The Genus Category and Cranial Morphometrics of the Catarrhini with Implications for Fossil Hominins

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The genus category and cranial morphometrics of the Catarrhini with implications for fossil hominins

Jack Andrew Coate

A thesis submitted in fulfillment of the requirements for the degree of Doctor of Philosophy

Postgraduate Board University of New South Wales

September 2007

PLEASE TYPE THE UNIVERSITY OF NEW SOUTH WALES Thesis/Dissertation Sheet

Surname or Family name: Coate

First name: Jack Other name/s: Andrew

Abbreviation for degree as given in the University calendar: PhD

School: SOMS Faculty: Anatomy

Title: The genus category and cranial morphometrics of the Catarrhini with implications for fossil hominins

Abstract 350 words maximum: (PLEASE TYPE) Recently, the number of hominin genera has increased dramatically. Prior to the announcement of Ardipithecus, only two genera were used by paleoanthropologists: Australopithecus and Homo. Presently, up to eight hominin genera are used: Sahelanthropus, Orrorin, Ardipithecus, Australopithecus, Praeanthropus, Kenyanthropus, Paranthropus and Homo. Unlike species concepts, the genus category has not received wide critical examination. To investigate the use of the genus category in paleoanthropology, a comparative framework drawing on morphometric data from a large number of catarrhines is developed. Cranial variables include 36 standard linear measurements from representatives of catarrhine genera across the major tribes/families. This study seeks to assess whether too few or too many hominin genera have been recognized compared with extant catarrhines. Moreover, two published hypotheses about the use of Homo are examined: 1) Wood & Collard’s (1999) proposal to transfer Homo habilis/rudolfensis to Australopithecus; and 2) Goodman et al’s (1998) classification of both humans and chimpanzees in Homo. To analyze these cranial variables and a number of shape indices calculated from them, as well as to assess competing hypotheses, univariate, bivariate and multivariate statistical approaches are used. The results allow the identification of a set of variables and shape indices which distinguish genera across the catarrhines. Importantly, body size seems to be the major separator of catarrhine genera, reinforcing the idea that they occupy discrete adaptive zones. Moreover, differences between these genera mostly represent contrasts in the size of the neuro- versus the viscerocranium. When applied to hominins, a picture emerges which distinguishes them from extant catarrhines: cranial shape rather than size is the major component distinguishing them; this suggests that extinct hominins occupied similar habitats and adaptive zones; variability in size and shape within hominin genera is much lower than extant catarrhines; and the major differences seen in shape among hominins are the result of encephalization in Homo. It is concluded here that both Wood & Collard’s (1999) and Goodman et al.’s (1998) proposals appear to be premature. Moreover, while the earliest hominins may be too finely split at the genus level, the evidence for distinction of Australopithecus and Paranthropus is solid.

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Recently, the number of hominin genera has increased dramatically. Prior to the announcement of Ardipithecus, only two genera were used by paleoanthropologists: Australopithecus and Homo. Presently, up to eight hominin genera are used: Sahelanthropus, Orrorin, Ardipithecus, Australopithecus, Praeanthropus, Kenyanthropus, Paranthropus and Homo. Unlike species concepts, the genus category has not received wide critical examination. To investigate the use of the genus category in paleoanthropology, a comparative framework drawing on morphometric data from a large number of catarrhines is developed. Cranial variables include 36 standard linear measurements from representatives of catarrhine genera across the major tribes/families. This study seeks to assess whether too few or too many hominin genera have been recognized compared with extant catarrhines. Moreover, two published hypotheses about the use of Homo are examined: 1) Wood & Collard’s (1999) proposal to transfer Homo habilis/rudolfensis to Australopithecus; and 2) Goodman et al.’s (1998) classification of both humans and chimpanzees in Homo. To analyze these cranial variables and a number of shape indices calculated from them, as well as to assess competing hypotheses, univariate, bivariate and multivariate statistical approaches are used. The results allow the identification of a set of variables and shape indices which distinguish genera across the catarrhines. Importantly, body size seems to be the major separator of catarrhine genera, reinforcing the idea that they occupy discrete adaptive zones. Moreover, differences between these genera mostly represent contrasts in the size of the neuro- versus the viscerocranium. When applied to hominins, a picture emerges which distinguishes them from extant catarrhines: cranial shape rather than size is the major component distinguishing them; this suggests that extinct hominins occupied similar habitats and adaptive zones; variability in size and shape within hominin genera is much lower than extant catarrhines; and the major differences seen in shape among hominins are the result of encephalization in Homo. It is concluded here that both Wood & Collard’s (1999) and Goodman et al.’s (1998) proposals appear to be premature. Moreover, while the earliest hominins may be too finely split at the genus level, the evidence for distinction of Australopithecus and Paranthropus is solid.

II Acknowledgements: I would personally like to thank my supervisor and co-supervisor, Dr. D. Curnoe and Prof. K. Ashwell, as well as, many other close friends and colleagues, including; H. Cohen, E. Danielson and family, T. Furlong, H. Green, D. Harris, Dr. A. Herries, S. Kiker, J. Louys, K. McQualter, D. Neuweger, Dr. S. van Holst, and L. van der Weyde. In addition, I would like to especially thank my parents and family for emotional and financial support throughout the years and during this thesis project. Furthermore, I would also like to personally thank the following institutions and their Physical Anthropology, Zoology and/or Mammalogy staff for allowing me to gather craniometric data from primate skulls in their care – Dr. S. Ingleby and T. Ennis at the Australian Museum, Sydney, Australia; Prof. F. Thackeray, Dr. T. Kearney, S. Potze and T. Perregil at the Transvaal Museum, Pretoria, South Africa; Prof. M. Raath, M. Dayal, M. Spocter, B. Billings and J. Hemingway at the University of Witwatersrand, School of Anatomical Sciences, Jo-burg, South Africa; J. Spence and E. Sarmiento at the American Museum of Natural History, NYC, NY, USA; Dr. Y. Haile-Selassie, L. Jellema and Prof. B. Latimer at the Cleveland Museum of Natural History, Cleveland, OH, USA, which houses the Hamann-Todd Collection; M. Schulenburg at the Field Museum, Chicago, IL, USA; L. Gordon at the Smithsonian Institute, National Museum of Natural History, Washington, D.C., USA; and P. Jenkins and D. Hills at the Natural History Museum, London, UK. In addition, Dr. R. Raaum and Dr. A. Tosi provided genetic information and distance matrices for some of the primates studied herein and both are greatly appreciated by the author for their data and assistance.

III Table of contents:

Abstract: II Acknowledgements: III Table of contents: IV Abbreviations: V-VII

Chapter 1: Introduction, pp.1-18. Chapter 2: Materials and Methods, pp.15-50. Chapter 3: Results for genera of Cercopithecini, pp.51-113. Chapter 4: Results for genera of Papionini, pp.114-179. Chapter 5: Results for genera of Colobinae, pp.180-277 Chapter 6: Results for genera of Hominoidea, pp.278-337 Chapter 7: Summary and Discussion of results for the extant Catarrhini genera, pp.338- 356. Chapter 8: Results for genera of the Hominina, pp.357-412. Chapter 9: Conclusions, pp.413-420. References: pp.421-489. Appendix: pp.490-500.

Anatomical, Genus and Journal Abbreviations (in alphabetical order): AJP – American Journal of Primatology AJPA – American Journal of Physical Anthropology Al. – Allenopithecus An. – Ankarapithecus Ar. – Ardipithecus Au. – Australopithecus Bas – Basion Bien – Interentoglenoid Breadth bp – basepairs Br - Bregma Cc. – Cercocebus Cp. – Cercopithecus Ch. – Chlorocebus Co. – Colobus Cum. – Cumulative CV – Coefficient of Variation or Canonical Variate(s) CVA – Canonical Variate Analysis D. – Dryopithecus Df: - Degrees of Freedom Dm3 – Distal M3 E. – Erythrocebus Ecm – Ectomolare Enm – Endomolare EQ – Encephalization Quotient Ex. – Excluding FAD – First Appearance Date Fmo – Frontomalarorbitale Fmt – Frontomalartemporale G – Glabella Go. – Gorilla H. – Homo Ho. – Homo H.he – “early Homo”, H. habilis sensu lato and H. erectus sensu lato Homo. - Homogeneity Hy. – Hylobates IJP – International Journal of Primatology Iv – Incisivion JAS – Journal of Anthropological Science JHE – Journal of Human Evolution Ka – Thousand years ago Ka. – Kenyanthropus Kp. – Kenyapithecus L. – Lophocebus V LAD – Last Appearance Date Lam – Lambda Ln – Natural Log Mn. – Mandrillus Mc. – Macaca Ma. – Mega Annum (or Megaannum; i.e. One million years) Max. – Maximum MANOVA – Multivariate Analysis of Variation Maxalvlen – Maxillo-alveolar Length (pros-dm3) Maxnawi – Maximum Nasal Aperture Width Mi. – Miopithecus Min. – Minimum MPE – Molecular Phylogenetics and Evolution MSV – Mosimann Shape Variable(s) Myr – Million years N. – Nasalis Nas – Nasion Non-metric MDS – Non-Metric Multidimensional Scaling No. – Number(s) Ns – Nasospinale O – Opisthiocranion Ol – Orale Opn - Opisthion Or. - Orrorin Ou. – Ouranopithecus p. - page Para. – Paranthropus PC – Principal Component(s) PCA – Principal Component Analysis Pi. – Piliocolobus PNAS – Proceedings of the National Academy of Sciences, USA Pms – Palatomaxillary suture Po. – Pongo Pp. – Papio pp. – pages Prae. – Praeanthropus Pre. – Presbytis Pro. – Procolobus Pros – Prosthion Py. – Pygathrix R. – Rhinopithecus Rhi – Rhinion SD – Standard Deviation Sa. – Samburupithecus SAJS – South Africa Journal of Science Se. – Semnopithecus Sh. - Sahelanthropus VI Si. – Simias – Small Sigma = Standard deviation SS – Social Structure Sta – Staphylion Sv. – Sivapithecus Th. – Theropithecus Tr. – Trachypithecus TREE – Trends in Evolution and Ecology Var. – Variance or Variation ¯x – Sample mean or average Zi – Zygomaxillare inferior Zs – Zygomaxillare superior

VII The genus category and cranial morphometrics of the Catarrhini with implications for fossil hominins

Chapter 1: Introduction - Historical and current trends in anthropology & taxonomy The genus and species categories are the foundation of Linnaean binomial nomenclature and modern taxonomy. The official commencement of Linnaean binomial nomenclature for many vertebrates and invertebrates begins with the tenth edition of Linneaus’ Systema Naturae, published in 1758. This hierarchical classification scheme is internationally accepted and has become embodied in the International Commission of Zoological Nomenclature (ICZN). In fact, the combination of these categorical names brought order and provided stability to taxonomy, which is something that had eluded Linneaus’ naturalist predecessors. While a considerable amount of time and energy has been devoted to species concepts, the genus category has received little attention and is generally thought to be ‘unproblematic’ (Wood & Collard, 1999a). In its most basic capacity, the genus category is simply either one monotypic species or a group of closely related species. Despite this simple definition, implementing the genus category in practice is much more difficult, especially when evaluating living species which may have fossil ancestors. The problem lies in the philosophical reasoning and varied evidence (morphology, genetics, ecology, behavior, etc.) that are employed to demarcate generic boundaries, which can, depending on the researcher, produce different results. The study and classification of human evolution has been and continues to be a contentious area of research. Despite nearly 250 years (2007 C.E. minus 1758 C.E. = 249 years) of taxonomic nomenclature and meticulous research, the placement of modern humans and their closest living relatives, the great apes, remains a debated topic. In fact, major interpretative (i.e. philosophical) differences and analyses of varied evidence (biomolecular and morphological) exist in paleoanthropology and primate systematics as to how and why modern humans, fossil species and extant primates, are classified. The biological classification of humans and closely related fossil species has nearly always been a highly debated topic, engendering multidisciplinary research and taxonomic refinement or revision (Marks, 2005). However, this should not come as a surprise considering that humans are the most studied of all primates with no lack of scientific theory. Modern humans belong to the genus Homo Linneaus, 1758 and are designated specifically by H.

sapiens - a fact which is not questioned. Although created in a pre-evolutionary framework, the development and use of the genus Homo includes many complementary and diverse forms of evidence and inferences. However, currently within the paleoanthropological literature there are numerous proposed hominin genera in use. In fact, there are polarized extremes of their possible relationships. Although the ‘robust australopithecines’ or ‘paranthropines’ of Paranthropus Broom, 1938 are not considered to be on the direct line to modern humans, they are thought to be closely related because of bipedal adaptations and cranial synapomorphies (Tobias, 1988). For this reason, the taxonomy of genera employed in paleoanthropology is as “chaotic” as it was in the first half of the twentieth century (Marks, 2005) when purported hominin genera were proposed for fossil remains, most of which were thought to date to the Early and Middle Pleistocene. In fact, Groves (2001a) lists nineteen generic synonyms for the genus Homo (see also Smith, 2002). However, the reasons for their creation are different to those proposed in recent years. While the genera of the first half of the twentieth century were created due to a lack of evolutionary theory and misunderstanding of the purpose of taxonomy, the recently proposed hominin genera are more a product of the predominance of cladistic theory and phylogenetic systematics. Lastly, the definition of the family Hominidae and genus Homo, and the genera, and species (extinct and extant) attributed to both are under critical debate (Wood & Collard, 1999a; Groves, 1999 & 2001b; Begun, 2001 & 2004a; Cela-Conde, 2001; Wildman et al, 2003). This thesis hopes to address and contribute to these issues. Over the last decade and a half four purported hominin genera have been described. Prior to these only two genera were commonly used or necessary, Australopithecus Dart, 1925 and Homo Linnaeus, 1758. Once not generally used in the literature (Tobias, 1967 & 1968; cf. Robinson, 1972), the genus Paranthropus is now readily employed by paleoanthropologists (e.g. Wood & Strait, 2004; cf. Suwa et al, 1997; Alemseged et al, 2002). Ardipithecus ramidus was initially announced as a species of Australopithecus (White et al, 1994), but White et al (1995) later decided a new genus was necessary to accommodate the fossil material. From what has been published, Ardipithecus is very primitive and bipedality for this species remains to be substantiated (Haile-Selassie, 2001; Haile-Selassie et al, 2004). The announcement, analysis and subsequent debate of Orrorin continues to draw controversy (Senut, 2001; Aiello & Collard, 2001; Sawada et al, 2002;

Pickford et al, 2002; Galik et al, 2004 & 2005;Ohman et al, 2005; White, 2006). Aside from its initial announcement, Kenyanthropus (Leakey, 2001; Leiberman, 2001) has received little attention (two exceptions, Cameron, 2003; Cela-Conde & Ayala, 2003); no doubt due to its poor preservation (White, 2003). In addition, large and statistically robust cladistic analyses of craniodental variables by Strait et al (1997) and Strait & Grine (2005) have argued for the removal of Australopithecus afarensis from Australopithecus and instead be placed in its own genus, Praeanthropus in an effort to remove paraphyly (an argument which has gained support and use by others - Wood & Collard, 1999a; Cameron, 2003; Cela-Conde & Ayala, 2003; Grine et al, 2006). Interestingly, the afarensis hypodigm has for years been subject to debate, particularly in regard to its locomotor repertoire, species composition and phyletic position (Tobias, 1980; Olson, 1981 & 1985; White et al, 1981; Susman et al, 1984 & 1985; Kimbel et al, 1984, 1985 & 1988; White, 1985; Ward, 2002). Despite this, strong evidence presented by Kimbel et al (2006) and White et al (2006) may demonstrate a ‘chrono-species’ relationship between Au. anamensis and Au. afarensis based on fossil mandibular and dental specimens from sites in Ethiopia, Kenya and Tanzania, between ~ 4.2 to 3 Mya. This casts doubts on the cladistic analyses by Strait et al (1997). Lastly, Sahelanthropus, dated to nearly ~ 7 million years, is perhaps the most thoroughly documented and analyzed taxon (Brunet et al, 2005; Zollikofer et al, 2005; Guy et al, 2005) – and may be a close relative, but not the last common ancestor of humans and chimpanzees (Wolpoff et al, 2002 & 2006). Understanding an evolutionary lineage from only fossilized skeletal remains is extremely difficult, both practically and theoretically (Tattersall, 1986 & 1992; Wood, 1988, 1991 & 1993; Cope, 1993; Collard & Wood, 2000; Plavcan & Cope, 2002). Still confounding matters is the great potential for homoplasy and differing amounts of variation-covatiation in closely, and even distantly related species (Lieberman et al, 1996; Larson, 1998; Lockwood & Fleagle, 1999; Ackermann, 2002 & 2003). As to taxonomy below the genus level, some workers argue for many species of hominins (Tattersall, 1986 & 2000; Cameron, 2003). For example, in Wood & Richmond’s (2000) review of hominin taxonomy and paleobiology, based on fossil, geological age and paleoecological evidence, four genera compromising more than 15 species are discussed (not including Sahelanthropus, Orrorin or Kenyanthropus). Justification for these species

mainly rests on morphological differences, although geological age, location of discovery and associated fauna play a role as well. What is more, the possible inferences from these data may elucidate behavioral and ecological differences. Different analyses based on ecological variables such as home range size and body mass (Foley, 1991; and Conroy, 2002 & 2003) indicates that the number of species used within paleoanthropology is reasonable within a mammalian or primate context. Yet, Eckhardt (2000) and Curnoe & Thorne (2003) have suggested far fewer species, <5, based on molecular data. On the other hand, most researchers would classify modern humans, chimpanzees and gorillas into separate genera. In general, the modern usage of the genus Homo is relatively agreed upon and represents an evolutionary unit or threshold bound by anatomy, locomotion, archaeology, ecology, life-history variables and inferred behaviors (Hughes & Tobias, 1977; Duchin, 1990; Stanley, 1992; Wood, 1992; Grine et al, 1996; Kimbel et al, 1996; McHenry & Berger, 1998; Aiello & Wells, 2000; McHenry & Coffing, 2000; Moggi- Cecchi, 2001; Dunsworth & Walker, 2002; Stiner, 2002; Bramble & Lieberman, 2004; Bobe & Behrensmeyer, 2004). However, recently, some anthropologists and molecular researchers have suggested placing humans and chimpanzees in the same genus and perhaps the gorilla as well, because of their small genetic distance from each other and their estimated recent divergence in the Late Miocene/Early Pliocene (Avise & Johns, 1999; Castresana, 2001; Curnoe & Thorne, 2003; Watson et al, 2001 and Wildman et al, 2003). However, even extant species are difficult to demarcate, evident from numerous observations of natural hybridizations between species in the wild and zoos (Van Gelder, 1977; Bernstein & Gordon, 1980; Struhsaker et al, 1988; Vervaecke & Van Elsacker, 1992; Jolly et al, 1997; Detwiler, 2002; Grubb et al, 2003; Brandon-Jones, 2004). Yet, some could argue the species category is the most objective category in any classification design (Simpson, 1961; Mayr, 1969; Groves, 2004). Still others have discussed the arbitrary nature of Linnaean taxonomy above the species level (see Anderson, 1975; Groves, 1986; Cela-Conde, 1998; de Quieroz & Gautheir, 1992; McKenna et al, 1997; Ereshefsky, 2001). Furthermore, there is no consensus on the definition, identification or delimitation for any of the lower categories within Linnaean hierarchal taxonomy for extant populations or species known only from fossils. Additionally, researchers use different paradigms, definitions and identification

procedures which they feel is the most appropriate or correct (e.g. cladistics or evolutionary systematics). There are also strong arguments for abandoning Linnaean hierarchal taxonomy and instead using nonranked monophyletic clades – e.g. the Phylocode (see de Quieroz & Gautheir, 1992; Benton, 2000; Groves, 2004). Most importantly though, there are inherent limitations as to how theoretical paradigms and hierarchical systems can be applied to the natural world and its phenomena (Mayr, 1974 & 1981; Knox, 1998; Szalay, 1999; Vrba, 1999; Mayr & Bock, 2002). Taking these comments into consideration, it is peculiar that over the past decade and half there have been numerous genera and species created or resurrected within paleoanthropology, each one (excluding Paranthropus and perhaps Kenyanthropus) vying for the position of the ‘direct ancestor’ of modern humans. As these examples and others make clear, recent analyses and discoveries have made hominin taxonomy volatile and unstable. This is in part by order of discovery, analysis and scientific announcement but another problem is the inadequate guidelines in demarcating higher taxa. Hominin genera currently used in paleoanthropology include, in nearly geological order from the oldest to the most recent:

Sahelanthropus Brunet et al, 2002, ~ 6 - 7 Ma; Orrorin Senut et al, 2001, ~ 6 Ma; Ardipithecus White et al, 1995, ~ 5.8 - 4.4 Ma; Australopithecus Dart, 1925, ~ 4.2 - 2 Ma; Praeanthropus Senuyek, 1955; ~ 3.7 - 2.9 Ma (Resurrected by Strait et al, 1997; see also Day et al, 1980; and Strait & Grine, 2004); Kenyanthropus Leakey et al, 2001, ~ 3.5 Ma; Paranthropus Broom, 1938, ~ 2.5 - ~ 1 Ma; & Homo Linneaus, 1758, ~ 2.5 - Present.

Please note, other fossil hominids which most likely contributed to the evolution of African hominid genera, such as Morotopithecus, Kenyapithecus, Samburupithecus, Dryopithecus and Ouranopithecus, are not examined within this thesis (see Leakey, 1967; de Bonis et al, 1990; McCrossin & Benefit, 1993; Ishida & Pickford, 1997; de Bonis &

Kuofos, 1997; Begun, 1992a & 1994; Begun et al, 1997; MacLatchy et al, 2000; Kordos & Begun, 2002; Young & MacLatchy, 2004; MacLatchy, 2004; Finarelli & Clyde, 2004). This large number of closely related genera within the span of less than seven million years is unusual when compared to the evolution and classification of other primates. In addition, other generic and species names are available for fossil hominin material (Groves, 1999). There are several extant and extinct primate genera which span nearly four to five million years or greater - Colobus Illiger, 1811; Macaca Lacepede, 1799; Victoriapithecus von Koenigswald, 1969; Sivapithecus Pilgrim, 1910; Libypithecus Stromer, 1913; Theropithecus Andrews, 1916; Gigantopithecus von Koenigswald, 1935; Aotus Illiger, 1811 and Tarsius Storr, 1780 (see Delson, 1975 & 1980; Andrews, 1981; Szalay & Delson, 1979; Tattersall et al, 1988; Kohler et al, 2000; Jablonski, 2002; Simons, 2003; Raaum et al, 2005). Groves (2001a, p.19; 2001b, p.296) also provides many nonprimate examples of genera and families with genera ~ 6 to 4 million years in time length; e.g. Ursus, Canis, Elephantidae, Bovidae. Nonetheless, the likely length of time for any genus should be kept in mind (Stanley, 1978; Simons, 2003). Moreover, there are several other hominoid genera from the Middle and Late Miocene of Africa and Eurasia and Africa (Hill & Ward, 1988; Andrews, 1992; Andrews et al, 1997; Andrews & Humphrey, 1999) which most likely contributed to, and are closely related to, the origin of the African ape clade - extant Subfamily Homininae (= Gorilla, Pan and Homo) (Groves, 1986; Cote, 2004). Some of these may include Kenyapithecus (McCrossin & Benefit, 1993 & 1997); Samburupithecus (Ishida & Pickford, 1997; Pickford & Ishida, 1998); Dryopithecus (Begun, 1992a, 1994 & 1995; Moya-Sola & Kohler, 1993 & 1996; Begun & Kordos, 1997; Kordos & Begun 2002); and Ouranopithecus (De Bonis et al, 1990; De Bonis & Kuofos, 1993 & 1997). Wood & Collard (1999a) are correct in stating that the genus category has received little attention in comparison to the consideration given to various species concepts, which is perhaps reason for its misuse and misunderstanding. Some notable exceptions to this would be discussions and work by Inger (1958), Tappen, (1960), Wolpoff (1978), Wolpoff & Lovejoy (1974), Walker (1976), Stanley (1978), Tobias (1978 & 1991), Wood (1992) and Groves (1989 & 2001a, b). The genus is the first rank above species in Linnaean binomial nomenclature - the first grouping, or hierarchical rank, of closely related species.

Figuratively, the genus is the fundamental ‘keystone’ of many more overarching, inclusive categories; the maximum and minimum number of which has estimated to be ~ 11 to 16 categories for a group of 100 species (Anderson, 1975). Klein (1999, p.11) lists a potential twenty-one levels in Linnaean hierarchal taxonomy and McKenna et al (1997) use twenty- five in their classification of Mammalia above the species level (Class, subclass, infraclass, superlegion, legion, sublegion, infralegion, supercohort, cohort, magnorder, superorder, grandorder, mirorder, order, suborder, infraorder, parvorder, superfamily, family, subfamily, tribe, subtribe, infratribe, genus & subgenus). Whereas the possible number of evolutionary trees is nearly beyond human comprehension (Felsenstein, 1978). As quoted by Mayr (1969, p.93), Linnaeus’ dictum (1737), “It is the genus that gives the characters, and not the characters that make the genus”, does intuitively have some truth. A formal definition was offered by Mayr, “A genus is a taxonomic category containing a single species, or a monophyletic group of species, which is separated from other taxa of the same rank [other genera] by a decided gap” (p.92). Mayr continues, “It is recommended for practical reasons that the size of the gap be in inverse ratio to the size of the taxon. In other words, the more species in the group the smaller gap needed to recognize it as a separate taxon, and the smaller the species group the larger the gap needed to recognize it. One of the function of the genus, from Linnaeus’ time on, is to relieve the memory (to facilitate information retrieval), and the “inverse ratio” recommendation prevents recognition of a burdensome number of monotypic genera” (p. 92). To delimit genera as species groups of optimal size is an operation that requires experience, good judgment, and common sense” (p.92). Later, Mayr (1982) would also comment that a genus is, “…the lowest collective category, an aggregate of species sharing certain joint properties” (p. 175). A definition not too different from more recent researchers in standard biological and anthropological texts, such as Tattersall et al (1988), Jones et al (1992) and Harrison et al (1995). In an analysis of the genus Papio (the baboons), Buettner-Janusch (1966) commented that the genus category could be discussed “ad nauseam” and yet admits it does recognize “natural phenomenon” in the biological world (p.290). Unfortunately, the genus category is difficult to define and consistently implement, which can lead to differences in interpretation, accompanied with an inflation of genera that may or may not have any biological usefulness. For example, Groves (2001a,

p.308; see also Smith, 2002, p.442, table 26.1; Campbell, 1965; McKenna et al, 1997) lists twenty generic names that were proposed for fossils (most of which are thought to date to the Middle or Late Pleistocene) which are now all included within Homo. These examples are evidence for a lack of a solid philosophical and empirical basis for the genus category (and therefore, hierarchal rankings as well), which has been discussed by others (e.g. Groves, 1989, 2001a & 2004; de Quieroz & Gauthier, 1992; Valentine & May, 1996; Knox, 1998; Cela-Conde, 1998; Cantino et al, 1999; Wood & Collard, 1999a & b, 2001a & b; Ereshefsky, 2001; Sarmiento et al, 2002; Cela-Conde & Ayala, 2003; Wildman et al, 2003). This research hopes to contribute to this debate and refine generic classification. Yet, consensus remains elusive while the number of fossil primate genera increases unabated (e.g. pliopithecoids - Moya-Sola et al, 2001; Rossie & MacLatchy, 2006; cercopithecoids - Frost, 2001; Hlusko, 2006; hominoids - Moya-Sola et al, 2004; Ward et al, 1999; Gebo et al, 1997) alongside robust genetic data revealing previously unrecognized groupings (e.g. Catarrhines - Page & Goodman, 2001; Papionins - Disotell et al, 1992; Disotell, 1994; Harris& Disotell, 1998; Harris, 2000; Cercopithecins - Tosi et al, 2002a & b, 2003, 2004; African Bush babies - Delpero et al, 2000). However, attempts must be made and tested. As stated earlier, the incentive for the present thesis began with a critical examination of two very different proposals defining the genus Homo (Wood & Collard, 1999a; Goodman et al, 1998), which inevitably led to broader questions; 1) what is a genus and 2) is it applied to extant and extinct primates equivalently? Equivalency, here operationally defined as containing comparable amounts of biological content and potential inferences from available evidence. More precisely though, when evaluated comparatively with extant catarrhine genera, both quantitatively and qualitatively, are the recently proposed hominin genera (e.g. Sahelanthropus, Orrorin, Ardipithecus, Kenyanthropus) in paleoanthropological taxonomy and phylogenetic reconstructions equivalent and/or comparable to extant nonhominin genera? Certainly the available evidence on extant primate genera far outweighs the inferable evidence from the fossil record but can the currently used genera in taxonomic nomenclature and systematics serve as comparative benchmarks to gauge the validity of the recently proposed hominin genera?

Wood & Collard’s genus argument: Wood & Collard’s (1999a) analysis of the genus category was directly concerned with the fossil species attributed to the genus Homo and their relation to modern humans, Homo sapiens, which is also the type species of the genus. They concluded that a genus could be defined as “a species, or monophylum, whose members occupy a single adaptive zone” (Wood & Collard, 2001a; p.67). This definition is influenced and compromised by ideas from evolutionary systematics and cladistics. However, an ‘adaptive zone’ or ‘plateau’ is not a new idea (see Mayr, 1950; Van Valen, 1971; Wolpoff & Lovejoy, 1975). Thus defined, their critical examination of the available evidence led them to conclude that H. habilis and H. rudolfensis should be removed if paraphyly is to be avoided while a coherent adaptive zone is upheld. Their conclusions are very similar to Wolpoff’s (1999; see Wolpoff et al, 1999). This argument has been accepted by some, such as Aiello & Andrews (2000), while others are at least “sympathetic” to the proposal (Conroy, 2005, p. xxiii). Furthermore, for any species attributed to the genus Homo (or for that matter, any other genus or species - extant, extinct, and fossil lineages with living descendants), “First, the species should belong to the same monophyletic group as the type species of that genus. Second, the adaptive strategy of the species should be closer to the adaptive strategy of the type species of the genus in which it is included, than it is the type species of any other genus” (Wood & Collard, 2001a, p. 67). Moreover, these requirements agree quite well with taxonomic rules of the ICZN which does require a genus to be defined by its type species and any member compared to it (Tattersall et al, 1988, p.222). Justification for the removal of H. habilis and H. rudolfensis relies on a number of inferred life-history and ecological variables; as well as an objective evaluation of the assumed qualities or behaviors of the species generally allocated to the genus Homo. One implicit or explicit criterion of the genus Homo was spoken language. For example, Tobias (1991 & 1994) has argued that with the appearance of H. habilis there are diagnosable features on the endocasts of this species which indicate a human-like brain configuration, perhaps indicating the possession of spoken language. In addition, the appearance of stone tools was assumed to be made by members of Homo as well (Leakey et al, 1964; Tobias, 1967; however, bear in mind, there have been numerous field observations of many monkey and ape species that fashion and use tools; Yamakoshi, 2004). Although, some

have questioned this inference because gross fossilized endocast features do not necessarily indicate neurological capabilities as chimpanzees have similarly developed frontal and temporal cortical areas as well (Gannon et al, 1998; Semendeferi et al, 2002). Interestingly, Elton et al (2001) demonstrate that other hominins (e.g. Paranthropus) also experienced brain encephalization. Moreover, the Plio-Pleistocene deposits that yield the earliest stone tools also contain contemporaneous and sympatric hominin species, all of which predate any fossils that have been allocated to Homo, although temporally it is very close (Semaw et al, 1997; de Heinzelin et al, 1999; Kimbel et al, 1996). Still, another example includes the analysis of postcranial elements of H. habilis and H. rudolfensis, which Wood & Collard argue are quiet different from H. sapiens, most likely reflecting different locomotor behaviors and habitat use. However, Haeusler & McHenry (2004) have more recently reanalyzed OH 62, a fragmentary skeleton presumably representing H. habilis, and their results indicate that the lower limb was more human-like then had been originally reconstructed and interpreted but that the upper arms in this species were still relatively longer than modern humans.

Goodman et al genus argument: The definition of the genus category offered by Goodman et al (1998 and 2001; see also Wildman et al, 2003) was derived by the direct comparison of the DNA of living primate species complemented by fossil evidence, plus strict adherence to phylogenetic principles (e.g. taxa are monophyletic clades and the age of the clade decides the rank of the taxa; both originally proposed by Hennig, 1966). The nuclear DNA under study was the -globin gene cluster, ~ 60- to 80-kilobase (kb), and within this cluster are -type genes ( and ). Furthermore, the nuclear regions examined include a pseudogene, noncoding introns of the functional genes and the flanking regions of these sections which all evolve at a faster rate compared to the coding sequences. Thus, these regions are selectively neutral, provide local molecular clocks, and the -globin locus (a 1.7-kb section of three exons and two introns) has been sequenced for 43 primate species. The fossil record is used to calibrate these molecular results. For example, the fossil primate genera Victoriapithecus (a cercopithecoid) and Proconsul (a hominoid) both contemporaneous during the early Miocene of East Africa provide direct evidence that by ~

22 Ma (at the latest) the lineages leading to the Old World Monkeys and Apes had already diverged. As a result, 25 Ma is the temporal datum point for the cercopithecoid-hominoid split. In another example, Catopithecus (a eucatarrhine) at 35 Ma (late Eocene, at the latest) again provides direct evidence that the platyrrhine and catarrhine lineages had separated by this time. Consequently, 40 Ma is the inferred timing of divergence for the platyrrhine and catarrhine lineages. In addition the results presented by Goodman et al (1998 & 2001) are congruent with many other primate morphological and molecular studies (e.g. Primates - Shoshani et al, 1996 and Poux & Douzery, 2004; Strepsirhines -Yoder et al, 1996; Platyrrhines - Schneider, 1996 and Horovitz et al, 1998; Catarrhines - Raaum et al, 2004; Hominoids - Caccone & Powell, 1989; Ruvolo, 1997a, b). The conclusions reached by Goodman et al (1998 & 2001) suggest that the age for a genus treated as ‘crown’ groups should 6-4 Ma; and 11-7 Ma if the ‘total’ group is considered (“A crown group is an extant monophyletic taxon defined by its LCA [Last Common Ancestor]; therefore it includes the LCA and all lineages that descend from this LCA to both extinct and extant species, but it does not include the stem of the LCA, whereas the total group does include the stem” (ibid. p. 592).). Autonomously, similar generic age estimates were obtained by Curnoe et al (2006). Primate genera defined as such, treated either as a crown or total group, requires modern humans and the chimpanzees both to occupy the genus Homo because they are estimated to have diverged only 6 Ma and are sister species (Groves, 1986; Martin, 1986; Page & Goodman, 2001; Gibbs et al, 2002; Wildman et al, 2003; Uddin et al, 2004); thus resulting in, H. (Homo) sapiens, H. (Pan) troglodytes & H. (Pan) paniscus. The fruition of J. Diamond’s (1988 & 1992) prediction has been realized (DNA-based phylogenies of the three chimpanzees. Nature 332, 685- 686; and The rise and fall of the third chimpanzee. London: Vintage.). Consequently, all putative hominin fossil genera are then grouped under Homo and then separated subgenerically. Independently, other molecular researchers and anthropologists such as Avise & Johns (1999), Watson et al (2001), Castresana (2001) and Curnoe & Thorne (2003) have also proposed the congeneric status of humans and chimpanzees. However, genetic techniques and hypotheses generated to demarcate taxonomic boundaries are not without criticism (e.g. Ferguson, 2002).

This is counter to other anthropologists who recognize up to 8 different hominin genera (Cameron, 2003; Strait & Grine, 2004; Wolpoff et al, 2006) excluding Pan, which to some is not a hominid (e.g. Martin, 1990; Fleagle, 1998; Sarmiento, 1998; Foley, 1999b; Nowak, 1999; Haile-Selassie et al, 2004). Despite theoretical developments (e.g. Hennig, 1966), modern genetics (e.g. Kumar, 2005) and an ever expanding fossil record (e.g. Hartwig, 2002), have Simpson’s (1945) critical comments regarding primate taxonomy influenced many investigators (see also Martin, 1981; Eckhardt, 2000; Tattersall, 2000; Foley, 2001; Wildman et al, 2003; Marks, 2005)? For example, Simpson wrote, “The primates are inevitably the most interesting of mammals to an egocentric species that belongs to this order. No other mammals have been studied in such detail, yet from a taxonomic point of view this cannot be considered the best-known order, and there is perhaps less agreement as to its classification than for most other orders. A major reason for this confusion is that much of the work on primates has been done by students who had no experience in taxonomy and who were completely incompetent to enter this field, however competent they may have been in other respects, and yet once their work is in print it becomes necessary to take cognizance of it. For this reason, if no other, it is not surprising that most primates have alternative names and that hardly two students use the same nomenclature for them. The importance of distinctions within the group has also been exaggerated that almost every color phase, aberrant individual, or scrap of fossil bone or tooth has been given a separate name, almost every really distinct species has been called a genus, and a large proportion of the genera have been called families. The peculiar fascination of the primates and their publicity value have almost taken the order out of the hands of sober and conservative mammalogists and have kept, and do keep, its taxonomy in a turmoil. Moreover, even mammalogists who might be entirely conservative in dealing, say, with rats are likely to lose a sense of perspective when they come to the primates, and many studies of this order are covertly or overtly emotional” (pp. 180-181); [and] “Perhaps it would be better for the

zoological taxonomist to set apart the family Hominidae and to exclude its nomenclature and classification from his studies” (188). Several other researchers have made similar comments – Anderson & Jones (1967); Simons (1963); Buettner-Janusch (1966); Tattersall & Eldredge (1977). Instead it would seem paleoanthropology and its taxonomic nomenclature are repeating past trends but for different reasons. While purported hominin genera prior to the evolutionary synthesis and cladistics were described because of non-population interpretations, misunderstandings of variation within species and misguided taxonomy, modern hominin genera have become splintered clades all deserving recognition. The similarities between Wood & Collard’s (1999a) arguments to those of Goodman et al (1998 & 2001) lie in their adherence to cladistic theory and recognition of monophyletic taxa only. However, the recognition of these taxa results from different forms of evidence (molecular and morphological) and analysis, which consequently result in dissimilar classifications (or each offers different resolution of macroevolutionary events), each potentially valid and useful; and both deserving attention and empirical testing. There has been a long debate and intense research on the relationship between molecular and morphological data and its potential inferences for primates and mammals in general (King & Wilson, 1975; Cronin & Meikle, 1979; Cherry et al, 1978 & 1982; Andrews, 1986 & 1987; Wayne et al, 1991; Smith & Littlewood, 1994; Purvis, 1995; Mann & Weiss, 1996; Shoshani et al, 1996; Pilbeam, 1996; Horovitz et al, 1998; Omland, 1997; Lovejoy et al, 1999 & 2000; Hillis & Wiens, 2000; Eckhardt, 2000; Weiss & Fullerton, 2000; Tosi et al, 2003; Curnoe & Thorne, 2003; Roseman, 2004). Needless to say, cladistic theory and associated methods (also having emerged alongside molecular biology in the second half of the twentieth century) have come to dominate particular aspects of biological classification and evolutionary theory, particularly within paleoanthropology. In fact, large and robust phylogenetic analyses for extinct and extant hominins and nonhuman primates have been performed, producing varied results with most disagreements the outcome of rank assignment and taxon content (Eldredge & Tattersall, 1975; Delson et al, 1977; Schwartz et al, 1978; Kluge, 1983; Boaz, 1983; Ciochon, 1983; Skelton et al, 1986; Strasser & Delson, 1987; Harrison, 1987; Stringer, 1987; Andrews & Martin, 1987; Chamberlain & Wood, 1987; Begun 1992a & 2001 and Begun et al, 1997; Groves, 2000;

Gibbs et al, 2003; Strait & Grine, 2004; Finarelli & Clyde, 2004). However, there have also been many other researchers who urge caution and question the results of cladistics and the demand for strict monophyly in classifications and instead argue for other forms of evidence with different approaches to taxonomic schemes (Mayr, 1974 & 1981; Hull, 1979b; Martin, 1981; Harrison, 1993; Corruccini, 1994; Begun, 1995 & 2004a; Eckhardt, 2000; Collard & Wood, 2000 & 2001b; Curnoe, 2003; Ackerman, 2003; Hawks, 2004; Hlusko, 2004; Nadal-Roberts & Collard, 2004; Andrews & Harrison, 2005). This thesis explores the definition and use of the genus category in extant Catarrhine systematics and paleoanthropology. In particular, what is the most appropriate and objective manner(s) in determining the boundaries for the genus Homo based on the available evidence. Likewise, can extant Catarrhini genera and the available evidence they provide be drawn upon to assess the recently proposed hominin genera; in addition to other arguments to include perhaps the chimpanzees within Homo. Biological classification above the species level is a strongly debated topic. Arguments and evidence used to define the genus Homo can be based on behavioral, morphological (including soft tissue, skeletal features and fossils) and genetic data. Two hypotheses defining the genus Homo tested here are those of Wood & Collard (1999a) and Goodman et al (1998). The former is grounded upon various forms of evidence which can be gleaned from the available extant species and fossil evidence, which in its totality apparently defines an ‘adaptive zone’ of one or more species. The latter relies almost exclusively on biomolecular evidence (specifically, sequenced nucleotides and protein structure) but is calibrated by the fossil record and insists time-ranked monophyletic clades. The cranium was chosen as a skeletal element to record morphological variation within and between genera and because many primate crania have been recovered from the fossil record.

Taxonomy and Nomenclature: One source of confusion in taxonomy is simply human language. Language, spoken and written, can be at times repetitive, ambiguous and contradictory. However, it is nonetheless the medium with which we physically and mentally engage with other humans; chimpanzees are anatomically incapable of articulate human-like speech. Taking this into consideration, it is paramount to know exactly the content of our subjects but also how to

appropriately and accurately discuss them. Any taxonomic discussion or analysis must first set out precise and clear definitions for the words (i.e. nomenclature) used (Simpson, 1961). The analysis herein has adopted the taxonomic nomenclature proposed by Groves (2001a) although species and subspecies assignments are not the main focal point. Please note however, this research in no way questions the validity of the species proposed by Groves. They most certainly are the most recent and accurate descriptions available of extant primate populations which are diagnosable from others. It should be noted that Groves uses the phylogenetic species concept (see Cracraft, 1983; and Kimbel & Rak, 1993). Therefore, other researchers can and do disagree with his species groupings and their hierarchical rank. However, his detailed research in the matter has produced a comprehensive taxonomic nomenclature with which we may accurately and appropriately (in terms of nomenclature history and priority) discuss the Order Primates. Furthermore, Groves’ taxonomic placement of taxa is not too radically different from other recent schemes (e.g. Hill, 1966, 1974 & 1970; Delson & Andrews, 1975; Delson, 1977; Schwartz et al, 1978; Wolfheim, 1983; Schwartz, 1986; Groves, 1970 &1989; Napier, 1981 & 1985; Conroy, 1990; Rowe, 1996; Shoshani et al, 1996; McKenna et al, 1997; Fleagle, 1998; Ankel-Simons, 1999; Nowak, 1999). Furthermore, Groves’ taxonomy is supplemented by the data and observations of Grubb et al (2003) and Brandon-Jones et al (2004); both of which Groves is a co-author and represent some of the most recent syntheses for the Old World Primates. Of course, though, some genera are monotypic which inevitably leads to comparing a genus of one species to a genus with many and vice versa; although, some hominin genera are monotypic as well (e.g. Kenyanthropus, Orrorin, Sahelanthropus). Yet, analyses presented here are directly concerned with the genus category and how this is used in modern catarrhine systematics compared to that of fossil hominin genera (Wood & Collard, 1999a,b & 2001a,b); as well as, considering molecularly derived time-ranked taxonomic hypotheses (Goodman et al, 1998 & 2001; Wildman et al, 2003). Accordingly, the genus is the ‘operational taxonomic unit’ for the analyses presented here and therefore is concerned with macroevolutionary events and their classification. Validation for this level of analysis is supported by the results from Purvis & Webster (1999; see also Purvis, 1995; Purvis et al, 1995) and Hardt & Wenke (2006).

Purvis & Webster’s (1999) macroevolutionary analysis suggest the use of generic means allows for some form of independent contrasts. While Hardt & Wenke’s (2006) study included geometric morphometrics of hominoids and cercopithecoids and despite heterogeneous craniometric samples their results still contain reliable biological information and are appropriate because of their ability to successfully discriminate superfamilies and genera. However, please note, this thesis is not directly concerned with the cladistic relationships of the genera under study, nor can the cranial morphometrics collected definitively answer specific phylogenetic questions - they are not meant to. (By and large the major phylogenetic relations of the Catarrhini have been determined.) Rather, do the species of the genera presently in use actually share particular cranial features and dimensions due to their supposed adaptive zones? Lastly, in accordance with molecular findings since the 1960’s (Goodman, 1963a & 1963b), humans (Homo) and the great apes (Pan, Gorilla & Pongo) together belong within the family Hominidae and thus all are considered ‘hominids’, a term traditionally used to describe only humans and their bipedal fossil ancestors (e.g. Campbell, 1962 & 1965; Pilbeam, 1962). Consequently, the term ‘hominin(s)’ (Tribe Hominini (= Pan-clade and Homo-clade), Subtribe Hominina (= Homo + bipedal species more closely related to modern humans) is necessary in discussion to distinguish fossil human ancestors from the great apes. Within the Hominidae are two subfamilies, the Ponginae which includes the only extant Asian ape, Pongo spp., and fossil genera such as Sivapithecus, Lufengpithecus and Khoratpithecus (Chaimanee et al, 2003 & 2004). The other subfamily, the Homininae designates the African Apes (Gorilla, Pan and Homo). Groves (2001a) goes no further then subfamilial classification for the African apes and will be dealt with more fully in Chapters 6-9. Thus, ‘hominins’ (as ‘homininans’ is a bit cumbersome and perhaps unnecessary; cf. Andrews & Harrison, 2005) in this analysis refers to fossil species thought to be more closely related to humans than to chimpanzees (Chapter 6, Figure 6.1). Additionally, there is no fossil evidence for chimpanzees older than the Middle Pleistocene (McBrearty & Jablonski, 2005). As such, there should be no ambiguity or confusion as to what is being discussed or referred to. That is, until ancient (>3-5 Ma) fossils are recovered and their relationship to modern chimpanzees is demonstrated.

However, there are still some who prefer the traditional, gradistic classification in which the great apes are placed in the Pongidae, while humans and their immediate ancestors are the only members of the Hominidae (e.g. Martin, 1990; Fleagle, 1999; Nowak, 1999; Haile-Selassie et al, 2004; Grehan, 2006). Moreover, ‘cercopithecines’ of the subfamily Cercopithecinae refers to the cheek-pouch monkeys; ‘cercopithecins’ refers to members of the tribe Cercopithecini (Allenopithecus, Miopithecus, Cercopithecus, Chlorocebus & Erythrocebus) (Figure 3.1); and lastly, ‘papionins’ refers to members of the tribe Papionini (Cercocebus, Mandrillus, Macaca, Lophocebus, Papio & Theropithecus) (Figure 4.1). ‘Colobines’ refers to any member of the subfamily Colobinae (multi- chambered stomach monkeys) but the subtribe Colobina will be used to discuss the African colobines (Colobus, Piliocolobus and Procolobus); Presbytina designates the South and Southeast Asian langurs and leaf-monkeys (Semnopithecus, Trachypithecus and Presbytis); and lastly, Nasalina will be employed for the East and Southeast Asian odd-nosed colobines (Pygathrix, Rhinopithecus, Nasalis and Simias) (Figure 5.1). Finally, ‘cercopithecoids’ refers to any member of the superfamily Cercopithecoidea (Old World Monkeys), while ‘hominoids’ denote the lesser and great apes (including modern humans) of the superfamily Hominoidea (Figure 6.1).

Classification outline for the extant Catarrhini: Order: Primates Suborder: Strepsirrhini Suborder: Haplorrhini Infraorder: Tarsiiformes Infraorder: Simiiformes (Eusimiiformes) Parvorder: Platyrrhini Parvorder: Catarrhini (Eucatarrhini) (Neocatarrhini) Superfamily: Cercopithecoidea Family: Cercopithecidae Subfamily: Cercopithecinae Tribe: Cercopithecini Allenopithecus nigroviridus Miopithecus spp. (2 species) Cercopithecus spp. (25 species) Chlorocebus spp. (6 species) Erythrocebus patas Tribe: Papionini Cercocebus spp. (6 species) Mandrillus spp. (2 species) Macaca spp. (20 species) Lophocebus spp. (3 species) Papio spp. (5 species) Theropithecus gelada Subfamily: Colobinae Subtribe1: Colobina (African colobines) Colobus spp. (5 species) Piliocolobus spp. (9 species) Procolobus verus Subtribe: Presbytina (Langurs and leaf-monkeys) Semnopithecus spp. (7 species) Trachypithecus spp. (17 species) Presbytis spp. (11 species) Subtribe: Nasalina (Odd-nosed colobines) Pygathrix spp. (3 species) Rhinopithecus spp. (4 species) Nasalis larvatus Simias concolor Superfamily: Hominoidea Family: Hylobatidae Hylobates spp. (4 subgenera and 14 species) Family: Hominidae Subfamily: Ponginae Pongo spp. (2 species) Subfamily: Homininae Gorilla spp. (2 species) Pan spp. (2 species) Homo sapiens

1 Please note, Groves (2001a) does not formally designate nor define colobine subtribes but these are employed herein to facilitate taxonomic discussion and provide comparative boundaries. Bold numbers indicate genera with >10 species. 18 Chapter 2: Material and Methods

2.1 Aims and Objectives: The impetus for this research commenced with a critical evaluation of two proposals defining the genus Homo; 1) Wood and Collard (1999a) and 2) Goodman et al (1998). Initially, this endeavor began with a thorough investigation of the anthropological and primatological literature for examples of use, and evidence employed to define the genus category in extant Catarrhini systematics because humans and their closest relatives, living or extinct, are catarrhines as well. However, this soon revealed that even many extant genera, similar in some ways to fossil hominin genera, are also marked by taxonomic differences of opinion and interpretation. To evaluate the genera used within Catarrhini systematics and paleoanthropology many lines of evidence must be assessed and attempting equal consideration. But how can biomolecular data, morphology, ecology, substrate use, diet and the fossil record be synthesized and interpreted on common ground (e.g. Pilbeam, 1996; Pilbeam & Young, 2004)? Questions one to six will be answered and discussed at the end of chapters three to six while questions seven to nine will be dealt with in the final chapter. Questions explored in this thesis include: 1. Are the extant Catarrhini genera adequately defined and do they compose an ‘adaptively coherent’ grouping of one or more species which occupy an ‘adaptive zone’, as defined by Wood & Collard (1999a)? 2. In terms of cranial morphology, how distinct are genera? In other words, how readily may a genus be determined from cranial dimensions, considered either singular or multiple (both metrical and nonmetrical traits)? 3. How much morphological variation is encompassed within a Catarrhine genus? Or, in other words, how does a ‘genus’ or ‘species’ samples behave statistically? 4. How much has one genus morphologically and genetically diverged from another? 5. Do cranial morphometric similarities and/or differences reflect adaptive zones? 6. What analogies may be drawn from extant Catarrhini genera for the interpretation of fossil hominin genera?

7. Are the recently proposed fossil hominin genera adaptively coherent occupying an adaptive zone different from that of other hominin genera based on comparisons with extant Catarrhini genera? 8. Is the congeneric placement of humans and chimpanzees (Goodman et al, 1998 & 2001 and others) more parsimonious than each occupying separate genera? 9. Would the congeneric placement of humans and chimpanzees better explain and synthesize the available evidence or oversimplify evolutionary differences? Or put more simply, what types of evidence best delineate generic boundaries?

2.2 Introduction: The guiding principles which this thesis is built upon are taxonomic uniformitarianism and equivalence. Modern osteological and dental specimens provide a baseline from which the fossil record may be gauged and interpreted (Wolpoff, 1971 & 1978; Wood, 1991; Plavcan & Cope, 2002). In addition, by knowing the behavioral and ecological context of extant skeletal material the paleobiology of fossil genera and species may also be inferred (Chiochon, 1993; Bromage, 1999; Elton, 2001; Reed, 2002). Quantifiable and morphological features, either singular or multiple, further reveal critical similarities and important functional differences, both of which may be due to either close phylogenetic relationship or evolutionary convergence (or homoplasy). Of course phenetic studies have practical and theoretical limitations (de Quieroz & Good, 1997) but they do nonetheless organize, quantify and synthesize biological data, from which many sound inferences and arguments are possible (Olson & Miller, 1958; Sneath & Sokal, 1973; Corruccini, 1975). The evolution and classification of modern Homo sapiens is perhaps best achieved and interpreted within a comparative evolutionary framework which also provides for some degree of objectivity via numerous pairwise comparisons (Gittleman & Luh, 1992; MacLarnon, 1999). Modern humans belong to the genus Homo Linnaeus, 1758, a member of the Catarrhini (Old World monkeys (OWM) and apes), which is the focus of this study. The Catarrhini were chosen because they contain relatives of Homo, and represent a natural grouping bound by anatomy, time, paleoclimatic events and biogeography (Delson & Andrews, 1975; Fleagle & Kay, 1985; Delson, 1985; Andrews, 1985; Andrews et al, 1996;

Jablonski & Kelley, 1997; Harrison, 2004). More importantly, there is a long and productive history of comparative studies, providing behavioral, morphological, and genetic insights (Wolpoff, 1978; Dunbar, 1983; Foley, 1987; Fleagle & Reed, 1996 & 2004; Reed, 1997; Isbell et al, 1998; Jolly, 2001; Elton, 2006). The definitions of the genus category examined here are those of Wood & Collard (1999a) and Goodman et al (1998 & 2001). Both definitions are quite different yet do share some theoretical criteria. As discussed previously, Wood & Collard’s solution to this problem was to use elements from both evolutionary systematics and cladistics. Their definition of a genus is, “a species, or monophylum, whose members occupy a single adaptive zone” (2001a, p. 67), defined by such features as body size, relative brain size, diet, locomotion and masticatory features, some of which may be inferred from cranial morphologies and other forms of complementary evidence. However, the definition provided by Wood & Collard, if useful and compelling, should apply not only to modern humans and their fossil ancestors but also to extant primates and their fossil antecedents. This requirement also applies to the time- ranked genus categories proposed by Goodman et al (1998 & 2001), which if convincing should more accurately depict evolutionary relationships then previous classifications as well as synthesizing the available evidence. To test these definitions, direct comparisons between primate genera were necessary. Comparisons between genera will remain within their respective superfamily, subfamily, subtribe and tribe. In addition, interpreting results from different (though intimately related) forms of evidence (molecular and morphological) is crucial. The skeletal complex chosen for generic comparison in this investigation are the dimensions of the neuro- and viscerocranium (ex. the mandible) of provenienced adult catarrhines, which provide many indicators of the genera’s adaptive zone and ecomorphology (Bromage, 1999). In addition, interpreting results from different (though intimately related) types of evidence (biomolecular and morphological) is crucial. However, Hillis & Wiens (2000) are quite astute in stating, “Morphological and molecular studies can each address questions that cannot be addressed by the other” (p.1).

Table 2.1: Catarrhini genera and sample sizes included in this study (listed in alphabetical order).

Males Females 4 2 Allenopithecus nigroviridis (Lang, 1923) Allen’s Swamp Monkey; Africa 10 9 Cercocebus spp. (E. Geoffroy, 1812) Mangabeys; Tropical Africa 12 12 Cercopithecus spp. (Linnaeus, 1758) Guenons; Tropical Africa 13 11 Chlorocebus spp. (Gray, 1870) Vervets or Grivets; Africa 15 11 Colobus spp. (Illiger, 1811) Black & White colobines; Tropical Africa 9 7 Erythrocebus patas (Schreber, 1774) Red Patas Monkey; Africa 17 17 Gorilla spp. (I. Geoffroy, 1853) Gorillas; Tropical Africa 11 9 Homo sapiens (Linnaeus, 1758) Humans; Global 16 15 Hylobates spp. (Illiger, 1811) Gibbons & Siamangs; East & Southeast Asia 11 12 Lophocebus spp. (Palmer, 1903) Pseudo-mangabeys; Tropical Africa 15 19 Macaca spp. (Lacepede, 1799) Macaques; N. Africa, Gibraltar & East Asia 8 14 Mandrillus spp. (Ritgen, 1824) Mandrill & Drill; Tropical Africa 9 10 Miopithecus spp. (I. Geoffroy, 1842) Talapoin; Tropical Africa 18 10 Nasalis larvatus (E. Geoffroy, 1812) Proboscis Monkey; Borneo 19 14 Pan spp. (Oken, 1816) Chimpanzees & Bonobos; Tropical Africa 19 12 Papio spp. (Erxleben, 1777) Baboons; Africa & Arabian Peninsula 15 13 Piliocolobus spp. (Rochebrune, 1887) Red colobines; Tropical Africa 14 12 Pongo pygmaeus (Lacepede, 1799) Orangutans; Borneo & Sumatra 8 10 Presbytis spp. (Eschscholtz, 1821) Surilis; East & Southeast Asia 3 2 Procolobus verus (Rochebrune, 1887) Olive colobine; West Africa 14 11 Pygathrix spp. (E. Geoffroy, 1812) Douc Langurs; East Asia 5 12 Rhinopithecus spp. (Milne-Edwards, 1872) Snub-nosed colobines; E. Asia 5 4 Semnopithecus spp. (Desmarest, 1822) Hanuman langurs; S. & E. Asia 2 5 Simias concolor (Miller, 1903) Simakobu; Mentawai Islands 7 7 Theropithecus gelada (I. Geoffroy, 1843) Gelada; Ethiopia, Africa 15 18 Trachypithecus spp. (Reichenbach, 1862) Lutungs; South & East Asia 294 278 Total 572

2.3 The Primate Cranium: The skeletal complex chosen for study was the primate skull and its various dimensions (Figures 2.1 & 2.2). The skull provides many indications of species life history, evolution and behavior, providing adaptively and phylogenetically significant data and observations for all species of primates thus far studied (Albrecht & Miller, 1993). For example, Aiello & Wood (1994) and Spocter & Manger (2007) successfully employ cranial variables to estimate body mass of hominin fossil species. Furthermore, cranial morphology may also reveal ecomorphologies (Bromage, 1999). Some of these include nuchal breadth relative to body size and the presence or absence of a sagittal crest. Additionally, just as the size and shape of the eye orbits can reflect diurnal or nocturnal behaviors (Martin, 1990), are there any traits which can differentiate genera and their adaptive zones? The cranial measurements collected for this study serve many functions. First, many variables of an adaptive zone, as defined by Wood & Collard (1999a), may be inferred from cranial dimensions; some of these include brain size and masticatory apparatus dimensions. Second, the cranial dimensions recorded serve as a surrogate measure of generic ranges of morphological variation and composition. Third, craniometric study is a historically established form of anthropological discourse generating both continued scientific debate and research (Howells, 1973 & 1989; Liebermann, 1995; Lahr, 1996) and also allows the results from this analysis to be compared and evaluated with previous work (e.g. Verheyen, 1962; Vogel, 1966 & 1968; Howells, 1973 & 1989; Creel & Preuschoft, 1976; Marroig & Cheverud, 2001a & 2001b; Pan & Groves, 2004; O’Higgins, & Pan, 2004; Miller et al, 2004). Incidentally, many primate crania and cranial fragments have been recovered from the fossil record of Africa and Eurasia (see Wood, 1991; Tobias, 1991; Schwartz & Tattersall, 2002, 2003 & 2005). Lastly, recent cranial studies have suggested that when cranial morphometrics are treated with multivariate statistics the results are similar to and behave like neutral genetic evolution (Relethford, 1994 & 2002; Roseman, 2004; Brace, 2005; Harvati & Weaver, 2006), which better allows these results to be compared to genetic data. The data presented here are based on primate cranial morphometrics, whilst the genetic, postcranial and ecological data were obtained through the published literature. Fortunately primate skulls are heavily sampled by national museums and universities.

Determination of sex with many primate genera was relatively easy, especially those with extreme sexual dimorphism, such as, Papio or Pongo. Still, if there were questions regarding sexual assignment, the institute’s records and notes were employed. Extant primate crania measured and included within this study were gathered from the following institutions (listed in order of visit); The Australian Museum, Sydney, Australia; University of Witwatersrand, School of Anatomical Sciences, Johannesburg, South Africa; Transvaal Museum, Pretoria, South Africa; American Museum of Natural History, New York City, New York, USA; Hamann-Todd Collection at the Cleveland Museum of Natural History, Cleveland, Ohio, USA; Chicago Field Museum, Chicago, Illinois, USA; British Museum of Natural History, London, UK; and the Smithsonian Institute, Washington, D.C., USA. Craniometric data of fossil hominins was gathered from the pertinent literature (e.g. Weindenreich, 1943; Santa Luca, 1980; Pilbeam, 1982; Kimbel et al, 1984, 1994 & 2004; De Bonis et al, 1990; Kidder et al, 1992; Alpagut et al, 1996; Arsuaga et al, 1997; Lockwood & Tobias, 1999 & 2002; Gabunia et al, 2000; Vekua et al, 2002; De Lumley et al, 2006; Curnoe & Tobias, 2006; Rightmire et al, 2006). But of course, any phenetic examination depends on the subject and medium of investigation, including the biological populations (of any taxonomic rank) involved. In other words, “In interpreting these results it would seem wise to bear in mind that any conclusions which are drawn must relate to the parameters which were selected for analysis” (Day, 1967, p. 324). Day continues to reason, “A multivariate technique, by taking account of the correlations between individual parameters in each bone, can present the functional morphology of that bone as a unified concept and assign it a position in relation to others; clustering of points will produce a grouping of bones with similar functional affinities. It is likely that these groups will represent populations of a Linnaean species, but this is not inevitable because Linnaean taxonomy is usually based on a wider selection of parameters than those which can be realized by examining a single bone” (p. 324).

2.4 Cranial Measurements:

The linear measurements chosen for this investigation fall into two categories; 1) the absolute size of a particular structure or complex, and 2) the measured distance between suture junctions or their length. In turn, these two categories document the two

developmental and functional regions of the skull, the neurocranium and viscerocranium. Length (or height) measurements are generally in line the median plane whereas breadth (or width) dimensions are usually transverse to the median plane. However, taking into consideration of the extent to which modern humans have experienced unprecedented encephalization (e.g. Ruff et al, 1997), a limited number of measurements of the neurocranium were selected. This was done so as not to unduly overweight measurements of the neurocranium. Instead, several facial and suture length dimensions were collected because these are more indicative of phylogenetic relationships (Ackermann, 2002; Young, 2005) and social (e.g. sexual dimorphism between males and females) and/or dietary or positional adaptations (e.g. a sagittal crest or strongly developed nuchal crest). Furthermore, cranial morphometric studies in general reveal phylogenetic signals and/or disposition while postcranial morphometric examinations retrieve functional similarities and/or convergence (Oxnard, 1975 & 1998). Additionally, most researchers agree that genera should be monophyletic (see Wood & Collard, 2001a & 2001b; Groves, 2001a & 2004; Tosi et al, 2004 for examples and discussion). Thus, it is perhaps more appropriate to record and examine crania rather than the postcranial skeleton for testing generic boundaries. Wood’s (1991) monograph on the Plio-Pleistocene hominin cranial remains from Koobi Fora, Kenya, served as a guide and reference to the craniometric data collected for the analysis herein, including some abbreviations, although other measurements and terminology have been employed. These include Buikstra & Ubelaker (1994), Bass (1995), White (2000), Ackermann (2002) and Ackermann & Cheverud (2004). Moreover, many other researchers influenced theoretical, practical and descriptive aspects of the present examination, such as, Albrecht & Miller (1993), Relethford (1994 & 2002), Aiello et al (2000), Pan & Oxnard (2001), Frost et al (2003), Guy et al (2003), and Lieberman et al (2004). All measurements are chord lengths recorded in millimeters, many divided by others to produce proportional measures. Additionally, measuring suture lengths between cranial elements provides important phylogenetic information; also allowing homologous features across diverse primate genera to be measured and compared (Ackermann 2002; Ackermann & Cheverud, 2004). For paired measurements the average of right and left was recorded, although, if a specimen was damaged, measurements were taken on the intact and

preserved portions. Occasionally specimens are damaged and cannot be recorded. Thus, substituting the sample’s mean is perhaps necessary for some statistical computation (Rightmire, 1969; Howells, 1973). Additionally, statistical programs (PAST ver. 1.53- Hammer et al, 2006; and SPSS 11.0.1, 2001) optionally insert means for various tests or provide casewise deletion for missing values. Substituting the sample mean for certain measurements of particular individuals was necessary for only a limited number of individuals. Furthermore, these individuals were both included and excluded from statistic analyses to gauge their impact. From a preliminary investigation it was determined that this procedure did not greatly alter the results and thus were included. For assorted portions of this analysis different spreadsheet and statistical computer programs were utilized. These include, Microsoft Office XP Excel 2003 with the add-in StatistiXL ver. 1.5; SPSS 11.0.1 (2001); and PAST ver. 1.53 (Hammer et al, 2006). In addition, StatSoft, Inc. (2007; see http://www.statsoft.com/textbook/stathome.html) was routinely consulted for statistical advice, underlying assumptions and explanations of statistical methods. Lastly, the measurement of error (see White, 2000, p.307) determined for eighteen skulls (comprising eleven catarrhine genera) measured twice for thirty-six measurements was < 3.3% (range 1.7-6.4) with a standard deviation of 0.02. In addition, the average difference in distance between standard calipers and 3-D digitizing microscribe (using Rhinoceros 3.0 to generate linear distances between craniometric points) was 1.4 mm (range 0.61 - 2.8) with a standard deviation of 0.60. There should be no discrepancy between measurements taken with calipers or microscribe. Franklin et al (2005) produced very similar results using a 3-D digitizer compared to those based on data collected with standard calipers.

1. Cranial vault length - The chord distance between glabella and opisthiocranion (g-o) was determined with spreading (or hinge) calipers with the basicranium placed on a cranial rest and the right or left lateral side facing the observer. The chord length from glabella to opisthiocranion was included because this dimension represents the greatest length of the neurocranium in the median plane (Wood, 1991 - no. 1, p. 287).

2. Maximum cranial length - The chord distance between prosthion and opisthiocranion (pros-o) was determined with spreading calipers with the basicranium placed on a cranial rest and the right or left lateral side facing the observer. Please note, in some primate genera opisthiocranion is inion, or immediately superior to it – e.g. Colobus, Macaca, Papio, etc. The chord length from prosthion to opisthiocranion was chosen because it represents the maximum length of the entire cranium in the median plane (Wood, 1991 - no. 2, p. 287). 3. Biasterionic breadth (biast) - The chord distance between left and right asterion was measured by dial (or sliding) calipers using the lower arms, with the neurocranium placed on a cranial rest and the occiput facing the observer. If the area of asterion is complicated with swirling ossicles, the greatest width and extent of the occipital bone (although near the junction of the temporal and parietal bones) was considered to be asterion. Biasterionic breadth was included because it represents the greatest width of the occipital bone and is the juncture of three important neurocranium bones (the parietal, occipital and temporal bones). Furthermore, the occipital bone provides surface area for many functionally important nuchal muscle attachments (Wood, 1991 - no. 14, p. 288). 4. Biporionic breadth (bipor) - The chord distance between left and right porion was measured with dial calipers with the lower arms positioned superolaterally to the most lateral extent of the bony tube of the external auditory meatus with the tips of the calipers resting on the inferior root of the zygomatic process with the neurocranium placed on a cranial rest and the occiput facing the observer. Biporionic breadth was chosen because it is an excellent indicator of basicranium breadth perpendicular to the median plane and is significantly correlated with body weight (Aiello & Wood, 1994; Wood, 1991 - no. 11, p. 287). 5. Biauriculare breadth (biaur) - The chord distance between left and right auriculare was measured using spreading calipers using the rounded end points of the calipers resting in the deepest incurvature at the root of the zygomatic processes, with the basicranium placed on a cranial rest and the occiput facing the observer. Biauriculare breadth for this study is the operational measurement for the greatest width of the cranium perpendicular to the median plane because it was a

homologous region across the entire sample. If bieuryonic breadth was recorded, the greatest width for humans would be located on the parietals, but for many other catarrhines, the widest point of the skull (ex. bizygomatic breadth) would be the lateral development of the temporal bones associated with nuchal muscle attachments. Thus, biauriculare breadth served as the measurement depicting the greatest cranial width across all genera by measuring from homologous features (Buikstra & Ubelaker, 1994 - no. 9, p. 75). 6. Bizygomatic breadth (bizygo) was determined using spreading calipers with the basicranium placed on a cranial rest and the viscerocranium facing the observer. Bizygomatic breadth was chosen because it represents one of the widest points of the skull perpendicular to the median plane and is a homologous and reproducible dimension across taxa (Wood, 1991 - no. 52, p. 289). 7. Bifrontomalarorbitale breadth (bifmo) was measured using dial calipers by placing the upper arms at the junction of the fronto-zygomatic suture on the lateral borders of the right and left eye orbits, with the basicranium placed on a cranial rest and the viscerocranium facing the observer. Bifrontomalarorbitale breadth was included because this dimension is correlated with body weight in hominins (Aiello & Wood, 1994) and describes the upper facial breadth excluding the lateral development of the zygomatic and frontal bones. Furthermore, bifmo is very similar or nearly approximate to biorbital breadth (bi-ektoconchion, ek-ek; Wood, 1991 - no. 50, p. 289) and is reported for many fossil specimens. 8. Superior facial breadth (bifrontomalartemporale breadth; bifmt) was determined using dial calipers by placing the lower arms at the junction of the frontal and zygomatic bones on the most superolateral extent of the fronto-zygomatic suture, with the basicranium placed on a cranial rest and the viscerocranium facing the observer. Bifrontomalartemporale breadth was chosen because this dimension represents the greatest breadth of the upper face, including the lateral development of the zygomatic and frontal bones (Wood, 1991 - no. 49, p. 289). 9. Bizygomaxillare superior breadth (bizs) was measured using the lower arms of dial calipers and placing them upon the superior junction of the maxilla and zygomatic bones on the inferior borders of the left and right orbits, with the basicranium placed

on a cranial rest and the viscerocranium facing the observer. Superior bizygomaxillare breadth was included because this measurement describes the greatest superior breadth of the maxilla and its contribution to the lower borders of the eye orbits (Ackermann, 2002, p.171). 10. Bizygomaxillare inferior breadth (bizi) was measured using the lower arms of dial calipers and placing them upon the most inferolateral right and left junctions of the zygomatico-maxillary suture with the neurocranium placed on a cranial rest and the anterior basicranium and palate facing the observer. Inferior bizygomaxillare breadth was included because this dimension depicts the greatest inferior breadth of the maxilla and this region is associated with vital mastication muscles and their attachment (Ackermann, 2002, p.171). Furthermore, this measurement is similar or near approximate to bimaxillary breadth (zygomaxillare, zm-zm; Wood, 1991 - no. 53, p. 289), which is also reported for some fossil specimens. 11. Zygomatico-maxillary suture length (superior zygomaxillare to inferior zygomaxillare; zs-zi) was measured using the lower arms of dial calipers, from zygomaxillare superior to zygomaxillare inferior. Position or hold cranium as necessary. This measurement was chosen to document the chord length of the zygomatico-maxillary suture (Ackermann, 2002, p.171). 12. Maximum length of the zygomatic bone (zs-zgyi) was measured with the lower arms of dial calipers from superior zygomaxillare to the posteroinferior point of the zygo-temporal suture along the zygomatic arch. Position or hold cranium as necessary. This measurement was included because the inferior border of the zygomatic bone is an important region associated with masticatory muscles, particularly the masseter muscles (Ackermann, 2002, p.171). 13. Superior facial length (basion to prosthion; bas-pros) was measured with spreading calipers with the skull resting on its left lateral side on a cranial rest and the basicranium facing the observer. The distance between basion and prosthion was chosen because this dimension describes the size and projection of the face (Wood, 1991 - no. 44, p. 289). 14. Inferior cranial length (basion to nasion; bas-nas) was measured with spreading calipers; if nasion was a complicated area with ossicles, the most superiomedial

point of the discernable nasal bones and their junction with the frontal bone was nasion. Position or hold cranium as necessary. The length between basion and nasion was included because this measurement represents the area of juncture between the viscero- and neurocranium (Wood, 1991 - no. 5, p. 287). 15. Cranial height (basion to bregma; bas-br) was determined with spreading calipers; if bregma was a complicated juncture between the frontal and parietals, the most posteromedial part of the frontal was taken to be bregma, including the sagittal crest if present (Whitehead et al, 2005). Position or hold cranium as necessary. Cranial height was chosen because this dimension represents the cranium’s greatest height in the median plane (Wood, 1991 - no. 4, p. 287). 16. Superior facial height (nasion to prosthion; nas-pros) was measured using the lower and/or upper (depending on size of specimen) arms of dial calipers depending on primate genera. Position or hold cranium as necessary. The chord length between nasion and prosthion was included because this measurement depicts the greatest length of the viscerocranium in the median plane (Wood, 1991 - no. 43, p. 289). 17. Frontal sagittal chord (nasion to bregma; nas-br) was measured using the lower arms of dial calipers, including the sagittal crest if present. Position or hold cranium as necessary. The frontal, parietal and occipital sagittal chord lengths were included to depict the dimensions of the neurocranium in the median plane, representing homologous dimensions across all primates. 18. Parietal sagittal chord (bregma to lambda; br-lam) was measured using the lower arms of dial calipers, although if lambda is a complicated area with ossicles the most superiomedial point of the occipital where it meets the parietals was considered lambda. Position or hold cranium as necessary (Wood, 1991 - no. 25, p. 288). 19. Occipital sagittal chord (lambda to opisthion; lam-opn) was measured using the lower arms of dial calipers. Position or hold cranium as necessary (Wood, 1991 - no. 39, p. 288). 20. Sagittal length of the nasal bones (nasion to rhinion; nas-rhi) was measured using the lower arms of dial calipers from nasion to the rhinion. Please note, in some genera, particularly Rhinopithecus (also observed in Pygathrix and Pongo), the

nasal bones may or may not be present (Wang et al, 1998). For some specimens, nasion may be a complicated area with ossicles. If so, the most mediosuperior point of the discernable nasal bones and their juncture with the frontal bone was considered nasion. Position or hold cranium as necessary. The sagittal length of the nasal bones and nasal aperture (no. 22, below) was included because the nasal region and its configuration are highly diagnostic for many primate genera (Verheyen, 1962; Wood, 1991 - no. 71, p. 290). 21. Inferior breadth of the nasal bones (inbrnabo) was measured with the lower arms of dial calipers placing each arm on the most right and left lateral points of the nasal bones where they meet the nasal aperture. Place the basicranium on a cranial rest with the viscerocranium facing the observer. The inferior breadth of the nasal bones was included to examine the nasal bones contribution to the superior border of the nasal aperture (Wood, 1991 - no. 74, p. 290). 22. Sagittal length of the nasal aperture (rhinion to nasospinale; rhi-ns) was measured using the upper arms of dial calipers. Position or hold cranium as necessary (Wood, 1991 - no. 70, p. 290). 23. Maximum width of the nasal aperture (maxnawi) was determined using the upper arms of dial calipers. Place the basicranium on a cranial rest with the viscerocranium facing the observer (Wood, 1991 - no. 68, p. 290). Naso-facial measurements are important because they are highly diagnostic between species (e.g. Verheyen, 1962) and can be influenced by climate (e.g. Harvati & Weaver, 2006). 24. Nasal height (nasion to nasospinale; nas-ns) was measured using the upper arms of dial calipers. Position or hold cranium as necessary. The distance between nasion and nasospinale was included because this dimension describes the length of the nasal region only and not the further projection of the premaxilla (Wood, 1991 - no. 69, p. 290). 25. Maxillo-alveolar length (prosthion to distal M3; maxalvlen) was determined with spreading calipers from prosthion to the central midpoint of an imaginary line connecting the right and left points where the maxillo-alveolus meets distal M3, with the skull resting on its left lateral side on a cranial rest and the basicranium

facing the observer. Please note, Wood (1991; no. 87, p. 292) measures to the maxillary tuberosities, however, measuring to distal M3 proved to be a more definable and reliable point between species and many specimens’ maxillary tuberosities were damaged or perhaps very smooth and difficult to demarcate posteriorly. The maxillo-alveolar length was included because this dimension depicts the area necessary for the entire dental arcade in the median plane. Furthermore, the differences between these landmarks (distal M3 and maxillary tuberosity not terribly great. 26. Maxillo-alveolar breadth (or, biectomolare breadth or outer alveolar breadth; biecm) was measured using dial calipers by placing the lower arms on the maxillary alveoli just superior to the approximate center of M2, with the neurocranium placed on a cranial rest and the anterior basicranium and palate facing the observer. Biectomolare breadth is included because this measurement represents the maximum width of the maxilla including the contribution of the postcanine dentition perpendicular to the median plane (Wood, 1991 - no. 88, p. 292). 27. Incisivion to the palatomaxillary suture (iv-pms) - Length between the most posteriomedial point of foramen incisive (or incisive canal) to the central intersection of the palatomaxillary suture was measured using the upper arms of dial calipers, with the neurocranium placed on a cranial rest and the anterior basicranium and palate facing the observer. The chord length between incisivion and the central intersection of the palatomaxillary suture was chosen because this dimension describes the length of the maxilla only and not the contribution of the premaxilla and palatine bones in the median plane (Wood, 1991 - no. 92, p. 292). 28. I1- I2 alveolar length (prosthion to the septum between I2 & C1; i1i2) was measured using the lower arms of the dial calipers from prosthion to the septum between I2 and C1. Please note, if large diastema exists between I2 and C, measure from prosthion (pros) to the maxillo-premaxillary suture in line with the dental arcade). I1- I2 alveolar length was included because this dimension records the area needed to house the medial and lateral incisors excluding the canines. Position or hold cranium as necessary (Wood, 1991 - no. 94, p. 292).

29. Internal breadth between the upper canines (bicanin) was measured using the upper arms of dial calipers with the tips of the arms on the palatal alveolus surrounding the canines perpendicular to the median plane with the neurocranium placed on a cranial rest and the anterior basicranium and palate facing the observer. Internal breadth between the upper canines was included to record the breadth of the anterior portion of the palate. 30. External breadth of the upper canines or intercanine distance (Wood, 1991 - no. 98, p. 293) (bicanex) was measured using the lower arms of dial calipers with the tips of the arms on the maxillary alveolus surrounding the canines perpendicular to the median plane with the basicranium placed on a cranial rest and the viscerocranium facing the observer. External breadth between the upper canines was included to record the maximum breadth of the anterior portion of the maxilla including the area necessary for the canines. 31. Palatal height at M2 (palhei) was determined using coordinate calipers by placing the long arms near the left and right ectomolare, allowing the measuring arm to be centered on the palatal roof with the neurocranium placed on a cranial rest and the posterior basicranium and occiput facing the observer (please note, Wood, 1991 measures palatal height at M1, no. 103, p. 293). 32. Biseptal breadth (biseptal) was measured using the lower arms of dial calipers. Please note, if large diastema exists between I2 and C, measure from prosthion to the maxillo-premaxillary suture centered in line with the dental arcade. Position or hold cranium as necessary. 33. Palatal breadth (biendomolare breadth; bienm) was measured using the lower arms of dial calipers not on the M2 with the tips of the arms on the maxillary alveolus surrounding M2 perpendicular to the median plane with the neurocranium placed on a cranial rest and the anterior basicranium and palate facing the observer. Biendomolare breadth was included as this measurement describes the maximum breadth of the palate excluding the contributing dimensions of the postcanine dentition (Wood, 1991- no. 91, p. 292). 34. Palatal length A was measured using the upper arms of the dial calipers from orale to staphylion (ol-sta) with the skull resting on its left lateral side on a cranial rest

and the basicranium and palate facing the observer. Palatal length A was chosen because this dimension is the maximum length of the entire palate in the median plane (Wood, 1991 - no. 89, p. 292). 35. Palatal length B was measured using the upper arms of the dial calipers from orale to the central intersection of palato-maxillary suture (ol-pms) with the skull resting on its left lateral side on a cranial rest and the basicranium and palate facing the observer. Palatal length B was recorded because this distance provides the length of the palate minus the contribution of the palatines in the median plane (Wood, 1991 - no. 90, p. 292). 36. Interentoglenoid breadth (bien) was measured using dial calipers by placing the lower arms on the tips of the left and right entoglenoid processes with the neurocranium placed on a cranial rest and the anterior basicranium and palate facing the observer. Interentoglenoid breadth was chosen because it represents a vital dimension of the basicranium and its relationship with the mandible (Wood, 1991 - no. 15, p. 288). Other cranial measurements (orbital height and width, interorbital pillar width, superior breadth of the nasal bones and bi-mastoid breadth) were initially collected but were later excluded from the analyses because 1) there was a question of their accuracy and repeatability, 2) they were highly variable 3) specimens were damaged or 4) measured incorrectly by the author and were thus omitted.

br fmt

g lam nas fmo

zs o rhi aur

ns por zgyi zi pros ast

opn bas ol

ecm

iv pms

zi zi zi

sta enm

dm3 zgyi zgyi

en en bas por por aur aur

opn ast ast

Figure 2.1: Adapted from Lang (1923) with some modification. A. Norma lateralis. B. Norma inferior.

From Ackermann, 2002

From Singleton, 2002

Figure 2.2: Arrows indicate abbreviations used within this thesis. Other measurements and abbreviations are from Wood, 1991; Buikstra & Ubelaker, 1994; Bass, 1995; & White, 2000.

2.5 The Craniometric Sample: The craniometric sample collected for this study contains representatives from all of the genera of the Catarrhini according to the taxonomy of Groves (2001a). The goal of this data collection was to document the comparative cranial diversity that can exist among populations of genera and species. The craniometric sample captures, morphometrically, broad macroevolutionary trends. This is validated by the results of discriminant function analysis of raw data. Whether entire multivariate dataset or partitioned groups (superfamily, family, subfamily, tribe and subtribe) more than 90% of individuals are correctly classified to their respective genus, cross-validated using SPSS 11.0. Thus, it fair to say that there must be some biological reality to the morphometric content of this dataset and genus level classification. Nevertheless, it must be admitted, the craniometric samples cannot possibly represent the full range of variation because some generic samples do not contain all species representatives. In addition, it was sometimes necessary to include a subadult specimen because no other specimens were available. Please note, the limited subadults (i.e. individuals with their third molar not fully erupted to the occlusal plane) included less than twenty individuals and had their third molar at least in the process of erupting. Although, if a particular measurement for a subadult fell well outside the range of adults, the dimension was excluded and the mean of the sample was substituted; this was only necessary for very few individuals. However, where deficiencies exist other published accounts from the primatological and anthropological literature can supplement results and conclusions. Furthermore, when pooled sex samples are less than ten, as is the case with some genera (Allenopithecus, Procolobus, Semnopithecus and Simias), results related to these genera should be considered approximate but are included nonetheless for heuristic and comparative purposes. In addition, genera with small sample sizes (<10) or subadults were both included and excluded from the multivariate statistic procedures to measure their impact. This revealed whether or not these genera skewed the results, which they did not, and so are included and reported.

2.5.1 Primates, Haplorrhini, Simiiformes, Catarrhini, Cercopithecoidea, Cercopithecidae, Cercopithecinae, Cercopithecini (n=89): The tribe’s craniometric sample is composed of 89 adults with subequal numbers of male and female adults, with a few subadults. While the samples for Erythrocebus, Allenopithecus and Miopithecus include all recognized species (although the first and second are monotypic genera) the same cannot be said for Cercopithecus and Chlorocebus. The sample for Cercopithecus includes only nine species although Groves (2001a) recognizes a potential 25. Thus, the sample for Cercopithecus is not exhaustive and cannot possibly represent the full range of variation within this genus. The same must be admitted for Chlorocebus, which is comprised mainly of vervets from South Africa even though Groves (2001a) has distinguished six species. However, this allows the comparison of samples from geographical extremes of supposed closely related species. That is, the specimens for Cercopithecus include species from tropical equatorial Africa, while specimens for Chlorocebus are from South Africa. Other previous craniometric studies of the guenons and their allies include the work of Verheyen (1962), Hill (1966), Vogel (1966 & 1968), Alexander (1981), Martin & MacLarnon (1988), Shea (1992) Jungers et al (1995) Ravosa & Profant (2000) and Cardini et al (2007). Table 2.2: Craniometric sample for the Cercopithecini. Allenopithecus Craniometric sample includes four males and two females, three of which were subadults nigroviridis (n=6). Miopithecus spp. Craniometric sample includes both species, M. talapoin (-8, -7) and M. ogouensis (- 1, -3) (n=19, -9, -10). Cercopithecus spp. Craniometric sample includes nine species, C. ascanius (-1, -2), C. erythrogaster (- 1, -1), C. mitis (-2, -2), C. neglectus (-2, -2), C. diana (-1, -1), C. petaurista (-2), C. cephus (-1), C. nictitans (-2, -2) and C. wolfi (-2) (n=24, -12, -12). Chlorocebus spp. Craniometric sample includes specimens mainly C. p. pygerythrus from Southern Africa although two male C. p. hilgerti were also included (n=24, -13, -11). Erythrocebus patas Craniometric sample includes nine males and seven females (n=16).

2.5.2 Primates, Haplorrhini, Simiiformes, Catarrhini, Cercopithecoidea, Cercopithecidae, Cercopithecinae, Papionini (n=143): The tribe’s craniometric sample is represented by 143 adults with a few subadults. While the samples for Mandrillus and Theropithecus document all species, the same cannot be said for Cercocebus, Macaca, Lophocebus or Papio. The sample for Cercocebus contains representatives from all species except C. sanjei. The sample for Macaca includes only nine species although Groves (2001a) recognizes a possible twenty. The sample for Lophocebus includes mainly L. albigena, with only three L. aterrimus and no specimens of L. opdenboschi. The sample for Papio comprises mainly specimens of P. ursinus although there are some representatives of P. anubis, P. papio and P. cynocephalus but P. hamadryas is not represented. Other craniometric studies of the papionins include the work of include Hill (1970 & 1974), Albrecht (1978), Groves (1978), Alexander (1982), Cheverud (1989), Fooden & Albrecht (1993); Martin (1993), Setchell et al (2001); Singleton (2002 & 2004a), Pan et al (2003), Frost et al (2003), Pan & Oxnard (2004), Hamada et al (2005) and Elton & Morgan (2006).

Table 2.3: Craniometric sample for the Papionini. Cercocebus spp. Craniometric sample includes four species - C. atys (-1, -2), C. torquatus (-4, -5), C. agilis (-5, -1) and C. galeritus (-1) (n=19, -10, -9). Mandrillus spp. Craniometric sample includes both species, M. sphinx (-3, -7) and M. leucophaeus (-5, -7) (n=22, -8, -14). Macaca spp. Craniometric sample includes ten species, M. sinica (-1), M. nigra (-2, -4), M. arctoides (-4, -3), M. nemestrina (-3, -1), M. ochreata (-1, -1), M. assamensis (-1, -1), M. fascicularis (-2, -3), M. hecki (-2), M. mulatta (-1, -1), M. silenus (-1), and M. thibetana (-2) (n=34, -15, -19). Lophocebus spp. Craniometric sample includes specimens mainly of L. albigena (-9, -11) but a few L. aterrimus (-2, -1) are included (n=23, -11, -12). Papio spp. Craniometric sample for Papio is mainly comprised of P. ursinus (-12, -7) from South Africa, although some P. anubis (-4, -4), P. cynocephalus (-2, -1) and P. papio (-1) were collected (n=31, -19, -12). Theropithecus gelada Craniometric sample includes seven males and seven females (n=14).

2.5.3 Primates, Haplorrhini, Simiiformes, Catarrhini, Cercopithecoidea, Cercopithecidae, Colobinae (n=196): Other craniometric studies of the African colobines include the work of Schultz (1958), Verheyen (1962), Hull (1979) and O’Higgins & Pan (2004). Further craniometric studies of the Asian langurs include the work of Hooijer (1962), Aimi & Bakar (1992), and Pan & Groves (2004). Lastly, craniometric studies of the Asian odd-nosed colobines include the work of Groves (1970) and Pan & Oxnard (2001).

Table 2.4: Craniometric sample for the Colobinae. Colobini, Colobina Craniometric sample includes three species - C. angolensis (-2, -2), (n=59), C. polykomos (-1, -1), and C. guereza (-12, -8) Colobus spp. (n=26, -15, -11). Piliocolobus spp. Craniometric sample includes four species - P. badius (-7, -6), P. rufomitratus (-5, -1), P. tephrosceles (-3, -1) and P. tholloni (-3, -2) (n=28, -15, -13). Procolobus verus Craniometric sample includes three males and two females (n=5). Presbytini, Presbytina Craniometric sample includes three species - S. entellus (-2, -1), (n=60), Semnopithecus S. schistaceus (-1, -1) and S. priam (-2, -2) (n=9, -5, -4). spp. Trachypithecus spp Craniometric sample includes six species - T. obscurus (-2, -2), Tr. pileatus (-1, - 3), T. phayerei (-2, -2), T. cristatus (-6, -7), T. (Kasi) johnii (-1, -1), and T. (Kasi) vetulus (-3, -3) (n=33, -15, -18). Presbytis spp. Craniometric sample includes five species - P. rubicunda (-6, -6), P. potenziani (- 2), P. melalophos (-1), P. natunae (-2), and P. thomasi (-1) (n=18, -8, -10). Presbytini, Nasalina Craniometric sample includes both species, P. nemaeus (-6, -5) (n=77), and P. nigripes (-8, -6) (n=25, -14, -11). Pygathrix spp. Rhinopithecus spp. Craniometric sample includes two species - R. roxellana (-3, -10) and R. avunculus (-2, -2) (n=17, -5, -12). Nasalis larvatus Craniometric sample includes eighteen males and ten females (n=28). Simias concolor Craniometric sample includes two males and five females, two of which were subadults (n=7).

2.5.4 Primates, Haplorrhini, Simiiformes, Catarrhini, Hominoidea (n=144): The superfamily is represented by 144 adults with very few subadults. The craniometric samples for the Hominoidea are not exhaustive, nor do they contain the full range of variation within these genera. The sample for Hylobates consists of seven species from four subgenera although Groves (2001a) has distinguished a possible 14. The sample for Pongo includes specimens of only P. pygmaeus (Bornean orangutan) and none for P. abelii (Sumatran orangutan). The sample for Gorilla contains specimens of only western lowland gorillas (G. gorilla) and the sample for Pan contains mainly specimens of Pan troglodytes although one female Pan paniscus was included. The sample for modern Homo contains adults from North America, South Africa and East Asia. Other craniometric studies of the hominoids include the work of Krogman (1969), Creel & Preuschoft (1971 & 1976), Cramer (1977), Shea (1985), Shea & Coolidge (1988), Groves et al (1992), Uchida (1996), Ackermann (2002), Albrecht et al (2003), Guy et al (2003), Miller et al (2004), Preuschoft (2004) and Brown et al (2005).

Table 2.5: Craniometric sample for the Hominoidea. Hylobatidae, Craniometric sample includes seven species from all four subgenera – Hylobates spp. H.(Hy.) lar (-4, -2), H. (Hy.) agilis (-1) and H. (Hy.) muelleri (-1, -4); H.(S.) syndactylus (-8, -5); H.(Ho.) (-1, -1); H.(N.) concolor (-1), H.(N.) gabriellae (-1) and H.(N.) leucogenys (-1, -1) (n=31, -16, -15). Hominidae, Craniometric sample includes only Pongo pygmaeus, fourteen males and twelve females Ponginae, (n=26). Pongo pygmaeus Hominidae, Craniometric sample includes only Gorilla gorilla, seventeen males and seventeen females Homininae, Gorilla (n=33). gorilla Pan spp. Craniometric sample includes specimens consists mostly of Pan troglodytes and one female Pan paniscus (n=33, -19, -14). Homo sapiens Craniometric sample includes eleven males and nine females from modern populations (N. America, South Africa and East Asia) (n=20).

2.6 Statistical Methods: 2.6.1 Descriptive, Univariate and Bivariate Statistics: The most commonly employed methods for making statistical inferences in biology are the parametric tests for mean differences, such as Student’s t-test and Analysis of Variance. These tests make several assumptions about the data under study. Important among them is the assumption that they follow a normal (Gaussian) distribution, and that variances and sample sizes are similar. Failure to meet these assumptions results in spurious results under a parametric model. They additionally assume that the data under study represent random samples and are independent. The first of these assumptions is difficult to test directly in paleontology. Fossil samples represent individuals whose remains have survived complex and unknown taphonomic histories. Thus, they might be biased in some unknown way, such as towards larger-bodied individuals such as males, or sample individuals or taxa from a narrow segment of their geographic or temporal distribution. There is simply no way of controlling for these factors, and in the absence of a complete record for hominins, fossils must and usually do get treated as random samples. The assumption of independence essentially requires that the value of one datum is unrelated to any other datum (i.e. knowing the value of one observation gives you no information about the value of any other). This is reasonable and uncontroversial here. Univariate data and descriptive statistics (mean, median, maximum (Max.), minimum (Min.), range, standard deviation (SD) and coefficient of variation (CV) of modern skeletal elements of known provenience provide many basic functions in comparative analysis and particularly in relation to biological classification of fossil material (Wood, 1991; Plavcan, 1993; Plavcan & Cope, 2002). In addition, examination of descriptive and univariate statistics identifies any errors in the craniometric datasets. Males and females were pooled together so as to capture and experience the full range of morphological variation and affects of sexual dimorphism within and between genera. Box plots are presented because they allow for quick assessment as to a sample’s spread and possible skewness, as well as identifying significant differences where no overlap of the

boxes occurs. Genera are arranged by mean generic body weight (grams - g) from least to greatest (left to right). Cranial variables and indices which distinguish at least two or more genera via examination of box-plots (i.e. where no overlap of boxes occur) were then subject to three different analyses.

2.6.2 Shapiro-Wilk Test: First, each generic sample was tested for normality by Shapiro-Wilk. If normality could not be assumed, that particular cranial variable was later excluded from one-way Analysis of Variation (ANOVA). The Shapiro-Wilk Test is a simple one-sample method for testing the assumption that a sample follows a normal distribution. In this study, Shapiro-Wilk tests were performed using the program PAST. The level of significance was set at p=0.05. Probability values shown indicate the probability that a sample is normally distributed (e.g. p=0.77 provides a probability of 77% that the sample is normally distributed).

2.6.3 Kruskal-Wallis Test: Second, to test statistical significance, all variables were subject to Kruskal-Wallis, which is a generally considered non-parametric ANOVA (Hammer et al, 2006). That is, it allows for comparison of multiple, randomly drawn, independent samples, and makes no assumptions about the data (i.e. normality and similar variances and sample sizes). It differs from ANOVA in providing a test of the equality of medians (instead of means). The Kruskal-Wallis procedure defines a ratio symbolized by H: numerator is the observed value of the sum-of-squares between group deviates (SSbg) based on ranks and the denominator is the mean of the sampling distribution of SSbg. As with ANOVA, the Kruskal-Wallis test does not indicate which of the samples compared show significantly (median) differences. Again, a post hoc test making pair-wise comparisons is required. Commonly, Mann-Whitney U tests are used for this purpose. Moreover, the Mann-Whitney test is usually altered with Bonferroni correction of p in order to maintain the overall probability of a Type I error at p=0.05. Here, Kruskal-Wallis tests were performed using PAST. The level of significance was set at p=0.05. The results of Bonferroni-corrected Mann-Whitney U-tests are also

given in PAST to allow assessment of pair-wise comparisons. Matrices of these the results are presented below.

2.6.4 One-way ANOVA Lastly, Where samples were found to follow a normal distribution using the Shapiro-Wilk test, the parametric inferential method Analysis of Variance (ANOVA) was employed to test the null hypothesis that sample means are identical. While the Student’s t- test is restricted to paired (two-sample) comparisons, ANOVA allows for multi-group comparisons to be undertaken (i.e. the means of more than two independent samples can be compared in a single test). When large numbers of paired tests of mean difference are undertaken it is a statistical inevitability that about 5% of such tests will be wrong (i.e. result in Type-I errors: rejection of a null hypothesis that is actually correct). For example, if 100 t-tests are undertaken then about 5 of them are expected to represent Type-I errors when the significance level is set at p=0.05. ANOVA allows this ambiguity to be overcome when more than one comparison is made. ANOVA essentially allows us to ask the question: Is there an overall indication of mean differences among the various samples? ANOVA is based on the comparison of variability (sums-of-squares) between groups (SSbg) and within groups (SSwg), the difference between these variance estimates provided by Fisher’s F-distributions. If the variance between two groups is large, the difference between means will be large, and the F-ratio will also be large. A major limitation of ANOVA is that is fails to indicate which of the samples compared are significantly different. To determine this, an additional (post hoc) test making pair-wise comparisons is required. Commonly, the Tukey’s HSD (Honest Significant Difference) is employed for such purposes as it is generally considered more robust than a t-test; being more stringent, and decreasing the probability of obtaining significant results by chance alone. Here, one-way ANOVA tests were performed using PAST. The level of significance was set at p=0.05. PAST also tests for equality of variance in all ANOVA’s, and only where this assumption was violated will these results be discussed in the test. Tukey’s HSD pair-wise comparisons are also automatically undertaken in PAST and the results (p-values) are presented in matrices in the results. Cranial variables which have a

normal distribution were subject one-way ANOVA. However, because of their small sample sizes, Allenopithecus, Procolobus, Semnopithecus and Simias were excluded from all ANOVA analyses but were examined via Shapiro-Wilk and Kruskal-Wallis. To examine the relationship between cranial size and body weight, the pooled sex mean for particular cranial dimensions were plotted against mean generic body weights collected from the published literature. Bivariate plots of body weight (x-variable) to cranial variables (y-variable) are a useful tool for examining large-scale macroevolutionary trends, including allometric trajectories and niche separation (Aiello & Day, 1982; Benefit & McCrossin, 1993; Aiello & Wood, 1994; Jungers et al, 1995).

2.7 Multivariate Statistics: 2.7.1 Principal Component Analysis (PCA): (cite Relethford, 1994 & 2002) Principal Components Analysis (PCA) is a robust ordination multivariate statistical procedure which can reduce a large number of variables to a few very important ‘principal components’ (PC) based on the variance-covariance (or correlation) matrix of the multivariate dataset. However, despite data reduction, very little information is lost. PCA has been applied successfully to numerous biological and anthropological studies for both morphometric and ecological data with significant results (Andrews & Williams, 1973; Lessertisseur et al, 1974; Albrecht, 1978; Howells, 1989; Strasser, 1992; Shea, 1992; Shea et al, 1993; Fleagle & Reed, 1996; Pan et al, 2003; Harvati, 2003; Frost et al, 2003; O’Higgins & Pan, 2004; Pan & Groves, 2004; Miller et al, 2004; Hamada et al, 2005). PCA was employed within this study because this technique has several features which are amenable to cranial morphometrics and their interpretation as well as identifying similar cases that may cluster together because of functional similarities or phylogenetic relationships. For example, linear measurements of cranial dimensions are highly interrelated which can produce spurious or false readings through redundancy. Complex interrelationships such as these are generally referred to as morphological integration or modularity (Olson & Miller, 1958; Cheverud, 1982; Ackermann & Cheverud, 2004b; Goswanji, 2006; Polanski & Franciscus, 2006). Fortunately, the computation for PCA uncorrelates each variable by creating linear combinations (or functions) of the original variables. These linear combinations maximize the inherent variation of the multivariate

dataset. In addition, PC’s are independent from each other by way of plotting axes perpendicular (or orthogonal) to one another. In theory, these PCs describe the underlying structure of the multivariate dataset. What’s more, these patterns may represent both phylogeny and adaptation. However, as with most statistical techniques there are assumptions and limitations to the methods. Multivariate statistics rely on conditions such as large representative samples, multivariate normality, and for some methods, the same units of measurements. Since this thesis is concerned with cranial variables which expose maximum generic differences or separation, only PCs 1 & 2 and PCs 2 & 3 will be discussed for Ln transformed data, whereas results for MSV will be limited to PCs 1 & 2 because size has been controlled for and PCs 2 & 3 rarely explain a large part of the variation between genera.

2.7.2 Canonical Variates Analysis (CVA): Canonical variates analysis (CVA) is another rigorous multivariate statistical technique which has some similarities to PCA, as well as sharing theoretical assumptions, although there are significant differences. As Albrecht (1978) has succinctly expressed, “The critical difference in the two techniques is that canonical analysis relies on the belief that it is of value to weight differences between the groups using the within-group dispersion as a standard of comparison” (p.37), which PCA does not (see also Albrecht, 1980). Within biological anthropology, CVA has been effectively applied to many primate genera and species and has produced statistically significant results (Albrecht & Miller, 1993). The computation of PCA and CVA produce similar results and graphical representation; and is used here in conjunction with PCA. In addition, both methods can be used to gauge the results of each other. Like PCA, CVA maximizes the variation between groups. Recently, Harvati (2003) has combined results of PCA and CVA demonstrating significant and interesting differences between Neandertals and Modern humans in temporal bone morphology. CVA has been successively applied to many primate species thus far studied morphometrically (cranial and postcranial), which have produced statistically significant results (Albrecht & Miller, 1993). CVA (is essentially multi-group discriminant function analysis; Albrecht, 1980; Hammer et al, 2006), like PCA, creates new, uncorrelated variables which are linear combinations of the original variables. Similar

to results for PCA, only CV axes 1 & 2 will be examined and discussed because these two axes explain the majority of the variation between genera while subsequent axes explain much less.

2.8 Morphological and Genetic Distances: To summarize the total morphological variation within a genus’ multivariate dataset, Euclidean distances between all individuals were recorded and the averages, maximums and minimums for each genus were presented. This summarizes the extent to which members of within a genus differ from other congenerics. Then, to measure morphological divergence between genera, again, Euclidean distances based on generic mean PCA and CVA object scores (for both Ln transformed and MSV data) from the first five PCs or CV axes were recorded generated to create inter-generic distance matrices. Generating Euclidean distances in this manner is very similar to Mahalanobis distances (Mimmack et al, 2001; Elmore & Richman, 2001). This allows for quick assessment as to how much one genus has diverged from other closely allied genera.

Morphological Distance – M: In addition, inter-generic M distances were computed for all groups studied. The “M” distance has been applied to several orders of animals with robust results (Cherry et al, 1978 & 1982; Atchley et al, 1980; Wilson et al, 1984). This computation provides evidence of shape divergence not size because each measured trait is converted into a proportion of the geometric mean, which is very similar to Mosimann shape variables. In addition, this statistic also utilizes the standard weighted deviation which corrects for small sample sizes. As Cherry et al (1978) describe, “M [morphological difference (M)] is the average number of standard deviations by which the two species [or in this analysis, genus] differ per trait measured” (p. 210).

Genetic Distances: Like morphological distances, genetic distances also summarize numerous variables and reduce data to one figure. However, unlike morphological distances, genetic distances increase with time and the same cannot be assumed for morphological features. Genetic

distances were compiled from the published literature (Tosi et al, 2003 & 2004; Page et al, 1999; Page & Goodman, 2001; Whittaker et al, 2006).

2.9 Data Transformation: Despite all measurements being recorded using the same unit of measurement, millimeters (mm), most statistical procedures advise or require data transformation prior to analysis. This is necessary for a number of reasons. Obviously the largest source of difference and variation between genera and species is absolute size (Jungers et al, 1995). In fact, larger species have larger ranges of variation (Albrecht, 1978). Accordingly, steps must be taken to control and account for the influence of size but it cannot be eliminated completely. Another consideration is the possibility of skewness in the data due to perhaps small sample size, sizes differences between males and females due to sexual dimorphism, or whether the dataset has multivariate normal distribution, as well as, similar variance- covariance patterns; the latter of which cannot be assumed for even closely related species (Ackermann, 2003). To adjust and comply with these requirements, prior to multivariate analysis all raw data was first examined for errors and then natural log (Ln) transformed. By doing this all measurements are distributed linearly with some correction by removing skewness as well as providing log normal distribution. In addition, the raw data was also converted to Mosimann Shape Variables (MSV); a method which in theory controls for size within the multivariate dataset (Nadal-Roberts & Collard, 2005). This is accomplished by computing the geometric mean of all measurements for one individual followed by dividing each measurement by the geometric for that individual (see Mosimann, 1970; Mosimann & James, 1979; Darroch & Mosimann, 1985; Jungers et al, 1995). Thus, two sets of multivariate analyses (PCA and CVA) were conducted on 1) A Ln transformed multivariate dataset; and 2) A size corrected multivariate dataset (Mosimann Shape Variables; MSV).

2.10 Structure of Results: Results reported in succeeding chapters are grouped at various taxonomic levels (tribe - Cercopithecini and Papionini; subfamily - Colobinae; and superfamily - Hominoidea) for comparative purposes. For example, the genera which are grouped within the tribe Cercopithecini are only compared with other genera of that tribe and so forth.

These groupings represent the comparative boundaries (defined by phylogeny, morphology, genetics, ecology and behavioral data) for the genera under study. By applying this criterion, group comparisons are naturally composed by a biological yardstick approach. Comparisons outside of these groupings are unnecessary because there are no generic taxonomic disagreements between these groupings. In other words, there are no taxonomic arguments for the inclusion of the Colobus within Papio or Hylobates and vice versa. Instead, differing generic taxonomic schemes are within these groupings. For example, within the Papionini tribe, Goodman et al (1998) suggests (based on genetic data) Mandrillus is a subgenus within Cercocebus. Another example is the gibbons which can be arranged into one genus with four subgenera or purely four genera. Still, another example includes the African colobines; most recognize Colobus as a distinct genus but disagreements remain concerning whether or not to recognize separate genera for Procolobus and Piliocolobus, or each a subgenus within the former due to shared derived characters (Oates et al, 1994). In fact, this last example is also true for the genera of Presbytina and Nasalina (Bennett & Davies, 1994). Due to the large amount of generic comparative data, results must follow similar and repetitive formats for clarity in the synthesis of evidence. First, each genus is briefly introduced noting some characteristic features, including body weight, habitat choice, diet, etc. Second, the taxonomic nomenclature history of the genera under consideration is provided, including differences of opinion and alternative classification schemes. Third, descriptive, box-plots and univariate statistics are presented and discussed for the cranial variables. Then, similar treatment is employed for the cranial indices. Third, a selection of cranial variables and associated sample mean, which significantly sorted genera, was then plotted against mean generic body weights (grams). Fourth, the results of multivariate statistics (PCA and CVA) of all cranial variables are offered and discussed. Fifth, important postcranial, behavioral, locomotion and ecological features are noted and discussed. These other types of evidence are necessary because genera are defined by many variables not including the cranium and many interact directly with a species’ habitat and locomotion. However, a complete review is beyond the scope of this thesis but features which differentiate genera and their adaptive zones are. Thus, data is limited mainly to skeletal elements which may be encountered in the fossil record. Sixth, results from recently

published genetic analyses are discussed and summarized. This is necessary to satisfy Wood & Collard’s (1999a) requirement of genus’ members to be more cladistically related to each than to any other species from another genus (i.e. taking into consideration the inter- and intra-generic genetic relationships). Lastly, each generic synopsis is outlined and evaluated along with morphological descriptions and adaptive features genus; and where necessary, genera which are in need of further refinement are mentioned. After these chapters per grouping are completed the analysis will then consider and evaluate fossil hominin genera based on the available evidence by similar statistical techniques. Through these generic comparisons, this thesis will answer the question - Are the recently proposed hominin genera, such as Sahelanthropus, Orrorin, Ardipithecus and Kenyanthropus, adequately defined and necessary when the extant Catarrhini genera are used as a comparative benchmark?

2.11 Limitations of data and analysis and theoretical assumptions The cranial morphometrics of extant catarrhines are interpreted within the principle of taxonomic uniformitarianism and evolutionary morphology or ecomorphology; both are equally important (Reed, 1997 & 2002; Bromage, 1999). The datasets provide a ‘bird’s eye’ as opposed to a ‘worm’s eye’ perspective (Andrews & Harrison, 2005). The majority of the comparative multivariate craniometric dataset is composed of monkeys from the Cercopithecoidea (n=428) while that of the apes from the Hominoidea is considerably less (n=144). However, this is more a product of macroevolutionary trends rather then deliberate choice. Since the Miocene there has been a dramatic decrease in the number of ape genera and species in comparison to monkeys (Andrews, 1981 & 1992; Fleagle, 1999).

50 Chapter 3: Results for genera of Cercopithecini

Allochrocebus Allenopithecus Miopithecus Cercopithecus Chlorocebus Erythrocebus

Cercopithecini Cercopithecinae Cercopithecidae Cercopithecoidea Figure 3.1: A taxonomic and phylogenetic Catarrhini diagram representing the likely relationships Simiiformes within the Cercopithecini based on Haplorrhini biomolecular and morphological data. Primates

3.1 Introduction: The purpose of this chapter is to report the results for genera of the tribe Cercopithecini. The Tribe Cercopithecini (Gray 1821) includes five genera: Allenopithecus nigroviridis (Allen’s swamp monkey), Miopithecus (Talapoins; 2 species), Cercopithecus (Guenons; 25 species clustered into 8 species-group; three of which contain only one species; the remaining five groups have species numbers ranging from two to six), Chlorocebus (Vervets or Grivets or Green Monkeys; 6 species) and Erythrocebus patas (Red or Patas monkey) (Figure 3.1). The average number of species per cercopithecin genus is 7.0, ranging from one to 25 with a standard deviation of 10.3. However, if the species of Cercopithecus are excluded, average number of species per genus drops to 2.5, ranging from one to six, with a standard deviation of 2.4.

The species of cercopithecin genera range in size from around one kilogram, (Miopithecus spp.) to up to ~ 13 kg (Erythrocebus patas), with Allen’s swamp monkey, the guenons and vervets falling between. Tables 3.1 to 3.5 list several key features which describe Cercopithecini genera and their respective adaptive zones; including body weight, limb indices, diet, social structure, etc. The cercopithecins in general may be characterized as frugivores although animal prey is consumed as well. Although all genera do exhibit some terrestrial behaviors, they nonetheless require trees for sleeping and evading predators. All genera are restricted to sub-Saharan Africa with only Erythrocebus and Chlorocebus sometimes extending into the Sahel. The extant cercopithecins can be viewed as a recent radiation because very few fossils have been discovered and Cercopithecus, Chlorocebus and Erythrocebus are the only genera within the tribe with reported fossil material; although the specific status of these fossils remains to be determined (Leakey, 1976 & 1988; Eck & Howell, 1972; Eck, 1976 & 1977; Pickford, 1987; Harrison & Harris, 1996; Jablonski, 2002; Frost & Alemseged, 2007). Genetic analyses by Goodman et al (1998 & 2001; see also Page et al, 1999) suggest extant lineages diverged from other cercopithecoids ~7-10 million years ago (Ma). Thus, the lack of cercopithecins from the fossil record indicates: 1) fossil species from tropical forested environments are less likely to be preserved in the fossil record and/or 2) tropical forested fossil deposits have not yet been adequately sampled. Table 3.1: Allenopithecus nigroviridis Body Weight1 Social (g) Limb Indices2 Substrate/ Structure (,; Average Locomotion2 Diet2 Habitat2 (SS)2/EQ3 Min.-Max.) 5500, 3700; Intermembral Quadrupedal- Frugivore + Arboreal & Multimale - 3000-6500 84 above branch animal prey, Terrestrial/ multifemale/ Brachial sitting & leaves, roots, Tropical .89-1.2 (62.5g 100.6 walking flowers lowland [wet] adult brain Crural forest using the weight) 97.5 lower strata

1 References for this chapter and chapters 4 to 6 - Weight data collected from Napier & Napier, 1967; Napier, 1981 & 1985; Jenkins, 1990; Rowe, 1996; Delson et al, 2000; Hutchins et al, 2003, Grzimek’s Animal Life Encyclopedia, 2nd edition. - * indicates >1 species, thus generic average is given for both sexes. 2 References for this chapter and chapters 4 to 6 - Data gathered from Napier & Napier, 1967; Napier, 1981 & 1985; Aiello, 1981a; Wolfheim, 1983; Andrews & Aiello, 1984; Schultz, 1986; Jenkins, 1990; Nakatsukasa, 1996; Rowe, 1996; Fleagle, 1998; Ankel-Simons, 1999; Jablonski, 2002; Hutchins et al, 2003, Grzimek’s Animal Life Encyclopedia, 2nd edition; and Primate Info Net, http://pin.primate.wisc.edu. 3References for this chapter and chapters 4 to 6 - Jerison, 1973; McHenry, 1988; Jungers, 1988b; Tattersall et al, 1988, p.103; Kappelman, 1996; Smith & Jungers, 1997; Collard & Wood, 1999. 52

Table 3.2: Miopithecus spp. Body Weight Limb Indices (g) Average Substrate/ (,; (range) Locomotion Diet Habitat SS/EQ Min.-Max.) 1396, 1135 Intermembral Quadrupedal- Frugivore + Arboreal & Multimale - (Mi. talapoin); 84 (82-85) above branch leaves, flowers, Terrestrial/ multifemale/ 900-1400 Brachial sitting & animal prey Tropical forest, 2.49-2.65 97 (93-101) walking mangrove (37.7g adult Crural swamp, lower brain weight 96 (93-102) & middle strata -M. talapoin)

Table 3.3: Cercopithecus spp. Body Weight Limb Indices (g) Average Substrate/ (,; (range) Locomotion Diet Habitat SS/EQ Min.-Max.) 6030.3, 3321*; Intermembral Quadrupedal- Frugivore + Arboreal & 1 male- 4000-9000 82 (77-88) above branch animal prey, Terrestrial/ multifemale & Brachial leaping, sitting leaves, flowers, Tropical forest/ multimale- 95 (87-102) & walking exudates woodland, multifemale/ Crural lower, middle C. nictitans- 93 (88-100) & upper 1.45-1.71 canopy C. pogonias- 1.63-1.96 (63.6-78.6g adult brain weight)

Table 3.4: Chlorocebus spp. Body Weight (g) Limb Indices Substrate/ (,; Average Locomotion Diet Habitat SS/EQ Min.-Max.) 5194, 3568.5*; Intermembral Quadrupedal- Frugivore + Terrestrial & Multimale - 3000-6500 83 above branch fruit, animal Arboreal/ multifemale/ Brachial sitting & prey, insects, Savanna .88-1.18 (59.8g 96.8 walking leaves woodland, near adult brain Crural water from weight – C. 92.7 lowland swamp aethiops) to the dry Sahel to montane forest up to 4500m

Table 3.5: Erythrocebus patas Body Weight Limb Indices (g) Average Substrate/ (,; (range) Locomotion Diet Habitat SS/EQ Min.-Max.) 8817, 4980; Intermembral Digitigrade Frugivore + Terrestrial & 1 male – 4500-13500 91 (89-96) Quadrupedal- seeds, animal Arboreal/ multifemale/ Brachial above branch prey, insects, open tropical 2.19 (106.6g 104 (100-105) sitting & very little grass savanna/ adult brain Crural walking woodland weight) 96 (94-98)

With regard to competing classificatory schemes, most workers now agree that Allenopithecus, Miopithecus and Erythrocebus require full generic rank because of numerous unique features, both behaviorally and morphologically (Groves, 1989; Groves, 2001; Napier, 1981; Grubb et al, 2004; cf. Napier & Napier, 1967; Verheyen, 1962; James, 1960; Chism & Rowel, 1988). However, Jolly (1966) questioned the placement of Miopithecus within the Cercopithecini. For example, unlike other Cercopithecini genera, female talapoins display cyclical sexual swellings. In addition, talapoin monkeys also use more facial-communicative gestures, more akin to the papionins. More recently, Groves (2000) has argued against the allocation of Allenopithecus to the Cercopithecinae. Reasons for this include the swamp monkey’s possession of ‘molar flare’ (a characteristic found among papionins; see Delson, 1975), female sexual swellings (shared with Miopithecus and papionins), its macaque-like body build and relatively long nasal bones. Furthermore, most disagreements about classification are concerned with the species composition of Cercopithecus and whether Chlorocebus is valid or should be considered a widespread polytypic species of Cercopithecus (Grubb et al, 2003; Enstam & Isbell, 2007; Ankel-Simons, 1999). Nevertheless, Groves (1989) lists several features which differentiate vervets from other guenons and aligns them with the patas monkey. Despite this, “Cercopithecus aethiops” continues to be used in recent literature (e.g. Bolter & Zihlman, 2003; Cardini et al, 2007).

3.2 Descriptive Statistics and Univariate Analyses for cranial variables: Table 3.6 provides pooled sex descriptive statistics for the Cercopithecini genera. The descriptive statistics reveal considerable sample overlap of Allenopithecus, Cercopithecus and Chlorocebus. Similarly, Cope (1993) and Plavcan (1993) reported

Table 3.6: Cercopithecini pooled sex. g-o pros-o biast bipor biaur bizygo bifmo bifmt bizs bizi zs-zi zs-zygi Allen. (n=6) Mean 71.9 98.0 40.9 50.9 55.9 63.8 45.9 50.1 27.4 42.9 16.9 34.0 Median 71.2 96.0 40.9 50.0 54.7 60.3 45.7 49.1 27.0 41.4 16.5 32.2 Maximum 77.0 110.0 46.0 58.0 63.0 77.0 50.5 58.1 31.0 48.4 19.6 43.0 Minimum 68.0 87.4 36.4 48.0 52.0 55.0 41.8 45.2 24.0 37.6 14.6 28.4 Range 9.0 22.6 9.6 10.0 11.0 22.0 8.7 12.9 7.0 10.8 5.0 14.6 Standard Deviation (SD) 3.8 9.2 3.4 3.7 4.1 8.4 3.1 4.3 2.5 4.4 2.2 5.2 Coefficient of Variation (CV) 5.3 9.4 8.4 7.2 7.4 13.2 6.7 8.5 9.3 10.3 13.3 15.2

Mio. (n=19) Mean 59.5 71.8 35.8 40.1 43.3 48.1 37.3 42.3 25.0 32.9 11.5 22.6 Median 59.1 72.9 36.0 40.0 43.0 48.0 37.5 42.6 25.4 32.8 11.2 22.7 Maximum 66.0 78.0 40.0 45.3 50.0 58.0 40.4 46.2 28.0 37.5 13.8 27.2 Minimum 56.0 65.0 32.9 35.8 38.2 41.0 32.3 35.9 18.0 28.5 9.7 16.9 Range 10.0 13.0 7.1 9.5 11.8 17.0 8.1 10.2 10.0 9.0 4.1 10.3 SD 2.3 3.3 2.0 2.8 3.4 4.2 2.1 2.5 2.3 2.2 1.2 2.9 CV 3.9 4.7 5.6 7.1 7.9 8.7 5.6 6.0 9.3 6.8 10.1 12.7

Cercop. (n=24) Mean 74.2 99.4 42.7 53.9 57.2 65.4 47.4 53.4 28.8 43.7 17.4 35.4 Median 75.1 98.1 42.6 54.0 56.8 64.0 46.7 52.9 28.7 42.7 17.5 35.7 Maximum 86.2 119.0 49.0 62.0 66.0 82.0 53.0 61.0 34.4 51.6 23.8 44.5 Minimum 67.5 84.0 37.6 47.7 52.0 56.5 42.1 45.5 24.4 38.3 13.3 27.9 Range 18.7 35.0 11.3 14.3 14.0 25.5 10.9 15.5 10.0 13.4 10.5 16.7 SD 4.9 9.8 3.3 3.8 4.2 6.6 3.0 4.9 2.7 3.7 2.1 4.2 CV 6.5 9.8 7.7 7.0 7.3 10.1 6.4 9.1 9.3 8.5 12.3 11.9

g-o pros-o biast bipor biaur bizygo bifmo bifmt bizs bizi zs-zi zs-zygi Chloro. (n=24) Mean 75.8 101.1 42.6 52.7 57.2 65.4 47.6 55.4 32.3 43.2 14.5 33.2 Median 74.6 101.9 42.8 52.2 56.4 62.8 47.2 55.0 33.1 42.2 14.6 33.3 Maximum 88.0 116.3 47.9 62.4 66.3 79.5 55.6 65.6 40.0 52.8 20.7 37.4 Minimum 67.6 84.8 33.9 44.9 50.4 58.9 41.1 47.4 23.9 38.2 11.8 29.2 Range 20.4 31.5 14.0 17.5 16.0 20.6 14.5 18.3 16.1 14.5 8.9 8.2 SD 5.1 8.1 3.4 4.2 3.8 5.9 3.5 4.8 4.0 3.6 1.9 2.7 CV 6.7 8.0 8.0 8.0 6.6 9.1 7.3 8.7 12.3 8.4 13.1 8.1

Erythro. (n=16) Mean 94.6 125.2 46.8 61.0 65.1 75.6 54.4 62.8 27.5 52.2 22.1 44.1 Median 93.0 117.0 45.5 58.2 63.5 71.6 52.9 60.4 29.3 50.3 22.0 42.6 Maximum 106.0 148.0 54.0 73.0 73.0 92.0 60.3 74.0 35.0 60.8 26.8 54.1 Minimum 86.0 110.0 40.0 54.7 59.0 66.0 48.4 55.3 18.2 44.6 18.6 38.5 Range 20.0 38.0 14.0 18.3 14.0 26.0 11.9 18.7 16.9 16.2 8.2 15.6 SD 6.6 13.4 3.7 6.1 4.7 8.8 4.2 6.8 4.8 5.3 2.4 5.6 CV 6.9 10.7 8.0 10.1 7.2 11.7 7.7 10.9 17.5 10.2 11.0 12.8

Table 3.6 continued: Cercopithecini pooled sex. bas-pros bas-nas bas-br nas-pros nas-br br-lam lam-opn nas-rhi inbrnabo rhi-ns maxnawi nas-ns Allen. (n=6) Mean 68.8 54.7 45.1 40.3 44.8 30.3 30.7 20.5 5.6 15.1 10.2 36.1 Median 67.3 55.3 45.0 40.5 44.4 30.0 30.9 19.7 5.7 15.7 10.2 35.1 Maximum 82.0 63.0 47.0 46.4 48.7 33.2 33.8 23.9 6.1 17.9 12.4 41.1 Minimum 57.0 47.0 43.8 34.0 40.5 28.8 27.4 17.7 4.8 12.0 8.1 31.1 Range 25.0 16.0 3.2 12.4 8.2 4.4 6.4 6.2 1.3 5.9 4.3 10.0 SD 9.1 5.7 1.2 5.1 3.4 1.6 2.1 2.8 0.5 2.1 1.8 4.1 CV 13.3 10.4 2.7 12.7 7.6 5.2 6.8 13.8 8.1 13.9 17.4 11.5

Mio. (n=19) Mean 45.6 41.3 38.2 22.1 37.7 29.5 23.1 8.6 4.8 11.0 6.9 19.2 Median 46.9 41.0 39.0 22.2 37.8 29.5 22.7 9.1 4.8 11.1 6.9 19.7 Maximum 53.0 46.0 42.0 23.7 41.7 32.8 26.5 10.8 5.6 13.0 8.1 21.5 Minimum 39.0 36.4 34.0 19.5 34.1 26.3 20.1 5.9 3.7 8.9 5.0 17.2 Range 14.0 9.6 8.0 4.2 7.6 6.5 6.4 4.9 1.9 4.1 3.1 4.4 SD 3.7 2.9 1.9 1.2 2.3 1.6 1.9 1.2 0.6 1.1 0.8 1.2 CV 8.0 7.1 5.1 5.6 6.1 5.4 8.1 14.4 12.6 9.6 12.2 6.4

Cercop. (n=24) Mean 71.4 58.3 48.5 39.1 45.1 34.6 28.5 17.2 8.4 17.9 10.4 35.3 Median 70.0 58.2 48.5 39.4 45.1 34.1 28.4 17.2 8.3 18.0 10.0 35.1 Maximum 91.0 69.7 53.5 52.0 52.4 43.6 32.3 22.2 10.3 22.9 13.1 43.9 Minimum 56.0 49.8 40.1 28.2 38.0 29.6 23.8 11.2 6.6 12.5 7.9 25.6 Range 35.0 19.9 13.3 23.8 14.4 14.1 8.6 11.0 3.7 10.5 5.2 18.3 SD 9.7 5.5 3.0 6.1 3.2 3.3 2.4 3.0 0.9 2.8 1.6 5.0 CV 13.6 9.4 6.2 15.7 7.0 9.4 8.4 17.5 10.9 15.5 15.6 14.2

bas-pros bas-nas bas-br nas-pros nas-br br-lam lam-opn nas-rhi inbrnabo rhi-ns maxnawi nas-ns Chloro. (n=24) Mean 70.6 58.0 49.0 38.0 46.9 35.9 29.4 16.2 5.8 18.3 10.3 34.4 Median 69.6 57.7 48.9 37.9 46.5 36.3 29.3 15.8 6.0 18.3 10.1 34.6 Maximum 81.3 68.6 54.7 44.1 55.5 41.3 35.7 21.7 8.0 22.8 14.5 39.9 Minimum 59.4 51.6 43.5 30.9 39.6 26.5 22.3 11.8 3.4 14.1 8.1 26.4 Range 21.9 17.0 11.2 13.3 15.9 14.8 13.4 9.9 4.6 8.7 6.4 13.5 SD 6.6 4.5 3.3 4.0 3.4 3.5 3.7 2.8 1.1 2.4 1.7 3.6 CV 9.4 7.7 6.8 10.5 7.2 9.8 12.6 17.0 19.5 13.0 16.1 10.6

Erythro. (n=16) Mean 87.5 70.4 54.8 50.8 56.1 40.8 35.7 21.1 9.3 24.7 12.8 45.2 Median 80.1 67.0 55.0 48.7 55.2 41.1 35.0 20.9 8.7 24.6 12.6 43.5 Maximum 109.0 83.0 60.0 64.0 62.2 44.9 44.1 28.1 13.4 33.6 16.0 56.4 Minimum 72.0 62.0 49.0 41.2 51.8 33.6 30.8 13.5 4.8 18.1 11.0 37.9 Range 37.0 21.0 11.0 22.8 10.4 11.2 13.3 14.6 8.7 15.5 5.0 18.5 SD 12.9 6.9 2.8 7.6 3.3 3.4 3.2 4.1 2.3 4.3 1.4 6.7 CV 14.7 9.8 5.1 15.1 5.9 8.2 9.0 19.7 24.9 17.3 11.1 14.9

Table 3.6 concluded: Cercopithecini pooled sex. maxalvlen biecm iv-pms i1i2 bicanin bicanex palhei biseptal bienm ol-sta ol-pms bien Allen. (n=6) Mean 39.8 33.6 14.1 10.9 15.0 25.7 7.2 17.9 16.3 36.9 23.80 33.9 Median 38.0 33.4 14.1 10.7 15.0 25.3 6.9 18.1 16.0 37.4 22.7 33.7 Maximum 44.0 35.1 16.5 12.3 16.0 30.0 10.5 19.2 18.7 43.1 22.7 39.0 Minimum 36.1 32.5 11.9 9.7 14.2 22.3 5.0 15.9 14.3 29.8 27.5 29.0 Range 7.9 2.7 4.6 2.5 1.8 7.6 5.5 3.3 4.4 13.3 22.4 10.0 SD 3.5 1.1 1.7 0.9 0.6 3.0 1.9 1.1 1.6 6.1 5.1 3.6 CV 8.7 3.4 12.3 8.0 4.1 11.8 26.4 6.3 9.7 16.4 2.1 10.8

Mio. (n=19) Mean 25.3 23.3 10.0 7.5 11.4 18.7 5.1 13.1 13.1 20.3 15.2 26.4 Median 25.0 23.2 10.0 7.5 11.3 18.9 5.0 13.1 12.9 20.5 15.5 26.0 Maximum 30.0 25.4 11.9 8.3 13.1 21.0 6.5 13.9 14.5 25.4 17.5 31.0 Minimum 23.0 21.1 8.0 6.5 9.4 15.8 4.0 12.3 11.5 16.6 12.7 22.3 Range 7.0 4.3 3.9 1.8 3.7 5.2 2.5 1.6 3.0 8.8 4.8 8.7 SD 1.9 1.3 1.1 0.4 1.0 1.5 0.8 0.5 0.9 2.3 1.2 2.1 CV 7.5 5.6 11.2 5.9 8.9 7.9 15.3 3.5 7.0 11.2 8.0 7.8

Cercop. (n=24) Mean 38.5 32.7 16.1 11.1 16.2 26.8 8.1 18.4 19.4 33.2 24.9 35.4 Median 37.7 32.8 15.7 11.1 15.8 25.4 7.8 18.4 19.2 32.1 24.7 35.0 Maximum 48.0 38.9 20.0 13.0 20.3 34.6 10.0 21.6 23.7 44.8 36.4 42.7 Minimum 32.0 28.6 12.6 9.5 13.2 21.8 6.0 16.0 15.5 25.1 18.6 29.2 Range 16.0 10.3 7.3 3.5 7.2 12.8 4.0 5.7 8.1 19.7 17.8 13.5 SD 4.6 2.8 2.1 0.9 1.9 3.7 1.1 1.6 2.4 4.9 4.3 3.3 CV 12.0 8.5 13.2 8.2 11.9 13.6 14.0 8.9 12.1 14.7 17.4 9.3

maxalvlen biecm iv-pms i1i2 bicanin bicanex palhei biseptal bienm ol-sta ol-pms bien Chloro. (n=24) Mean 38.7 31.6 16.3 10.6 14.1 25.3 6.7 16.5 18.8 34.2 26.2 34.3 Median 37.9 31.3 16.2 10.7 14.1 26.1 6.5 16.3 18.7 34.6 26.3 33.6 Maximum 45.9 35.5 19.7 11.7 16.6 28.9 8.4 18.3 21.2 38.5 31.2 40.0 Minimum 31.2 28.1 12.7 9.3 12.6 22.3 4.8 14.0 16.2 29.7 22.4 29.2 Range 14.7 7.4 7.0 2.4 4.1 6.6 3.6 4.3 5.0 8.8 8.8 10.7 SD 3.4 1.8 1.9 0.8 1.0 2.2 0.9 1.0 1.2 3.1 2.4 2.8 CV 8.8 5.7 11.7 7.4 6.8 8.8 13.8 6.1 6.6 9.0 9.3 8.1

Erythro. (n=16) Mean 49.2 35.9 20.4 13.8 19.1 32.2 8.0 22.1 21.4 41.6 32.3 40.6 Median 48.2 35.5 20.5 13.8 18.1 30.8 7.6 21.6 21.1 38.7 30.9 41.5 Maximum 58.0 41.8 26.0 16.2 23.1 40.8 11.0 25.5 26.1 56.0 41.2 45.0 Minimum 42.0 31.2 15.0 11.9 14.7 26.0 5.1 19.7 16.8 31.0 22.9 33.2 Range 16.0 10.6 11.0 4.3 8.4 14.8 5.9 5.8 9.3 25.0 18.3 11.9 SD 5.3 3.4 2.5 1.5 2.8 5.3 1.9 2.0 2.7 7.6 5.0 3.7 CV 10.7 9.4 12.1 10.6 14.7 16.6 24.3 8.9 12.8 18.3 15.5 9.0

odontometric overlap for many Cercopithecus species. Coefficient of variation (CV) range from less than 7.0% for variables such as cranial vault length (g-o) and maxillo-alveolar breadth (or maxillary breadth; biecm) to measurements which are near to or greater than 20.0%, such as height of the nasal aperture (rhi-ns), inferior breadth of the nasal bones (inbrnabo) and palatal height (palhei). Figures 3.2 to 3.16 are box-plots for the Cercopithecini cranial variables (and arranged similarly for cranial indices, see below) with genera arranged from smallest to largest generic mean body weight. Expectedly, the overall small cranial dimensions of Miopithecus and the large cranial dimensions of Erythrocebus separate these genera from other cercopithecins. As such, the cercopithecins may grouped into small (Miopithecus; <3kg), medium (Allenopithecus, Cercopithecus and Chlorocebus; ~4-12kg) and large (Erythrocebus; >10kg; which is also the most sexually dimorphic of the tribe) genera based largely on body size. For example, many cranial (Verheyen, 1962; Shea, 1992), postcranial (Strasser, 1992) and head-&-body lengths (Napier & Napier, 1967; Napier, 1981; Rowe, 1996) also conform to this pattern. Some dimensions which exhibit this pattern may be seen in biasterionic breadth (biast), cranial vault length (g-o), superior facial and nasal heights (nas-pros and nas-ns) and palatal length A (ol-sta) (Figures 3.3, 3.9, 3.5, 3.6 and 3.7). However, there are some cranial variables which do not match these scaling trends. For example, despite its stocky and robust bauplan, the inferior breadth of the nasal bones (inbrnabo), parietal sagittal chord (br-lam), palatal breadth (bienm) and cranial height (bas- br) of Allenopithecus are smaller than expected for its body size (Figures 3.8, 3.9, 3.10 and 3.11). The rather narrow palate is likely the result of this species ‘molar flare’. Still, the sample for Allenopithecus is small and includes three subadults. Thus, results for this genus should be considered approximate and may be unrepresentative of its true variation. In addition, two cranial measurements which did illustrate differences between the Cercopithecus and Chlorocebus samples include the bizygomaxillare superior breadth (bizs) and the zygomatico-maxillary suture length (zs-zi) (Figures 3.12 and 3.13). To assess for statistically significant median differences, cranial variables were first converted into box plots. Cranial variables which distinguished at least two or more genera (i.e. where no overlap of boxes occurred giving a heuristic probability of <0.01) were then subjected to Shapiro-Wilk, Kruskal-Wallis & Mann-Whitney, and One-Way Analysis of

Box Plots for cranial variables of the Cercopithecini (Mi. – Miopithecus, n=19; Ch. – Chlorocebus, n=24; Cp. – Cercopithecus, n=24; Al. – Allenopithecus, n=6; & E. – Erythrocebus, n=16; variables in alphabetical orber by abbreviation):

Figure 3.2 E. 60 60 Figure 3.3 Ch. Cp. E.

50 Cp. Al. 50 Ch. Al. mm

Mi. mm Mi. 40 40

1 2 3 4 5 6 1 2 3 4 5 6 Cercopithecini: Cranial Height (bas-br) Cercopithecini: Biasterionic Breadth (biast)

28 Figure 3.4 27 E. 26 25 Cp Figure 3.5 24 . 80 23 E. 22 Ch. 21 70 Ch. Cp. 20 Al. mm 19 60 Al. 18

17 mm 16 50 Mi. 15 Mi. 14 13 40 12

1 2 3 4 5 6 1 2 3 4 5 6 Cercopithecini: Palatal Breadth (bienm) Cercopithecini: Biporionic Breadth (bipor)

Figure 3.7 70 Figure 3.6 Ch. 40 E. 60 Cp. E. Ch. Cp. Al. 50 Al. 30 Mi. mm mm 40 Mi. 20 30

1 2 3 4 5 6 1 2 3 4 5 6

Cercopithecini:C i h i iBizygomaxillare f i i Inferior ill Breadth d h (bi (bizi) i) Cercopithecini: Bizygomaxillare Superior Breadth (bizs)

Figure 3.8 Figure 3.9 50 110 E. E. Cp. 100 Ch. 40 90 Ch. Cp.

Al Al. Mi . mm 80 mm .

30 70 Mi.

1 2 3 4 5 6 1 2 3 4 5 6 Cercopithecini: Parietal Sagittal Chord (br-lam) Cercopithecini: Cranial Vault Length (g-o)

Figure 3.10 Figure 3.11 E. 14 18 13 17 E. 12 16 Ch. 11 Cp. 15 14 Cp. 10 13 Al. 9 Ch. 12

mm 8 11 mm 7 Al. 10 Mi. 9 Mi. 6 8 5 7 4 6 3 5 1 2 3 4 5 6 1 2 3 4 5 6 Cercopithecini: Inferior Breadth of the Nasal Bones (inbrnabo) Cercopithecini: Maximum Nasal Aperture Width (maxnawi)

Figure 3.12 Figure 3.13 70 60 E. E. 50 Cp 60 Ch. . Cp. Ch. Al. 40 Al. 50 mm mm Mi. 30 Mi. 40 20

1 2 3 4 5 6 1 2 3 4 5 6 Cercopithecini: Frontal Sagittal Chord (nas-br) Cercopithecini: Nasal Height (nas-ns)

Figure 3.14 Figure 3.15 70 60 E. E. 60 Cp. 50 Cp. Al. 50 Al. Ch. 40 Ch. 40 mm mm 30 Mi. 30 Mi. 20 20

1 2 3 4 5 6 1 2 3 4 5 6 Cercopithecini: Superior Facial Height (nas-pros) Cercopithecini: Palatal Length A (ol-sta)

Figure 3.16 30 E. Cp. Ch. Al. 20 Mi. mm

1 2 3 4 5 6 Cercopithecini: Zygomatico-maxillary Suture Length (zs-zi)

Variation (ANOVA). Shapiro-Wilk was utilized to assess the normality of the datasets. If a sample was found to be non-normal, the genus was excluded from One-Way ANOVA. Allenopithecus was excluded from all ANOVA analyses because of its small sample size (n=6), which fails to meet the ANOVA assumption of similar sample sizes. First, Kruskal- Wallis, which is essentially a non-parametric ANOVA (Hammer et al, 2006), was used to measure statistical significance between generic medians. Then, one-way ANOVA was employed to determine significant differences in generic means. Finally, these procedures were then repeated and applied to cranial indices (%).

3.2.1 Shapiro-Wilk results for cranial variables: Most cranial variables have a normal distribution. Table 3.7 lists the results of Shapiro-Wilk tests for normality. The palatal length (ol-sta) and the zygomatico-maxillary suture length (zs-zi) for Chlorocebus was not normally distributed. The nasal height (nas- ns) and biporionic breadth (bipor) for Erythrocebus was also not normally distributed. Thus, when applying a One-Way ANOVA to these cranial variables Chlorocebus and Erythrocebus were excluded, respectively.

3.2.2 Kruskal-Wallis and Mann-Whitney results for cranial variables: When Kruskal-Wallis was applied to the cercopithecin cranial variables, many statistically significant results emerged. Table 3.8 displays the results for Kruskal-Wallis. Allenopithecus was included because Kruskal-Wallis is non-parametric technique, the small sample size does not alter results and the median is a more stable measure of central tendency. The outcome of Kruskal-Wallis again highlights the similarity in cranial size between Allenopithecus, Cercopithecus and Chlorocebus. Cranial variables in which these genera are not significantly different included biasterionic breadth (biast; Table 3.8, no. 2), bizygomaxillare inferior and superior breadths (bizi and bizs; no. 5 & 6), cranial vault length (g-o; no. 8), maximum width of the nasal aperture (maxnawi; no. 10), frontal sagittal chord (nas-br; no. 11), nasal height (nas-ns) and superior facial height (nas-pros; no. 13). Once more, Miopithecus and Erythrocebus are clearly distinct. As the box-plot indicated, the median parietal sagittal chord (br-lam; no. 7) between Miopithecus and Allenopithecus is not significantly different.

3.2.3 One-Way ANOVA results for cranial variables: Akin to the results for Kruskal-Wallis, one-way ANOVA found significant results between genera for a range of variables. Table 3.9 displays the results of one-way ANOVA. Miopithecus and Erythrocebus are significantly different for each cranial variable examined. For most variables, the majority (>70%) of variation is between groups. Variables in which the majority of the variation is not explained between groups, but instead within groups, include biasterionic (biast) and bizygomaxillare superior (bizs) breadths, parietal sagittal chord (br-lam) and maximum width of the nasal aperture

Table 3.7: Shapiro-Wilk Results for cranial variables of the Cercopithecini; those in bold represent significant departures from normality (variables listed in alphabetical order by abbreviation).

Mi. Ch. Cp. Al. E. n=19 n=24 n=24 n=6 n=16 1. Bas-br W: 0.9 0.96 0.95 0.93 0.98 p(normal): 0.06 0.41 0.23 0.59 0.95 2. Biast W: 0.94 0.96 0.96 0.99 0.96 p(normal): 0.3 0.51 0.47 0.97 0.67 3. Bienm W: 0.96 0.99 0.96 0.97 0.96 p(normal): 0.57 0.99 0.45 0.91 0.69 4. Bipor W: 0.95 0.96 0.96 0.81 0.86 p(normal): 0.41 0.8 0.48 0.08 0.02 5. Bizi W: 0.98 0.94 0.95 0.87 0.92 p(normal): 0.93 0.14 0.31 0.23 0.18 6. Bizs W: 0.97 0.98 0.95 0.97 0.93 p(normal): 0.69 0.86 0.31 0.9 0.26 7. Br-lam W: 0.98 0.96 0.98 0.88 0.91 p(normal): 0.98 0.5 0.83 0.29 0.13

8. G-o W: 0.92 0.95 0.94 0.89 0.92 p(normal): 0.14 0.28 0.17 0.33 0.14 9. Inbrnabo W: 0.93 0.96 0.99 0.92 0.89 p(normal): 0.21 0.49 0.98 0.48 0.06 10. Maxnawi W: 0.96 0.95 0.94 0.88 0.9 p(normal): 0.58 0.21 0.18 0.27 0.07 11. Nas-br W: 0.96 0.97 0.98 0.91 0.91 p(normal): 0.66 0.65 0.96 0.43 0.11 12. Nas-ns W: 0.94 0.96 0.97 0.97 0.86 p(normal): 0.29 0.4 0.76 0.89 0.02 13. Nas-pros W: 0.92 0.95 0.98 0.97 0.9 p(normal): 0.12 0.3 0.89 0.91 0.07 14. Ol-sta W: 0.97 0.9 0.97 0.82 0.92 p(normal): 0.84 0.02 0.73 0.09 0.17

15. Zs-zi W: 0.94 0.88 0.94 0.87 0.96 p(normal): 0.24 0.009 0.17 0.21 0.71

Table 3.8: Kruskal-Wallis results for cranial variables of the Cercopithecini with Mann-Whitney pairwise comparisons (p (same)) (variables listed in alphabetical order by abbreviation).

Mi. Ch. Cp. Al. E. n=19 n=24 n=24 n=6 n=16 1. Bas-br 0 2.64E-08 3.04E-08 0.0003 5.26E-07 Mi. H: 64.88 0 0.77 0.01 0.00001 Ch. Hc: 64.94 0 0.004 2.88E-06 Cp. p(same): 2.73E-13 0 0.0005 Al 0 E. 2. Biast 0 4.42E-07 1.04E-07 0.003 5.73E-07 Mi. H: 48.2 0 0.89 0.31 0.005 Ch. Hc: 48.2 0 0.31 0.002 Cp. p(same): 8.59E-10 0 0.01 Al 0 E. 3. Bienm 0 2.64E-08 2.64E-08 0.0005 2.26E-07 Mi. H: 55.56 0 0.5 0.0004 0.002 Ch. Hc: 55.56 0 0.007 0.45 Cp. p(same): 2.48E-11 0 0.001 Al 0 E. 4. Bipor 0 3.04E-08 2.64E-08 0.0003 5.26E-07 Mi. H: 61.06 0 0.28 0.10 0.00006 Ch. Hc: 61.06 0 0.007 0.00008 Cp. p(same): 1.74E-12 0 0.005 Al 0 E. 5. Bizi 0 2.64E-08 2.64E-08 0.0003 5.26E-07 Mi. H: 60.66 0 0.58 0.86 5.23E-06 Ch. Hc: 60.67 0 0.59 0.00001 Cp. p(same): 2.10E-12 0.004 Al 0 E. 6. Bizs 0 1.21E-06 0.00004 0.07 0.06 Mi. H: 32.79 0 0.0024 0.010 0.004 Ch. Hc: 32.79 0 0.32 0.95 Cp. p(same): 1.32E-06 0 0.80 Al 0 E. 7. Br-lam 0 7.33E-07 6.47E-07 0.32 5.26E-07 Mi. H: 58.38 0 0.04 0.002 0.0002 Ch. Hc: 58.38 0 0.003 6.37E-06 Cp. p(same): 6.34E-12 0 0.0004 Al 0 E. 8. G-o 0 2.64E-08 2.64E-08 0.0003 5.26E-07 Mi. H: 67.6 0 0.40 0.11 1.95E-07 Ch. Hc: 67.61 0 0.31 1.45E-07 Cp. p(same): 7.29E-14 0 0.0005 Al 0 E.

Mi. Ch. Cp. Al. E. n=19 n=24 n=24 n=6 n=16 9. Inbrnabo 0 0.0009 2.64E-08 0.006 5.26E-07 Mi. H: 69.95 0 2.44E-08 0.28 1.45E-07 Ch.

Hc: 69.65 0 0.0002 0.003 Cp. p(same): 2.70E-14 0 0.0005 Al 0 E. 10. Maxnawi 0 2.64E-08 3.74E-08 0.001 5.26E-07 Mi. H: 57.69 0 .74 0.17 0.00004 Ch. Hc: 57.69 0 0.20 0.0001 Cp. p(same): 8.88E-12 0 0.003 Al 0 E. 11. Nas-br 0 5.29E-08 2.31E-07 0.0007 5.26E-07 Mi. H: 64.85 0 0.10 0.22 4.69E-07 Ch. Hc: 64.85 0 0.76 1.45E-07 Cp. p(same): 2.77E-13 0 0.0005 Al 0 E. 12. Nas-ns 0 2.64E-08 2.64E-08 0.0003 5.26E-07 Mi. H: 61.23 0 0.58 0.39 1.27E-06 Ch. Hc: 61.23 0 0.70 0.00002 Cp. p(same): 1.60E-12 0 0.007 Al 0 E. 13. Nas-pros 0 2.64E-08 2.64E-08 0.0003 5.26E-07 Mi. H: 61.97 0 0.56 0.20 7.21E-07 Ch. Hc: 61.97 0 0.55 0.00002 Cp. p(same): 1.12E-12 0 0.006 Al 0 E. 14. Ol-sta 0 2.64E-08 3.04E-08 3.00E-04 5.26E-07 Mi. H: 54.24 0 0.40 0.07 0.002 Ch. Hc: 54.24 0 0.04 0.0009 Cp. p(same): 4.70E-11 0 0.44 Al 0 E. 15. Zs-zi 0 1.21E-06 4.01E-08 0.0003 5.26E-07 Mi. H: 72.68 0 8.75E-07 0.01 1.24E-07 Ch. Hc: 72.68 0 0.86 6.25E-07 Cp. p(same): 6.17E-15 0 0.002 Al 0 E.

Table 3.9: One-Way ANOVA results for cranial variables of the Cercopithecini (variables listed in alphabetical order by abbreviation). Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 1. Bas-br p(same): 0 0.0002 0.0002 0.0002 Between F: 159.3 0 0.95 0.0002 groups: 2566.57 79.82 3 855.52 104.2 2.25E-27 0.09 df: 41.8 0 0.0002 Within groups: 648.86 20.18 79 8.21 p: 6.69E-23 ex. Al. 0 Total: 3215.43 82 2. Biast Mi. Ch. Cp. E. Between F: 53.12 0 0.0002 0.0002 0.0002 groups: 1110.49 58.16 3 370.16 36.61 6.19E-15 0.07 df: 40.86 0 1 0.0007 Within groups: 798.73 41.84 79 10.11 p: 3.69E-14 0 .0007 Total: 1909.22 82 ex. Al. 0 3. Bienm F: 139.2 Mi. Ch. Cp. E. Between df: 39.1 0 0.0002 0.0002 0.0002 groups: 707.66 70.86 3 235.89 64.03 4.27E-21 0.000004 p: 6.64E-21 0 0.74 0.0005 Within groups: 291.03 29.14 79 3.68 ex. Al. 0.01 Total: 998.69 82 0 4. Bipor Between Mi. Ch. Cp. groups: 2390.01 73.22 2 1195 87.5 4.89E-19 0.28 F: 116.5 0 0.0001 0.0001 Within groups: 874.055 26.78 64 13.66 df: 42.61 0 0.48 Total: 3264.06 66 p: 5.34E-18 ex. Al.& E 0 5. Bizi Mi. Ch. Cp. E. Between F: 100.2 0 0.0002 0.0002 0.0002 groups: 3292.89 74.50 3 1097.63 76.92 2.26E-23 0.001 df: 40.05 0 0.98 0.0002 Within groups: 1127.33 25.50 79 14.27 p: 1.17E-18 0 0.0002 Total: 4420.22 82 ex. Al. 0

Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 6. Bizs p(same): Mi. Ch. Cp. E. Between F: 23.76 0 0.0002 0.01 0.2 Mi. groups: 540.55 37.07 3 180.18 15.51 5.04E-08 0.0002 df: 39.13 0 0.01 0.0003 Ch. Within groups: 917.48 62.93 79 11.61 p: 6.36E-09 0 0.60 Cp. Total: 1458.03 82 ex. Al. 0 E. 7. Br-lam Mi. Ch. Cp. E. Between F: 64.13 0 0.0002 0.0002 0.0002 Mi. groups: 1150.87 63.63 3 383.62 46.07 2.56E-17 0.03 df: 40.27 0 0.28 0.0002 Ch. Within groups: 657.782 36.37 79 8.33 p: 2.16E-15 0 0.0002 Cp. Total: 1808.65 82 ex. Al. 0 E. 8. G-o Mi. Ch. Cp. E. Between F: 196.9 0 0.0002 0.0002 0.0002 Mi. groups: 10777.5 85.15 3 3592.49 151 1.28E-32 0.0001 df: 39.16 0 0.74 0.0002 Ch. Within groups: 1879.95 14.85 79 23.80 p: 1.18E-23 0 0.0002 Cp. Total: 12657.4 82 ex. Al. 0 E. 9. Inbrnabo Mi. Ch. Cp. E. Between F: 105.7 0 0.01 0.0002 0.0002 Mi. groups: 309.45 76.21 3 103.15 84.35 1.47E-24 0.005 df: 39.63 0 0.0002 0.0002 Ch. Within groups: 96.61 23.79 79 1.22 p: 5.98E-19 0 0.0003 Cp. Total: 406.06 82 ex. Al. 0 E. 10. Maxnawi Mi. Ch. Cp. E. Between F: 88.24 0 0.0002 0.0002 0.0002 Mi. groups: 322.75 65.96 3 107.59 51.02 1.92E-18 0.006 df: 41.17 0 0.99 0.0002 Ch. Within groups: 166.59 34.04 79 2.11 p: 5.72E-18 0 0.0002 Cp. Total: 489.34 82 ex. Al. 0 E.

Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 11. Nas-br p(same): Mi. Ch. Cp. E. Between F: 120 0 0.0002 0.0002 0.0002 Mi. groups: 2958.4 79.61 3 986.13 102.8 3.38E-27 0.50 df: 41.54 0 0.26 0.0002 Ch. Within groups: 757.716 20.39 79 9.59 p: 1.71E-20 0 0.0002 Cp. Total: 3716.11 82 ex. Al. 0 E. 12. Nas-ns Mi. Ch. Cp. Between F: 265.8 0 0.0001 0.0001 Mi. groups: 3317.92 78.43 2 1658.96 116.3 4.84E-22 0.0002 df: 36.04 0 0.72 Ch. Within groups: 912.654 21.57 64 14.26 p: 2.67E-22 ex. Al.& E 0 Cp. Total: 4230.58 66 13. Nas-pros Mi. Ch. Cp. E. Between F: 214.3 0 0.0002 0.0002 0.0002 Mi. groups: 7388.9 77.59 3 2462.97 91.17 1.39E-25 4.44E-07 df: 35.55 0 0.90 0.0002 Ch. Within groups: 2134.19 22.41 79 27.02 p: 8.18E-23 0 0.0002 Cp. Total: 9523.09 82 ex. Al. 0 E. 14. Ol-sta Mi. Cp. E. Between F: 109.1 0 0.0001 0.0001 Mi. groups: 4109.22 73.12 2 2054.61 76.18 1.05E-16 4.77E-06 df: 29.36 0 0.0001 Cp. Within groups: 1510.34 26.88 56 26.97 p: 2.56E-14 ex. Al. & Ch. 0 E. Total: 5619.56 58 15. Zs-zi Mi. Cp. E. Between F: 162 0 0.0001 0.0001 Mi. groups: 1003.88 82.22 2 501.94 129.5 9.92E-22 0.05 df: 31.82 0 0.0001 Cp. Within groups: 217.054 17.78 56 3.88 p: 2.08E-17 ex. Al. & Ch. 0 E. Total: 1220.94 58

(maxnawi). Only the bizygomaxillare superior breadth (bizs) and inferior breadth of the nasal bones (inbrnabo) were significantly different between Cercopithecus and Chlorocebus. Interestingly, some variables between these two genera are virtually identical in terms of mean and variance values; some of these include biasterionic (biast) and bizygomaxillare inferior (bizi) breadths and superior facial height (nas-pros).

3.2.4 Summary for cranial variables: In summary, cranial variables easily distinguish Miopithecus and Erythrocebus, which is not surprising when considering that talapoins and the patas monkey are the smallest and largest of the cercopithecins, occupy very different habitats and possess opposing social organizations. In contrast, Cercopithecus, Chlorocebus and Allenopithecus, which are sympatric in certain areas, overlap considerably in cranial dimensions despite occupying different ecological niches.

3.3 Cranial Indices: Many have discussed that proportions are perhaps a better measure of differences then raw data when studying evolutionary trends as they are size standardized comparisons (i.e. expressing shape via proportions; see Simpson et al, 1960; Schultz, A.H. 1963b; Ashton et al, 1965 & 1975; Zuckerman et al, 1973; Wood & Chamberlain, 1986; Albrecht et al, 1993). Table 3.10 provides the generic mean of several relative cranial proportions which are percentages, not fixed dimensions, with standard deviations (SD). Results from cranial indices expose some commonalities between genera, but also important proportional dissimilarities. Figures 3.17-3.27 are box-plots of the cranial indices for the Cercopithecini. Proportionally, Miopithecus has the largest neurocranium with very little facial projection and a relatively small face (Table 3.10: No. 1, 2, 37, 38, 51, 60 & 62). Allenopithecus is distinguished from all other cercopithecins by its relatively long nasal and zygomatic bones and palate (Table 3.10: No. 62, 63, 64, 66 & 71). Unlike other cercopithecins, the maxillo-alveolar breadth (biecm) of Erythrocebus is proportionally the least in relation to the intercanine distance (bicanin). In other words, the patas monkey, males in particular, has relatively large canines, associated with substantial increase in body size and facial elongation (Table 3.10: No. 21, 60 & 69).

Table 3.10: Generic means for cranial indices (%; indices listed in alphabetical order by abbreviation).

Mi. Ch. Cp. Al. E. 1. bas-br/bas-nas 92.84 84.54 83.45 80.49 78.26 SD 4.60 4.22 5.26 4.78 5.60 2. bas-br/bas-pros 84.07 69.71 68.66 63.44 63.60 SD 4.82 5.18 6.95 5.42 7.20 3. bas-br/biast 106.86 115.22 113.92 110.91 117.69 SD 4.57 7.48 6.76 7.76 8.28 4. bas-br/biaur 88.54 85.72 85.00 80.99 84.46 SD 4.75 4.22 5.51 3.84 4.66 5. bas-br/g-o 64.2 64.6 65.4 62.8 58.0 SD 1.9 2.9 3.7 2.1 3.0 6. bas-br/bizygo 79.71 75.17 74.54 71.57 73.11 SD 4.62 4.71 6.04 7.20 6.56 7. bas-br/pros-o 53.22 48.54 49.02 46.29 44.09 SD 2.01 2.84 3.69 3.10 3.63 8. bas-nas/bas-pros 90.56 82.41 82.12 78.74 81.06 SD 2.67 3.50 4.07 3.14 4.27 9. bas-pros/pros-o 63.40 69.72 71.64 73.27 69.60 SD 2.50 1.77 3.27 6.30 3.07 10. biast/biaur 83.02 74.55 74.67 73.21 71.96 SD 5.94 3.67 3.57 4.35 4.42 11. biast/g-o 60.2 56.3 57.5 56.8 49.5 SD 2.7 3.2 2.8 2.6 3.2 12. biast/lam-opn 153.0 144.5 150.4 133.6 133.1 SD 9.7 13.2 13.2 11.6 11.8 13. biaur/bizygo 90.03 87.71 87.72 88.23 86.52 SD 2.51 3.85 4.95 5.86 5.61 14. biaur/pros-o 60.19 56.63 57.73 57.15 52.16 SD 2.31 1.78 3.31 2.44 2.41 15. bicanex/bas-pros 40.53 35.95 37.63 35.94 36.75 SD 3.35 1.69 2.08 3.27 2.54 16. bicanex/nas-pros 83.83 67.03 69.29 63.96 63.43 SD 9.37 5.05 7.24 5.98 5.66 17. bicanex/ol-sta 91.54 74.19 81.09 71.42 77.72 SD 8.30 3.08 6.19 7.26 6.34 18. biecm/bas-br 60.99 64.82 67.48 74.88 65.51 SD 2.65 4.93 4.42 3.02 4.38 19. biecm/bas-nas 56.56 54.71 56.13 60.27 51.14 SD 2.33 3.70 2.28 4.07 3.28 20. biecm/biast 65.22 74.59 76.71 83.05 76.94 SD 4.69 6.55 4.53 6.84 5.48 21. biecm/bicanex 127.24 125.51 122.79 132.96 113.20 SD 13.39 9.79 9.65 15.94 10.79 22. biecm/bifmt 55.12 57.35 61.30 67.85 57.36 SD 2.33 3.86 3.19 6.15 3.42 23. biecm/bizi 70.86 73.43 75.00 79.41 68.93 SD 3.10 4.26 4.46 8.14 3.18 73

Mi. Ch. Cp. Al. E. 24. biecm/bizygo 48.57 48.64 50.14 53.68 47.67 SD 2.59 3.72 3.38 6.62 2.45 25. biecm/g-o 39.17 41.84 44.05 47.06 37.95 SD 1.59 2.73 2.71 2.75 2.03 26. biecm/ol-sta 115.71 93.00 99.44 89.01 88.01 SD 7.37 6.65 9.37 12.32 11.39 27. biecm/pros-o 32.43 31.41 32.95 34.71 28.78 SD 1.12 2.29 1.41 3.27 1.79 28. biecm/zs-zgyi 104.06 95.65 93.17 101.08 81.96 SD 9.36 6.22 9.18 14.28 6.80 29. bien/bas-br 69.09 70.15 72.95 77.35 74.17 SD 4.02 5.73 5.21 4.93 5.63 30. bien/bas-nas 64.05 59.16 60.70 60.47 57.90 SD 3.35 3.72 3.32 6.81 4.52 31. bien/bas-pros 57.95 48.73 49.88 47.62 46.97 SD 2.55 3.31 4.28 5.81 4.72 32. bien/biast 73.84 80.69 82.91 85.61 87.12 SD 5.43 6.87 5.14 6.39 7.36 33. bifmt/biaur 97.92 96.77 93.50 89.60 96.39 SD 4.09 3.23 6.07 3.04 4.75 34. bifmt/bizi 128.59 128.30 122.49 117.12 120.47 SD 4.17 7.37 6.62 6.94 7.75 35. bifmt/bizygo 88.15 84.83 81.84 79.01 83.30 SD 4.13 3.69 4.24 5.05 5.59 36. bifmt/g-o 71.12 73.01 71.95 69.59 66.27 SD 2.94 2.81 4.25 3.74 3.56 37. bifmt/nas-pros 192.05 146.74 138.61 125.08 124.67 SD 14.73 13.24 15.35 12.38 9.35 38. bifmt/pros-o 58.87 54.77 53.83 51.21 50.18 SD 1.79 1.67 2.22 2.73 1.20 39. bipor/g-o 67.4 69.5 72.7 70.8 64.4 SD 3.6 2.8 2.8 2.8 3.7 40. bizi/bas-br 86.18 88.39 90.16 94.90 95.26 SD 4.15 6.24 6.58 7.59 8.00 41. bizi/biast 92.09 101.75 102.45 104.84 111.84 SD 5.90 8.72 5.89 4.46 9.27 42. bizi/bizygo 68.58 66.28 66.93 67.50 69.24 SD 3.14 4.18 3.82 2.89 3.75 43. bizi/nas-pros 149.5 114.7 111.9 107.0 103.8 SD 12.6 12.0 10.2 10.6 9.2 44. bizs/zs-zgyi 111.17 97.52 82.35 81.75 62.78 SD 8.89 11.69 10.26 10.77 10.75 45. bizs/zs-zi 219.42 226.35 168.27 164.26 126.05 SD 23.41 36.81 25.74 23.62 26.76 46. bizs/bizi 75.82 74.76 66.24 64.29 52.69 SD 4.50 7.52 6.16 6.42 7.37

Mi. Ch. Cp. Al. E. 47. bizygo/pros-o 66.89 64.65 65.85 64.89 60.49 SD 3.00 2.81 2.66 2.94 4.13 48. br-lam/g-o 49.6 47.8 46.1 42.2 43.1 SD 2.4 3.6 3.0 2.4 2.6 49. g-o/biaur 137.88 132.62 129.99 128.89 145.56 SD 7.34 3.72 4.88 3.59 4.65 50. g-o/bizygo 124.11 116.28 114.00 113.81 126.03 SD 7.04 5.21 7.15 9.34 10.45 51. g-o/pros-o 82.84 75.07 74.99 73.67 75.89 SD 2.08 2.47 4.39 3.93 3.53 52. iv-pms/bicanex 54.47 64.21 60.42 53.20 64.41 SD 7.68 5.33 5.86 10.04 7.93 53. iv-pms/ol-sta 49.57 47.59 48.78 35.59 49.84 SD 5.71 3.90 3.83 6.19 5.64 54. lam-opn/bas-br 60.54 60.00 58.76 67.91 65.27 SD 3.98 5.28 2.87 4.00 5.43 55. lam-opn/bas-pros 50.87 41.90 40.38 43.04 41.47 SD 4.09 5.39 4.92 3.91 5.45 56. lam-opn/biaur 53.57 51.52 49.96 55.01 55.11 SD 4.04 6.01 4.31 4.20 5.19 57. lam-opn/g-o 39.5 39.2 38.4 42.7 37.4 SD 2.1 4.0 3.1 2.6 3.1 58. maxnawi/nas-rhi 76.8 63.0 61.3 45.6 62.6 SD 11.1 10.4 9.8 5.5 10.6 59. nas-br/g-o 63.5 61.9 60.9 62.3 59.3 SD 2.8 2.9 3.1 1.7 2.3 60. nas-pros/biaur 51.31 66.38 68.14 72.14 77.79 SD 4.96 5.58 7.84 6.41 7.79 61. nas-pros/bien 84.10 110.99 110.25 119.46 124.94 SD 7.81 10.61 11.58 14.12 14.59 62. nas-pros/g-o 37.18 50.07 52.53 56.00 53.49 SD 2.53 4.18 6.75 5.21 5.53 63. nas-rhi/nas-pros 38.72 42.63 43.91 50.73 41.55 SD 4.74 4.94 3.52 2.10 6.13 64. ol-sta/bas-br 52.98 69.89 68.45 85.45 75.75 SD 4.92 5.62 7.96 11.85 12.07 65. ol-sta/bas-nas 49.06 58.94 56.82 68.38 58.75 SD 3.69 3.64 4.46 6.87 6.19 66. ol-sta/bas-pros 44.39 48.48 46.53 53.87 47.43 SD 3.03 1.74 2.38 6.25 3.26 67. ol-sta/bizygo 42.11 52.36 50.71 61.00 54.92 SD 3.42 3.04 4.17 9.89 6.95 68. palhei/ol-sta 25.38 19.74 24.46 18.84 19.12 SD 2.55 2.60 2.83 2.69 2.71 69. zs-zgyi/bizi 68.52 76.95 80.95 79.06 84.53 SD 5.86 4.81 6.13 5.36 6.58

Mi. Ch. Cp. Al. E. 70. zs-zgyi/bizygo 46.88 50.87 54.03 53.25 58.49 SD 3.10 2.55 3.07 1.52 5.45 71. zs-zgyi/ol-sta 111.65 97.27 107.17 89.18 107.34 SD 7.46 3.81 10.04 14.27 10.03 72. zs-zi/bizi 34.78 33.52 39.81 39.34 42.50 SD 2.61 3.35 3.65 1.82 3.98 73. zs-zi/bizygo 23.85 22.24 26.64 26.56 29.46 SD 2.03 3.09 2.78 1.78 3.34 74. zs-zi/zs-zgyi 51.01 43.61 49.43 49.95 50.46

SD 4.81 4.12 5.74 4.11 5.08

Box Plots for cranial indices of the Cercopithecini (Mi. – Miopithecus, n=19; Ch. – Chlorocebus, n=24; Cp. – Cercopithecus, n=24; Al. – Allenopithecus, n=6; & E. – Erythrocebus, n=16; indices in alphabetical order by abbreviation):

Figure 3.17 70 200 Figure 3.18 Mi 190 . Cp. Cp. 180 Ch. Al. Mi. Ch. 60 170 E. E. 160 Al. %

% 150 140 50 130 120 110

1 2 3 4 5 6 1 2 3 4 5 6 Cercopithecini: biast/g-o x 100 Cercopithecini: biast/lam-opn x 100

Figure 3.19 Figure 3.20 80 Cp 130 Mi. Mi. . Ch. Al. 120 E. Ch. Al. 110 Cp. 70 100 % E. % 90 60 80 70

1 2 3 4 5 6 1 2 3 4 5 6 Cercopithecini: biecm/zs-zgyi x 100 Cercopithecini: bipor/g-o x 100

Figure 3.21 Figure 3.22 90 Mi. Mi. Ch. 130 80 Cp. 120 Ch. Al. 110 100 Cp. 70 E. Al. 90 % % E. 60 80 70 50 60 50 40 1 2 3 4 5 6 1 2 3 4 5 6 Cercopithecini: bizs/bizi x 100 Cercopithecini: bizs/zs-zgyi x 100

Figure 3.23 Figure 3.24 300 Ch. 100 Mi. Mi. Ch. 90 Cp. Cp. Al. 200 80 E. 70 E. % % 60 Al. 100 50 40

1 2 3 4 5 6 1 2 3 4 5 6 Cercopithecini: bizs/zs-zi x 100 Cercopithecini: maxnawi/nas-rhi x 100

Figure 3.25 Figure 3.26 E. 60 100 Al. Al. Ch. Cp. E. 90 50 Mi. Cp. 80 Ch. % 40 % 70 Mi. 60 30 50

1 2 3 4 5 6 1 2 3 4 5 6 Cercopithecini: nas-rhi/nas-pros x 100 Cercopithecini: ol-sta/bas-br x 100

Figure 3.27 130 Mi. Cp. E. 120

110 Al. Ch.

% 100

1 2 3 4 5 6 Cercopithecini: zs-zgyi/ol-sta x 100

The box-plot for bizygomaxillare superior breadth (bizs) illustrates a difference between Cercopithecus and Chlorocebus, so to do relative proportions. Proportionally, the superior portion of the maxilla in Chlorocebus is wider than Cercopithecus in relation to the zygomatico-maxillary suture length (zs-zi) and maximum length of the zygomatic bone (zs-zgyi); both of which also reveal interesting trends related to body size (Table 3.10; No. 44, 45 & 46).

3.3.1 Shapiro-Wilk results for cranial indices: Table 3.11 presents the results of applying Shapiro-Wilk to the selected cranial indices. All cranial indices may be assumed to posses a normal distribution. Therefore, it is fair to say that by converting the raw data from the cercopithecins into cranial indices, which are measures of proportional shapes not size, males and females are very similar despite sexual dimorphism.

3.3.2 Kruskal-Wallis and Mann-Whitney results for cranial indices: Table 3.12 provides the results of employing Kruskal-Wallis and Mann-Whitney to several cranial indices. Most indices for Miopithecus and Erythrocebus are significantly different from Chlorocebus, Cercopithecus and Allenopithecus. Although one index, the relative proportion of the maximum length of the zygomatic (zs-zgyi) to palatal length A (ol-sta) was not significantly different between the talapoin and patas monkeys (Table 3.12, No. 11). In contrast, there were several indices in which Chlorocebus, Cercopithecus and Allenopithecus were not significantly different (Table 3.12, no. 1, 2, 4, 8, 9 and 10).

3.3.3 One-Way ANOVA results for cranial indices: Table 3.13 provides the results from subjecting the cranial indices to one-way ANOVA. Again, Allenopithecus was excluded from all ANOVA analyses due to small sample size. The results for one-way ANOVA of the cercopithecins confirm the results of Kruskal-Wallis and Mann-Whitney. Excluding three indices, Erythrocebus was significantly different to all other genera. The indices that were not significantly different include the relative proportion of the maximum width of the nasal aperture (maxnawi) to the sagittal length of the nasal bones (nas-rhi; Table 3.13, no. 8); the relative contribution

Table 3.11: Shapiro-Wilk Results for cranial indices of the Cercopithecini; those in bold represent significant departures from normality (indices (%) listed in alphabetical order by abbreviation).

Mi. Ch. Cp. Al. E. n=19 n=24 n=24 n=6 n=16 1. Biast/g-o W: 0.96 0.96 0.96 0.94 0.95 p(normal): 0.55 0.48 0.44 0.65 0.44 2. Biast/lam-opn W: 0.96 0.99 0.95 0.99 0.92 p(normal): 0.51 0.98 0.26 0.99 0.15 3. Biecm/zs-zgyi W: 0.92 0.95 0.98 0.96 0.97 p(normal): 0.10 0.24 0.94 0.83 0.90 4. Bipor/g-o W: 0.99 0.97 0.97 0.91 0.97 p(normal): 0.99 0.74 0.56 0.42 0.89 5. Bizs/bizi W: 0.96 0.98 0.97 0.92 0.95 p(normal): 0.58 0.88 0.63 0.53 0.46 6. Bizs/zs-zgyi W: 0.97 0.96 0.96 0.92 0.96 p(normal): 0.79 0.54 0.51 0.52 0.66 7. Bizs/zs-zi W: 0.96 0.98 0.96 0.96 0.96 p(normal): 0.5 0.93 0.50 0.82 0.69

8. Maxnawi/nas-rhi W: 0.96 0.97 0.97 0.98 0.19 p(normal): 0.66 0.76 0.64 0.97 0.12 9. Nas-rhi/nas-pros W: 0.94 0.97 0.97 0.90 0.96 p(normal): 0.29 0.70 0.59 0.35 0.72 10. Ol-sta/bas-br W: 0.96 0.98 0.97 0.86 0.94 p(normal): 0.63 0.95 0.73 0.18 0.40 11. Zs-zgyi/ol-sta W: 0.97 0.94 0.98 0.98 0.98 p(normal): 0.83 0.19 0.96 0.94 0.92

Table 3.12: Kruskal-Wallis results for cranial indices of the Cercopithecini with Mann-Whitney pairwise comparisons (p (same)) (indices (%) listed in alphabetical order by abbreviation).

Mi. Ch. Cp. Al. E. n=19 n=24 n=24 n=6 n=16 1. Biast/g-o 0 0.0006 0.004 0.02 6.24E-07 Mi. H: 45.53 0 0.17 0.17 3.76E-06 Ch. Hc: 45.54 0 0.38 4.06E-07 Cp. p(same): 3.08E-09 0 0.0008 Al 0 E. 2. Biast/lam-opn 0 0.02 0.46 0.003 0.00006 Mi. H: 26.34 0 0.19 0.08 0.006 Ch. Hc: 26.34 0 0.009 0.001 Cp. p(same): 0.00003 0 0.77 Al 0 E. 3. Biecm/zs-zgyi 0 0.006 0.00002 0.82 7.41E-07 Mi. H: 51.33 0 0.006 0.01 1.27E-06 Ch. Hc: 51.33 0 0.0007 0.0001 Cp. p(same): 1.91E-10 0 0.0005 Al 0 E. 4. Bipor/g-o 0 0.02 0.00003 0.03 0.04 Mi. H: 42.48 0 0.0001 0.21 0.0001 Ch. Hc: 42.49 0 0.12 7.74E-07 Cp. p(same): 1.33E-08 0 0.003 Al 0 E. 5. Bizs/bizi 0 0.50 3.41E-07 0.0008 5.26E-07 Mi. H: 64.78 0 7.09E-07 0.001 1.24E-07 Ch. Hc: 64.78 0 0.34 9.56E-07 Cp. p(same): 2.86E-13 0 0.006 Al 0 E. 6. Bizs/zs-zgyi 0 0.0003 2.64E-08 0.0003 5.26E-07 Mi. H: 70.81 0 4.15E-07 0.001 1.24E-07 Ch. Hc: 70.81 0 0.82 6.37E-06 Cp. p(same): 1.53E-14 0 0.001 Al 0 E. 7. Bizs/zs-zi 0 0.13 2.22E-06 0.0008 5.26E-07 Mi. H: 64.89 0 2.17E-08 0.0003 1.24E-07 Ch. Hc: 64.89 0 0.62 0.0001 Cp. p(same): 2.72E-13 0 0.02 Al 0 E.

Mi. Ch. Cp. Al. E. n=19 n=24 n=24 n=6 n=16 8. Maxnawi/nas-rhi 0 0.0005 0.00009 0.0003 0.00002 Mi. H: 34.48 0 0.55 0.001 0.18 Ch. Hc: 34.48 0 0.001 0.69 Cp. p(same): 5.95E-07 0 0.001 Al 0 E. 9. Nas-rhi/Nas-pros 0 0.02 0.0007 0.0003 0.007 Mi. H: 25.57 0 0.43 0.001 0.63 Ch. Hc: 25.57 0 0.0005 0.75 Cp. p(same): 0.00004 0 0.002 Al 0 E. 10. Ol-sta/bas-br 0 2.64E-08 6.50E-08 0.0003 5.26E-07 Mi. H: 51.40 0 0.38 0.007 0.12 Ch. Hc: 51.40 0 0.003 0.03 Cp. p(same): 1.84E-10 0 0.15 Al 0 E. 11. Zs-zgyi/ol-sta 0 4.01E-08 0.03 0.0004 0.44 Mi. H: 47.97 0 0.00002 0.34 8.31E-07 Ch. Hc: 47.98 0 0.009 0.16 Cp. p(same): 9.55E-10 0 0.001 Al 0 E.

Table 3.13: One-Way ANOVA results for cranial indices of the Cercopithecini (indices (%) listed in alphabetical order by abbreviation). Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 1. Biast/g-o p(same): Mi. Ch. Cp. E. Between F: 37.61 0 0.0005 0.02 0.0002 groups: 1068.02 60.51 3 356.00 40.34 6.49E-16 0.65 df: 41.30 0 0.56 0.0002 Within groups: 697.13 39.49 79 8.82 p: 0.00 ex. Al. 0 0.0002 Total: 1765.15 82 0 2. Biast/lam-opn Mi. Ch. Cp. E. Between F: 10.29 0 0.13 0.90 0.0002 groups: 4110.60 25.84 3 1370.20 9.18 0.00003 0.31 df: 42.15 0 0.43 0.02 Within groups: 11796.70 74.16 79 149.33 p: 3.30E-05 ex. Al. 0 0.0003 Total: 15907.30 82 0 3. Biecm/zs-zgyi Mi. Ch. Cp. E. Between F: 25.71 0 0.002 0.0002 0.0002 groups: 4515.81 55.85 3 1505.27 33.31 5.10E-14 0.01 df: 39.26 0 0.12 0.0002 Within groups: 3569.93 44.15 79 54.19 p: 2.29E-09 ex. Al. 0 0.0003 Total: 8085.74 82 0 4. Bipor/g-o Mi. Ch. Cp. E. Between F: 21.97 0 0.23 0.0001 0.01 groups: 710.38 48.39 3 236.79 24.69 2.27E-11 0.29 df: 39.34 0 0.005 0.0002 Within groups: 757.65 51.61 79 9.59 p: 1.60E-08 0 0.0002 Total: 1468.03 82 ex. Al. 0 5. Bizs/bizi Mi. Ch. Cp. E. Between F: 73.62 0 0.92 0.0002 0.0002 groups: 6294.31 78.08 3 2098.1 93.82 5.79E-26 0.20 df: 40.67 0 0.0002 0.0002 Within groups: 1766.61 21.92 79 22.36 p: 1.75E-15 0 0.0002 Total: 8060.92 82 ex. Al. 0

Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 6. Bizs/zs-zgyi p(same): Mi. Ch. Cp. E. Between F: 25.71 0 0.002 0.0002 0.0002 Mi. groups: 4515.81 55.85 3 1505.27 33.31 5.10E-14 0.01 df: 39.26 0 0.12 0.0002 Ch. Within groups: 3569.93 44.15 79 45.19 p: 2.29E-09 0 0.0003 Cp. Total: 8085.74 82 ex. Al. 0 E. 7. Bizs/zs-zi Mi. Ch. Cp. E. Between F: 67.69 0 0.34 0.0002 0.0002 Mi. groups: 136916.00 73.41 3 45638.70 72.71 1.16E-22 0.93 df: 41.34 0 0.0002 0.0002 Ch. Within groups: 49590.00 26.59 79 627.72 p: 0.00 0 0.0002 Cp. Total: 186506.00 82 ex. Al. 0 E. 8. Maxnawi/nas-rhi Mi. Ch. Cp. E. Between F: 12.06 0 0.0002 0.0002 0.0001 Mi. groups: 3597.14 32.82 3 1199.05 12.86 6.31E-07 0.11 df: 42.98 0 0.94 0.63 Ch. Within groups: 7363.9 67.18 79 93.21 p: 7.38E-06 0 0.92 Cp. Total: 10961 82 ex. Al. 0 E. 9. Nas-rhi/nas-pros Mi. Ch. Cp. E. Between F: 5.30 0 0.03 0.002 0.009 Mi. groups: 321.88 17.31 3 107.29 5.51 0.002 0.50 df: 40.61 0 0.79 0.98 Ch. Within groups: 1537.71 82.69 79 19.46 p: 0.004 0 0.96 Cp. Total: 1859.58 82 ex. Al. 0 E.

Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 10. Ol-sta/bas-br p(same): Mi. Ch. Cp. E. Between F: 67.49 0 0.0002 0.0002 0.0002 Mi. groups: 5317.64 57.94 3 1772.55 36.27 7.67E-15 3.24E-06 df: 39.08 0 0.90 0.02 Ch. Within groups: 3860.78 42.06 79 48.87 p: 1.63E-15 ex. Al. 0 0.003 Cp. Total: 9178.41 82 0 E. 11. Zs-zgyi/ol-sta Mi. Ch. Cp. E. Between F: 58.8 0 0.0002 0.08 0.94 Mi. groups: 3390.67 47.27 3 1130.22 23.61 5.24E-11 0.0001 df: 36.19 0 0.0002 0.0002 Ch. Within groups: 3782.51 52.73 79 47.88 p: 5.47E-14 ex. Al. 0 0.26 Cp. Total: 7173.18 82 0 E.

of the sagittal length of the nasal bones (nas-rhi) to the superior facial height (nas-pros; no. 9); and lastly, the relative proportion of the zygomatic bone’s length (zs-zgyi) to palatal length (ol-sta; no. 11); the latter index was also not significantly different between Erythrocebus and Cercopithecus. Three indices revealed proportional similarities between Miopithecus and Chlorocebus and were not significantly different (Table 3.13, no. 2, 4 & 5). In addition, Cercopithecus and Chlorocebus were not significantly different in regard to the relative contribution of the occipital bone’s breadth (biast) and neurocranial lengths (g-o and lam- opn; Table 3.13, no. 1 & 2). Other proportions which were not significantly different between these two genera included the maximum width of the nasal aperture (maxnawi) in relation to the length of the nasal bones (nas-rhi; Table 3.13, no. 8); the relative difference between the sagittal length of the nasal bones (nas-rhi) and superior facial height (nas-pros; Table 3.13, no. 9); and finally, the relative length of the palate (ol-sta) in relation to cranial height (bas-br; Table 3.13, no. 10).

3.3.4 Summary for cranial indices: In summary, like the univariate data, Miopithecus and Erythrocebus are once again very distinct from the other members of the tribe. In addition, excluding some the cranial indices discussed above, Allenopithecus, Cercopithecus and Chlorocebus overlap in many cranial proportions, although some indices reveal significance differences. Cranial indices also assist in reducing the differences in size between males and females.

3.4 Bivariate Results: To examine the relationship between cranial variables and body size the Ln transformed generic mean for particular variables was plotted against Ln transformed mean generic body weights (g). Table 3.14 presents the results of linear regression between cranial variables and body weight. Save for the bizygomaxillare superior breadth (bizs) all other cranial variable are significantly correlated to body size. In fact, the bivariate plots graphically illustrate how each genus is distributed across the potential or real landscape, one with unevenly distributed conspecifics and interspecific competitors, predators and food resources (Figure 3.28). For example, because of its small size, Miopithecus can pass

Table 3.14: Results of body weight (Ln (g)) plotted against the generic mean for particular cranial measurements (Ln (mm)) (cranial variables listed in alphabetical order by abbreviation; those in bold represent cranial variables with an adjusted R squared value >0.70 and a p-value <0.05).

Adjusted P-value/ Mean Generic Body Weight (g) Ln (mm) R squared Significance F (Napier, 1981; Rowe, 1996; bas-br 0.84 0.02 Fleagle, 1998; Delson et al, 2000): bas-nas 0.93 0.01 bas-pros 0.98 0.001 Miopithecus spp. - 1266 biast 0.88 0.01 Chlorocebus spp. - 4381 biaur 0.96 0.002 Cercopithecus spp. - 4676 Allenopithecus nigroviridis - 4750 bicanex 0.93 0.005 Erythrocebus patas - 6899 biecm 0.98 0.001 bien 0.93 0.005 bienm 0.80 0.03 bifmo 0.91 0.01 bifmt 0.80 0.03 bipor 0.93 0.005 bizi 0.91 0.01 bizs 0.10 0.32 bizygo 0.96 0.0002 br-lam 0.37 0.17 g-o 0.73 0.04 lam-opn 0.84 0.02 nas-br 0.73 0.04 nas-pros 0.98 0.001 nas-rhi 0.94 0.004 ol-sta 0.95 0.003 ol-pms 0.90 0.01 palhei 0.88 0.01 pros-o 0.90 0.01 zs-zgyi 0.94 0.004 zs-zi 0.78 0.03

4.40 Figure 3.28: Adjusted R2: 0.96 E. 4.30 Significance F & P-value 0.002 4.20 Ch. y = 0.2495x + 2.0815 Cp. Al 4.10 .

4.00

3.90

Ln Bizygomatic Breadth Bizygomatic Ln Mi. 3.80 6.50 7.00 7.50 8.00 8.50 9.00 Ln Body Weight (g)

and climb through thick vegetation, including dense Mangrove swamps and undergrowth, which they are known to utilize (Gautier-Horn, 1973; Wolfheim, 1983), and which would otherwise hinder or compromise another genus, particularly say Erythrocebus, which is very large (intermembral index of 92 - a high percentage much like terrestrial papionins) and has a long slender body. In contrast, Allenopithecus, Cercopithecus and Chlorocebus are closely grouped together around the linear trend line (Figure 3.28). Allenopithecus and Cercopithecus spp., particularly C. neglectus, do share ecological niches, both preferring inundated, swamp forests (McGraw, 1994). However, the majority of the geographic range for Chlorocebus far exceeds that of Cercopithecus spp. and only marginally overlaps and ecological niche separation limits competition (Tappen, 1960; Dunbar & Dunbar, 1974; Wolfheim, 1983; Isabell et al, 1998). Despite this, Chlorocebus is restricted to areas near permanent water sources (i.e. lakes or rivers). Only one figure is provided because despite which cranial variable is plotted against body weight, the pattern and distribution of genera remains very similar.

3.5 Multivariate Analyses and Morphological Distances: To consider and evaluate the entire multivariate datasets simultaneously Principal Components Analysis (PCA) and Canonical Variates Analysis (CVA) were utilized.

3.5.1 PCA results for Ln transformed data: To identify the underlying structure (i.e. the cranial variables which account for the most of the variation between genera) of the Cercopithecini multivariate dataset derived from Ln transformed generic means, PCA was employed. By doing so, PCA based on the variance-covariance matrix results in the first (Eigenvalue – 1.53, 93.76%), second (Eigenvalue – 0.06, 3.56%) and third (Eigenvalue – 0.03, 1.85%) PCs accounting for 99.17% of the variation within the tribe (Figure 3.29). One hundred percent of the variation is explained by four PCs. The low to moderate variable loadings (low - <0.20; moderate - >20 <0.80; high > 0.80) due to data transformation (i.e. Ln transformed) and homogeneity of dataset; see Tabachnick & Fidell, 2007) for the first PC are all positive and may be interpreted as differences due to size (Jolicoeur & Mosimann, 1960; Rao, 1964). Evidence

to substantiate this can be seen in the PCA object scores per genus. Miopithecus, the smallest Old World monkey, scores the lowest with 16.86 (succeeded by Allenopithecus at 18.95) and Erythrocebus, the largest cercopithecin, scores the highest with 20.27 (followed by Cercopithecus at 19.17). The PCA score for Chlorocebus is 18.97. The low to moderate variable loadings for PCs 2 and 3 have mixed values. The first PC is dominated by naso-facial and palatal lengths. The largest variable loadings are achieved by the sagittal length of the nasal bones (nas-rhi), nasal height (nas- ns), superior facial height (nas-pros) and the sagittal height of the nasal aperture (rhi-ns) with 0.28, 0.26, 0.25 and 0.23, respectively (Table 3.17). These dimensions were closely followed by palatal measurements, such as palatal lengths B and A, and the length between incisivion and the junction of the palato-maxillary suture (iv-pms) scoring 0.22, 0.22 and 0.21, correspondingly. Other high variable loadings go to the maximum length of the zygomatic (zs-zgyi) and inferior breadth of the nasal bones (inbrnabo), both with 0.20. The two largest positively loaded measurements for the second PC are, the sagittal length of the nasal bones (nas-rhi) and palatal length A (ol-sta) with 0.54 and 0.25. Contrasting with these dimensions are negatively loaded measurements such as the inferior breadth of the nasal bones (inbrnabo), sagittal height of the nasal aperture (rhi-ns) and parietal sagittal chord (br-lam) receiving -0.51, -0.29 and -0.27, respectively. The largest variable loadings for the third PC are achieved by the inferior breadth of the nasal bones (inbrnabo) with 0.49 and followed by the zygomatico-maxillary suture length (zs-zi) with 0.39. Other relatively high positive variable loadings go to palatal height at the second upper molar (palhei), internal bicanine (bicanin) and biseptal breadths with 0.26, 0.24 and 0.20, correspondingly. Juxtaposed against these measurements are negatively loaded dimensions such as bizygomaxillare superior breadth (bizs), palatal length B (ol-pms) and the sagittal height of the nasal aperture (rhi-ns) which procure variable loadings of -0.37, -0.25 and -0.22, respectively. Table 3.15 provides the inter-generic Euclidean distance matrix produced from PCA object scores per genus from the first to fifth principal components based on Ln transformed generic means (which are equivalent to Mahalanobis distances; see Elmore & Richman, 2001; and Mimmack et al, 2001). The average distance between cercopithecin genera is 1.57 (Range 2.89). The largest distance produced is between Miopithecus and

Figure 3.29: Cercopithecini scatterplot of PC 1 & 2 and PC 2 & 3 for Ln transformed generic means. Al. - Allenopithecus; Mi. - Miopithecus; Cp. - Cercopithecus; Ch. - Chlorocebus; and E. - Erythrocebus. Nasal & Palatal -1 -3.4 Inferior breadth of the lengths nasal bones & zs-zi -1.1 Cp.

-1.2 -3.5 Al. Al. -1.3 E. Mi. -3.6 PC 2: -1.4 PC 3: Increase in 3.56% -1.5 1.85% body size -3.7 -1.6 Ch. -1.7 -3.8 Cp. -1.8 Mi. E. -1.9 Nasal & Facial lengths Ch. Nasal & Palatal lengths

17 18 19 20 -1.9 -1.8 -1.7 -1.6 -1.5 -1.4 -1.3 -1.2 -1.1 -1 Inferior breadth of PC 1: 93.76% PC 2: 3.56% the nasal bones & Bizygomaxillare height of the nasal Superior aperture

% of PCA PC Eigenvalue Var. Cum. %. scores Axis 1 Axis 2 Axis 3 1 1.53 93.76 93.76 Al. 18.95 -1.23 -3.51 2 0.06 3.56 97.33 Mi. 16.86 -1.77 -3.56 3 0.03 1.85 99.17 Cp. 19.17 -1.74 -3.45 Ch. 18.97 -1.61 -3.90 E. 20.27 -1.83 -3.56 Table 3.15: Inter-generic Euclidean distances based on PCA Mean 18.84 -1.64 -3.60 scores from PC 1-5 per genus from Ln transformed generic Max. 20.27 -1.83 -3.90 means. Min. 16.86 -1.23 -3.45 Al. 0 Mean 1.57 Range 3.41 0.60 0.44 Mi. 2.16 0 Range 2.89 SD 1.23 0.24 0.17 Cp. 0.60 2.33 0 CV 6.55 14.68 4.82 Ch. 0.55 2.15 0.52 0 E. 1.45 3.41 1.15 1.37 0 Al. Mi. Cp. Ch. E. Table 3.16: Intra-generic Euclidean distances based on Ln transformed data. Mean Max Min Range Al. 0.90 1.32 0.51 0.81 Mi. 0.71 1.53 0.35 1.18 Cp. 0.91 2.00 0.42 1.58 Ch. 0.83 1.52 0.30 1.22 E. 1.06 1.73 0.43 1.30

Table 3.17: Variable loadings for Cercopithecini Ln transformed generic means. PC loadings PC 1 PC 2 PC 3 bizs 0.04 inbrnabo -0.51 bizs -0.37 biast 0.08 rhi-ns -0.29 ol-pms -0.25 br-lam 0.09 br-lam -0.27 rhi-ns -0.22 bas-br 0.11 bienm -0.17 br-lam -0.18 nas-br 0.11 iv-pms -0.14 iv-pms -0.17 bifmo 0.11 g-o -0.13 bifmt -0.15 bifmt 0.11 bifmt -0.11 bienm -0.13 biaur 0.12 bas-br -0.09 bas-br -0.11 lam-opn 0.12 nas-br -0.09 nas-br -0.09 bipor 0.12 bicanin -0.08 ol-sta -0.08 bien 0.13 ol-pms -0.07 pros-o -0.08 g-o 0.13 biast -0.05 maxnawi -0.07 biecm 0.13 bipor -0.04 g-o -0.07 bizi 0.13 bifmo -0.03 bas-pros -0.06 bizygo 0.13 zs-zi -0.03 bizygo -0.06 palhei 0.14 pros-o -0.03 bifmo -0.06 bienm 0.15 bien -0.02 biaur -0.06 bicanin 0.15 bas-nas -0.02 bas-nas -0.06 biseptal 0.15 biseptal -0.02 biast -0.05 bas-nas 0.16 bizi -0.02 maxalvlen -0.03 pros-o 0.16 bicanex -0.02 bipor -0.03 bicanex 0.16 maxnawi -0.01 lam-opn -0.02 i1i2 0.18 zs-zygi 0.00 bizi -0.01 maxnawi 0.18 i1i2 0.00 nas-ns -0.01 zs-zi 0.19 bizygo 0.01 bien 0.01 bas-pros 0.19 biaur 0.01 biecm 0.01 maxalvlen 0.19 bas-pros 0.02 nas-pros 0.02 inbrnabo 0.20 bizs 0.05 i1i2 0.06 zs-zygi 0.20 lam-opn 0.06 bicanex 0.06 iv-pms 0.21 palhei 0.10 zs-zygi 0.07 ol-sta 0.22 maxalvlen 0.11 nas-rhi 0.14 ol-pms 0.22 biecm 0.11 biseptal 0.20 rhi-ns 0.23 nas-pros 0.17 bicanin 0.24 nas-pros 0.25 nas-ns 0.19 palhei 0.26 nas-ns 0.26 ol-sta 0.25 zs-zi 0.39 nas-rhi 0.28 nas-rhi 0.54 inbrnabo 0.49

Erythrocebus at 3.41 while the smallest distance is between Cercopithecus and Chlorocebus with 0.52 succeeded by Cercopithecus and Allenopithecus, 0.60. For comparison, Table 3.16 provides intra-generic Euclidean distances based on Ln transformed data. The largest mean intra-generic Euclidean distance was produced by Erythrocebus with 1.06 (followed by Cercopithecus and Allenopithecus with 0.91 and 0.90). This is due to the sexual dimorphism between males and females. The smallest mean intra-generic Euclidean distance was produced by Miopithecus with 0.71 (succeeded by Chlorocebus, 0.83). Male and female talapoins are not extremely sexually dimorphic (Rowe, 1996) and smaller species have smaller ranges of variation (Albrecht, 1978). In summary, for Ln transformed data the major distinguishing features between genera of the Cercopithecini are the dimensions of the naso-facial region. These results are similar to those reported and discussed by Verheyen (1962). In particular, the sagittal length of the nasal bones (nas-rhi), inferior breadth of the nasal bones (inbrnabo), nasal height (nas-ns) and palatal length A (ol-sta) received some of the highest variable loadings (Table 3.17). In contrast, neurocranial measurements did not figure prominently in PCA for Ln transformed data.

3.5.2 PCA results for MSV Mosimann Shape Variables (MSV): To identify the underlying structure (i.e. the cranial variables which account for the most of the variation between genera) of the Cercopithecini multivariate dataset derived from the generic mean Mosimann Shape Variables (MSV; Mosimann, 1970; Mosimann & James, 1979; Darroch & Mosimann, 1985), PCA was employed. By doing so, PCA based on the variance-covariance results in the first (Eigenvalue – 0.22, 78.35%) and second (Eigenvalue – 0.03, 11.32%) PCs accounting for 89.67% of the variation within the tribe (7.65% less variation explained than PCs 1 & 2 for Ln transformed generic means but 84.26% more variation explained than PCs 2 & 3 for Ln transformed generic means; Figure 3.30). One hundred percent of the variation is explained by four PCs. The variable loadings for the first PC are positive and negative (Tables 3.18.), which again vary from low to moderate, and thus cannot be interpreted as size differences but instead shape dissimilarities; the second PC also has mixed values. PCA object scores per genus reveal this as well. The largest PCA score is achieved by Miopithecus with 3.44 (followed by

Figure 3.30: Cercopithecini scatterplot of PC 1 & 2 for generic mean MSV. Al. - Allenopithecus; Mi. - Miopithecus; Cp. - Cercopithecus; Ch. - Chlorocebus; and E. - Erythrocebus.

Table 3.18: Variable loadings for 0.8 E. Cercopithecini generic mean MSV. PC loadings Cranial vault 0.7 PC 1 PC 2 & Nasal nas-pros -0.30 ol-sta -0.37 lengths nas-ns -0.30 bizs -0.34 0.6 ol-sta -0.22 biecm -0.31

Mi. nas-rhi -0.20 nas-rhi -0.28 PC 2: Ch. bas-pros -0.20 biaur -0.20 0.5 11.32% Cp. maxalvlen -0.14 bizygo -0.16 zs-zygi -0.11 biast -0.16 Biasterionic & ol-pms -0.10 bipor -0.11 0.4 Bizygomaxillare rhi-ns -0.07 bifmo -0.09 Facial & Nasal lengths Superior breadths iv-pms -0.05 palhei -0.06 0.3 zs-zi -0.05 lam-opn -0.06 Al. maxnawi -0.03 bien -0.05 i1i2 -0.02 bizi -0.04 Palatal length & 3 4 inbrnabo -0.01 bas-br -0.03 Bizygomaxillare PC 1: 78.35% bicanex -0.01 nas-ns -0.03 Superior breadth palhei 0.00 maxalvlen -0.02 % of Cum. biseptal 0.01 maxnawi -0.02 bicanin 0.01 nas-pros -0.01 PC Eigenvalue Var. %. 78.35 pros-o 0.02 biseptal 0.01 1 0.22 78.35 bienm 0.03 i1i2 0.02 2 0.03 11.32 89.67 3 0.02 5.73 95.40 biecm 0.03 bicanin 0.02 bas-nas 0.04 bicanex 0.02 lam-opn 0.08 bienm 0.07 PCA scores Axis 1 Axis 2 bien 0.09 bifmt 0.07 Al. 2.35 0.29 bizi 0.10 zs-zi 0.08 Mi. 3.44 0.55 bizygo 0.13 bas-pros 0.09 Cp. 2.52 0.50 bipor 0.16 nas-br 0.09 Ch. 2.70 0.52 biaur 0.17 inbrnabo 0.11 E. 2.29 0.79 bifmo 0.20 zs-zygi 0.11 Mean 2.66 0.53 nas-br 0.23 iv-pms 0.13 Max. 3.44 0.79 bas-br 0.24 bas-nas 0.14 Table 3.19: Inter-generic Min. 2.29 0.29 bifmt 0.25 ol-pms 0.18 Euclidean distances based Range 1.15 0.50 g-o 0.26 br-lam 0.21 on PCA scores from PC SD 0.47 0.18 br-lam 0.28 pros-o 0.23 1-5 per genus from CV 17.50 33.41 generic mean MSV. bizs 0.30 rhi-ns 0.31 Al. 0 Mean 0.69 biast 0.31 g-o 0.37 Mi. 1.12 0 Range 0.8 Table 3.20: Mean Max Min Range Cp. 0.42 0.96 0 Intra-generic Al. 0.70 0.97 0.42 0.55 Ch. 0.50 0.78 0.37 0 Euclidean Mi. 0.51 0.83 0.29 0.54 E. 0.50 1.18 0.47 0.55 0 distances Cp. 0.65 1.28 0.36 0.92 Al. Mi. Cp. Ch. E. based on Ch. 0.56 1.14 0.24 0.91 MSV. E. 0.64 1.11 0.32 0.79

Chlorocebus at 2.7) but the smallest PCA score goes to Erythrocebus with 2.29 (succeeded by Allenopithecus at 2.35). The first PC (Table 3.18) is an interesting mix of breadth and length MSV of the neuro- and viscerocranium. Interestingly, the neurocranium MSV are positive while those of the viscerocranium are negative. Positively loaded MSV include biasterionic and bizygomaxillare superior breadths (biast and bizs), parietal sagittal chord (br-lam), cranial vault length (g-o), superior facial breadth (bifmt), cranial height (bas-br) and frontal sagittal chord (nas-br), which attain variable loadings of 0.31, 0.30, 0.28, 0.26, 0.25, 0.24 and 0.23, respectively. Juxtaposed against these measurements are negatively loaded dimensions such as superior facial height (nas-pros), nasal height (nas-ns), palatal length A (ol-sta), sagittal length of the nasal bones (nas-rhi) and superior facial length (bas-pros), which achieve variable loadings of -0.30, -0.30, -0.22, -0.20 and -0.20, respectively. The largest (Table 3.18) positively loaded MSV for the second PC include cranial vault length (g-o), sagittal height of the nasal aperture (rhi-ns), maximum cranial length (pros-o) and parietal sagittal chord (br-lam) which obtain variable loadings of 0.37, 0.31, 0.23 and 0.21, respectively. Working against these MSV are negatively loaded dimensions such as palatal length A (ol-sta), bizygomaxillare superior (bizs), sagittal length of the nasal bones (nas-rhi) and biauriculare breadth (operationally, maximum cranial width for this study) with negative variable loadings of -0.37, -0.34, -0.28 and -0.20, correspondingly. Table 3.19 provides the inter-generic Euclidean distance matrix produced from PCA object scores from the first to fifth principal components based on generic mean MSV. The average distance between cercopithecin genera is 0.69 (Range 0.80). Again, the largest distance is between Miopithecus and Erythrocebus at 1.18 while the smallest distance is between Cercopithecus and Chlorocebus with 0.37 followed by Cercopithecus and Allenopithecus, 0.42. For comparison, Table 3.20 provides intra-generic Euclidean distances based on MSV. The largest mean intra-generic Euclidean distance was produced by Allenopithecus at 0.70 (closely followed by Cercopithecus and Erythrocebus with 0.65 and 0.64). The smallest mean intra-generic Euclidean distance was again produced by Miopithecus at 0.51 (succeeded by Chlorocebus, 0.56). In summary, in contrast to Ln transformed data, for MSV the major distinguishing features between genera of the Cercopithecini are the dimensions of the neurocranium but

still naso-facial and palatal measurements contribute to discriminating genera. In particular, biasterionic breadth (biast), cranial vault length (g-o), parietal sagittal chord (br-lam) received some of the highest variable loadings.

3.5.3 CVA results for Ln transformed data: To consider and summarize the entire Cercopithecini (n = 89) multivariate datasets, as well as, “…weight differences between groups [or, genus, in this study] using the within- group dispersion as a standard of comparison” (Albrecht, 1978; p.37), canonical variates analysis (CVA) is most appropriate (in addition to confirming the results of PCA). By subjecting the Ln transformed pooled sex multivariate dataset to multivariate analysis of variation (MANOVA) and CVA, the first (Eigenvalue - 33.40; 68.46%) and second (Eigenvalue - 6.78; 13.90%) canonical variate axes were found to account for 82.36% of the variation (Figure 3.31). Similar to PCA, the mean CVA object scores for the first canonical axis per genus also seem to be sized related. Miopithecus has the lowest, 8.54 (Max. - Min., 8.72 - 8.28), with Erythrocebus receiving the highest, 10.47 (Max. - Min., 10.85 - 10.17) (Table 3.22). To test for significance of results, CVA object scores per genus (for both Ln transformed and MSV multivariate datasets) from the first ten canonical variate axes were subjected to a two-sample multivariate t-test using StatistiXL (version 1.5). For each pairwise generic comparison, the null hypothesis of no significant difference was rejected (p < 0.0001). Measurements which are positively correlated with canonical variates axis 1 include the maxillo-alveolar length (pros-dm3) and I1I2 alveolar length (i1i2, pros to septum between I2 & C1), 0.23 and 0.22 (Table 3.23). Working against these are measurements which are negatively correlated with canonical variates axis 1 include palatal height (palhei), biasterionic (biast), bifrontomalarorbitale (bifmt), bizygomaxillare superior (bizs) and bifrontomalartemporale (bifmt) breadths and cranial height (bas-br) with -0.42, -0.40, - 0.36, -0.29, -0.27 and -0.20, respectively. Cranial measurements that are positively correlated with the second CV axis includes the maxillo-alveolar breadth (biecm), palatal length A (ol-sta), biauriculare breadth (biaur), nasal height (nas-ns), sagittal length of the nasal bones (nas-rhi), bifrontomalarorbitale breadth (bifmo) and superior facial length (bas-pros) attaining

Figure 3.31: Cercopithecini scatterplot of canonical variate axes 1 & 2 for Ln transformed pooled sex data with 95% confidence ellipses. Allenopithecus - dot ; Miopithecus - cross +; Cercopithecus - square ; Chlorocebus - x; and Erythrocebus - circle .

3 Biecm & Palatal length

Axis 2: 2

13.90%

Bifmt Occipital sagittal chord & cranial vault length Cranial vault length & 8 9 10 11 CV Axis 1: 68.46 Bifmt MANOVA Wilks's Pillai lambda: 0.00 trace: 3.47 df1: 144.00 df1: 144.00 df2: 197.80 df2: 208.00 F: 11.54 F: 9.41 p(same): 0.00 p(same): 0.00 CVA Eigenvalue 1: 33.40 Percent: 68.46 Eigenvalue 2: 6.78 Percent: 13.90 Total %: 82.36

Table 3.21: Inter-generic Euclidean distances based on mean CVA scores from axes 1-5 per genus from Ln transformed data.

Al. 0 Mean 1.05 Mi. 1.47 0 Range 1.8 Cp. 0.44 1.50 0 Ch. 0.41 1.38 0.44 0 E. 0.91 2.22 0.78 0.96 0 Al. Mi. Cp. Ch. E.

Table 3.23: Variable loadings for Cercopithecini Table 3.22: Allenopithecus nigroviridis (n=6; 4, 2). Ln transformed pooled sex data. CVA scores Axis 1 Axis 2 CVA Mean 9.70 1.88 Eigen- Eigen- Max. 9.87 2.00 vectors Axis 1 vectors Axis 2 Min. 9.56 1.70 bifmt -0.21 g-o -0.37 Range 0.31 0.30 bicanex -0.19 bifmt -0.32 SD 0.13 0.11 bas-pros -0.17 bizi -0.16 CV 1.38 6.10 pros-o -0.15 nas-br -0.15 Miopithecus spp. (n=19; 9, 10). biast -0.13 nas-pros -0.13 CVA scores Axis 1 Axis 2 bizs -0.12 br-lam -0.12 Mean 8.54 (5.2) 1.28 palhei -0.12 bicanex -0.12 Max. 8.72 (1.1) 1.42 bizygo -0.02 bienm -0.11 Min. 8.28 1.17 ol-sta 0.00 iv-pms -0.11 Range 0.44 0.25 bipor 0.02 inbrnabo -0.10 SD 0.11 0.07 bien 0.05 bas-nas -0.10 CV 1.30 5.48 maxnawi 0.05 maxalvlen -0.09 Cercopithecus spp. (n=24; 12, 12). bienm 0.05 rhi-ns -0.07 CVA scores Axis 1 Axis 2 bifmo 0.06 bicanin -0.06 Mean 9.74 1.68 bas-br 0.07 zs-zygi -0.06 Max. 10.13 1.88 inbrnabo 0.07 pros-o -0.05 Min. 9.32 1.49 bizi 0.08 biast -0.04 Range 0.80 0.39 biaur 0.08 bien 0.00 SD 0.23 0.10 ol-pms 0.11 lam-opn 0.03 CV 2.35 5.98 bicanin 0.13 zs-zi 0.03 Chlorocebus pygerythrus (n=24; 13, 11). bas-nas 0.14 bizygo 0.05 CVA scores Axis 1 Axis 2 biseptal 0.16 i1i2 0.05 Mean 9.65 1.67 nas-pros 0.16 ol-pms 0.06 Max. 9.95 1.75 zs-zygi 0.16 maxnawi 0.07 Min. 9.33 1.56 br-lam 0.17 palhei 0.09 Range 0.62 0.20 biecm 0.17 bas-br 0.09 SD 0.18 0.05 zs-zi 0.18 biseptal 0.11 CV 1.91 3.05 iv-pms 0.18 bipor 0.12 Erythrocebus patas (n=16; 9, 7). rhi-ns 0.18 bizs 0.15 CVA scores Axis 1 Axis 2 nas-rhi 0.19 bas-pros 0.20 Mean 10.47 1.67 nas-ns 0.21 bifmo 0.24 Max. 10.85 1.83 nas-br 0.27 nas-rhi 0.25 Min. 10.17 1.50 i1i2 0.27 nas-ns 0.26 Range 0.69 0.33 maxalvlen 0.29 biaur 0.28 SD 0.23 0.10 g-o 0.29 ol-sta 0.30 CV 2.21 5.83 lam-opn 0.33 biecm 0.35

variable loadings of 0.35, 0.30, 0.28, 0.26, 0.25, 0.24 and 0.20, correspondingly. Juxtaposed against these measurements are dimensions which are negatively correlated with the second CV axis such as cranial vault length (g-o) and superior facial breadth (bifmt) with loadings of -0.37 and -0.32. The scatterplot for CV axes 1 & 2 of pooled sex Ln transformed data reveal the distinctiveness of Miopithecus and Erythrocebus (Figure 3.31); neither sample nor their 95% confidence ellipsis overlap with that of the other cercopithecins. Cercopithecus and Chlorocebus overlap considerably. The Allenopithecus sample and its 95% confidence ellipsis overlaps slightly with that of Cercopithecus and Chlorocebus, although this is most likely the result of small sample size and the inclusion of some subadults and with a larger more comprehensive sample would most likely not overlap. Table 3.21 provides the inter-generic Euclidean distance matrix produced from generic mean CVA object scores from the first to fifth canonical variate axes based on Ln transformed pooled sex data. The average Euclidean distance between cercopithecin genera is 1.05 (Range 1.80). The largest distance is between Miopithecus and Erythrocebus at 2.22 while the smallest distance is between Chlorocebus and Allenopithecus with 0.41. For comparison, Table 3.16 provides intra-generic Euclidean distances based on Ln transformed data. The largest mean intra-generic Euclidean distance was produced by Erythrocebus with 1.06 (followed by Cercopithecus and Allenopithecus with 0.91 and 0.90). This is due to the sexual dimorphism between males and females. The smallest mean intra-generic Euclidean distance was produced by Miopithecus with 0.71 (succeeded by Chlorocebus, 0.83). In summary, the CVA results for pooled sex Ln transformed data are similar to PCA results. Again, naso-facial measurements contribute most to discriminating genera but neurocranial dimensions are also amongst the higher loading variables. However, there are some measurements which receive high variable loadings for CVA but did not for PCA, such as the frontal sagittal chord (nas-br) and I1I2 alveolar length (i1i2).

3.5.4 CVA results for MSV: After converting the raw data of the Cercopithecini (n=89) into Mosimann Shape Variables (MSV) and subjecting the entire multivariate dataset to MANOVA and CVA the

first (Eigenvalue - 13.67; 45.47%) and second (Eigenvalue - 7.91; 26.31%) canonical variate axes account for 71.78% of the variation (10.58% less variation explained than Ln transformed pooled sex data; Figure 3.32). CVA object scores for the first and second canonical variates axes are both negative. Miopithecus on average scored the largest for CV axis 1, -2.18 (Max. - Min., -2.04 - -2.32) while Erythrocebus on average scored the least, - 1.34 (Max. - Min., -1.08 - -1.54), indicating differences due to size have been removed or at least corrected for (Jungers et al, 1995; Table 3.25). The two largest loaded MSV that are positively correlated with the first CV axis are the maxillo-alveolar length (pros-dm3) and I1I2 alveolar length (pros-to septum between C- I2 with 0.23 and 0.22 (Table 3.26). Juxtaposed against these two dimensions are MSV which are negatively correlated with second CV axis such as palatal height (palhei), biasterionic breadth (biast), bifrontomalarorbitale breadth (bifmo), bizygomaxillare superior (bizs), superior facial breadth (bifmt) and cranial height (bas-br) with variable loadings of -0.42, -0.40, -0.36, -0.29, -0.27 and -0.20, respectively. Cranial dimensions which are positively correlated with the second CV axis include palatal length A (ol-sta), maxillo-alveolar breadth (biecm), sagittal length of the nasal bones (nas-rhi) and biauriculare breadth (biaur) procuring variable loadings of 0.31, 0.26, 0.24, 0.22 and 0.21, correspondingly. Contrasting with these MSV are cranial dimensions that are negatively loaded with the second CV axis including inferior breadth of the nasal bones (inbrnabo), cranial vault length (g-o), palatal breadth (bienm), superior facial breadth (bifmt) and the length between incisivion and the central junction of the palato-maxillary suture (iv-pms) with variable loadings of -0.36, -0.32, -0.27, -0.26 and -0.25, respectively. Table 3.24 provides the inter-generic Euclidean distances matrix produced from generic mean CVA object scores from the first to fifth canonical variate axes based on pooled sex MSV. The average distance between cercopithecin genera is 0.52 (Range 0.58). Again, the largest distance is between Miopithecus and Erythrocebus at 0.89 but the smallest distance is between Cercopithecus and Allenopithecus with 0.33. For comparison, Table 3.20 provides intra-generic Euclidean distances based on MSV. The largest mean intra-generic Euclidean distance was produced by Allenopithecus at 0.70 (closely followed by Cercopithecus and Erythrocebus with 0.65 and 0.64). The smallest mean intra-generic Euclidean distance was again produced by Miopithecus at 0.51 (succeeded by Chlorocebus,

Figure 3.32: Cercopithecini scatterplot of canonical variate axes 1 & 2 for pooled sex MSV with 95% confidence ellipses. Allenopithecus - dot ; Miopithecus - cross +; Cercopithecus - square ; Chlorocebus - x; and Erythrocebus - circle .

0 -0.1 Palatal length -0.2 & Biecm -0.3 -0.4 -0.5 -0.6 -0.7 CV -0.8 -0.9 Axis 2: -1 26.31% -1.1 -1.2 -1.3 -1.4 -1.5 -1.6 Palatal height -1.7 & Biasterionic -1.8 Maxillo-alveolar Breadth -1.9 length & i1i2 Inferior breadth -2.9-2.8-2.7-2.6 -2.5-2.4-2.3-2.2 -2.1 -2 -1.9 -1.8 -1.7 -1.6 -1.5 -1.4 -1.3 -1.2 -1.1 -1 of the nasal bones & g-o CV Axis 1: 45.47%

MANOVA Wilks's Pillai lambda: 0.00 trace: 3.44 df1: 144.00 df1: 144.00 df2: 197.80 df2: 208.00 F: 9.35 F: 8.86 p(same): 0.00 p(same): 0.00 CVA Eigenvalue 1: 13.67 Percent: 45.47 Eigenvalue 2: 7.91 Percent: 26.31 Total %: 71.78

Table 3.24: Inter-generic Euclidean distances based on mean CVA scores from axes 1-5 per genus from MSV.

Al. 0 Mean 0.52 Mi. 0.80 0 Range 0.58 Cp. 0.33 0.72 0 Ch. 0.38 0.55 0.31 0 E. 0.39 0.89 0.38 0.46 0 Al. Mi. Cp. Ch. E.

100

Table 3.25: Allenopithecus nigroviridis (n=6; 4, 2). CVA scores Axis 1 Axis 2 Table 3.26: Variable loadings for Mean -1.58 -0.90 Cercopithecini pooled sex MSV. Max. -1.71 -1.06 CVA Min. -1.51 -0.76 Eigen- Eigen- Range 0.20 0.30 vectors Axis 1 vectors Axis 2 SD 0.08 0.10 palhei -0.42 inbrnabo -0.36 CV 5.14 10.95 biast -0.40 g-o -0.32 Miopithecus spp. (n=19; 9, 10). bifmo -0.36 bienm -0.27 CVA scores Axis 1 Axis 2 bizs -0.29 bifmt -0.26 Mean -2.18 -1.37 bifmt -0.27 iv-pms -0.25 Max. -2.32 -1.47 bas-br -0.20 bicanex -0.18 Min. -2.04 -1.25 biaur -0.12 bicanin -0.17 Range 0.27 0.22 bipor -0.12 nas-pros -0.15 SD 0.07 0.06 bizygo -0.11 zs-zygi -0.13 CV 3.11 4.23 biecm -0.10 bizi -0.13 Cercopithecus spp. (n=24; 12, 12). bicanin -0.09 nas-br -0.12 CVA scores Axis 1 Axis 2 bien -0.09 biast -0.12 Mean -1.65 -1.11 br-lam -0.08 bas-nas -0.11 Max. -1.95 -1.30 bicanex -0.06 br-lam -0.10 Min. -1.40 -0.95 inbrnabo -0.05 palhei -0.08 Range 0.55 0.36 pros-o -0.04 maxalvlen -0.08 SD 0.12 0.08 nas-br -0.03 rhi-ns -0.07 CV 7.43 7.22 bienm -0.02 bien -0.06 Chlorocebus pygerythrus (n=24; 13, 11). ol-sta -0.01 pros-o -0.03 CVA scores Axis 1 Axis 2 nas-rhi -0.01 zs-zi -0.02 Mean -1.73 -1.11 maxnawi -0.01 i1i2 0.00 Max. -1.90 -1.20 biseptal 0.00 bas-br 0.01 Min. -1.53 -1.04 bizi 0.00 bizygo 0.02 Range 0.37 0.17 zs-zi 0.03 maxnawi 0.02 SD 0.10 0.04 bas-nas 0.06 bipor 0.03 CV 5.89 3.65 iv-pms 0.09 ol-pms 0.05 Erythrocebus patas (n=16; 9, 7). bas-pros 0.10 lam-opn 0.06 CVA scores Axis 1 Axis 2 g-o 0.11 bizs 0.09 Mean -1.34 -1.20 lam-opn 0.13 biseptal 0.09 Max. -1.54 -1.29 rhi-ns 0.13 bifmo 0.12 Min. -1.08 -1.10 nas-ns 0.14 bas-pros 0.14 Range 0.46 0.19 ol-pms 0.14 biaur 0.21 SD 0.11 0.06 zs-zygi 0.15 nas-ns 0.22 CV 8.50 5.00 nas-pros 0.17 nas-rhi 0.24 i1i2 0.22 biecm 0.26 maxalvlen 0.23 ol-sta 0.31

101

0.56). In addition, Table 3.27 provides Cherry et al’s (1978) M distance between cercopithecin genera. The average M distance between genera is 1.19 (Range 1.53). The largest morphological divergence is achieved by Erythrocebus and Miopithecus, 2.14 (followed by Miopithecus and Allenopithecus at 1.83); the smallest is between, Cercopithecus and Allenopithecus, 0.61 (succeeded by Cercopithecus and Chlorocebus at 0.65).

Table 3.27: Inter-generic Cherry et al’s M distance between cercopithecin genera. Al. 0 Mean 1.19 Mi. 1.83 0 Range 1.53 Cp. 0.61 1.52 0 Ch. 0.9 1.63 0.65 0 E. 0.79 2.14 0.81 1.03 0 Al. Mi. Cp. Ch. E.

Again, the scatterplot for CV axes 1 & 2 of pooled sex MSV reveals the distinctiveness of Miopithecus and Erythrocebus (Figure 3.32). Again, Cercopithecus and Chlorocebus overlap considerably, with Allenopithecus only slightly so. In summary, CVA results for pooled sex MSV are similar to those for PCA. However, there are some measurements which receive high variable loadings that did not for PCA, such as I1I2 alveolar length (i1i2) and palatal height (palhei) (Table 3.18).

3.6 Limb Proportions and Postcranial Features: Published data on postcranial elements and limb proportions for the cercopithecins (Tables 3.1-3.5) was gathered from Hill (1966), Manaster (1979), Napier (1981), Strasser & Delson (1987), Strasser (1988, 1992 & 1994), Groves (1989 & 2001a), Nakatsukasa (1994), Gebo & Sargis (1994), Anapol & Gray (2003), Anapol et al (2004 & 2005) and Whitehead et al (2005). Postcranially, many distinctions between genera are possible due locomotor behaviors and preferred substrate. Allenopithecus is distinguished from the other guenons by its stockily-built body (Hill, 1966) as well as its large first digit of the foot and dissimilar form of walking compared to other cercopithecins (Kingdon, 1988). Miopithecus is peculiar due to its small size. According to Gebo & Sargis (1994) some features associated with arboreality include a smaller body size, shortened navicular for climbing and longer legs for leaping along with a proximal location of the tibial tubercle. Many

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indices relating to the fore- and hindlimb such as the intermembral index are an excellent indicator of locomotor behavior across the primates (Fleagle, 1998; Watkins, 2003). Within the cercopithecini, the patas is the most derived at 91-92% compared to the others which range from 79% to 86%; not surprising since Erythrocebus is the most terrestrial of this group. Of particular interest is the range of variation seen in the genus Cercopithecus (Tables 3.1-3.5). The fore- and hindlimb indices listed for Cercopithecus have ranges of variation larger than the other cercopithecins, which suggests different forms of locomotion and substrate use and/or preference within this genus. For example, the intermembral index for C. ascanius, C. diana and C. cephus is 79% while C. mona has 86&; and C. neglectus and C. preussi have 82% (Napier, 1981; Rowe, 1996; Fleagle, 1998). In addition, Kingdon (1988) and Gebo & Sargis (1994) list several postcranial features which the patas, vervets and the lhoesti-group guenons share due to locomotor terrestrial adaptations. Features associated with some form of terrestriality include reduction of the hand and foot digits, a limited humeral facet upon the ulna and slim patellar facet; the former indicates a cursorial locomotor repertoire while the latter two reflect reduced joint size and mobility. However, it must be admitted that all members of the cercopithecini occasionally spend time on the ground and in trees either to gather food or sleep. For example, patas monkeys must climb trees to sleep and avoid predators (similar to the behaviors of baboons and chimpanzees) whereas other species rarely come to the ground (C. mitis and C. ascanius). Thus, the postcranial skeleton contains a mosaic of features allowing travel in both substrates. In addition, “semiterrestrial guenons [e.g. A. nigroviridis & C. neglectus] do not appear to be strikingly different in their overall limb morphology from that of the arboreal guenons [e.g. C. mitis and C. ascanius] (Gebo & Sargis, 1994, pp. 350-351). These resemblances they conclude must result from similar limb mechanics of arboreal and semiterrestrial travel. Overall, traits associated with terrestrialism include a high intermembral index, longer tarsals (navicular and cuboid), reduced joint mobility at the elbow and shortened digits. With the genetic data from Tosi et al (see below) and the many observations made by Hill (1966), a grouping of the semiterrestrial and terrestrial guenons separate from the more arboreal species seems appropriate. For example, Hill described the scapula of C. aethiops and C. lhoesti as having “the classical quadrupedal form” (longer in the coronal

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than sagittal plane) (p. 597). Again concerning the scapula, Hill (p. 665) discusses the similarities of Erythrocebus, C. lhoesti and the C. aethiops superspecies (note, in Hill’s classification scheme Chlorocebus is not recognized; he uses Cercopithecus aethiops.). In particular he notes that they share long spinous processes following perpendicular to the vertebral border, as well as an increase in the size of their supraspinous fossa. The scapula is an important determinant in locomotion and all the semi- and terrestrial guenons have a scapular index above 100% while the mainly arboreal guenons have less (ibid p. 252), although as Hill and Gebo & Sargis (1994) note, C. ascanius is very close with 97%. To summarize, postcranially the cercopithecins are quite varied. The range of variation within Cercopithecus is perhaps more than what should be expected of a genus whose members share an adaptive zone.

3.7 Genetics of the Cercopithecini: Of all the extant catarrhines, although oddly familiar to the Hylobatidae, members of the Cercopithecini have highly varied diploid numbers revealing a complex chromosomal evolution within this tribe, most likely the result of vicariance and isolation events due to fluctuations in tropical forest range and fragmentation during the Late Miocene and into the Plio-Pleistocene (Dutrillaux, 1988; Grubb, 1990 & 1999). These include, 2n=48 for Allenopithecus nigroviridus; 2n=54 for Miopithecus spp.; for the guenons of Cercopithecus the 2n is quite varied - 58 diana-group; 70-72 mitis-group; 66-68 mona-group; 66 cephus-group; 60 lhoesti-group; 64 hamlyni; & 58-62 neglectus-group; 2n=60 for Chlorocebus spp.; and lastly, 2n=54 for Erythrocebus patas. Recent genetic analyses by Tosi et al (2002a & 2002b; 2003a; 2004) have greatly clarified the relationships within the cercopithecini using multiple independent loci, including, Y- chromosomal DNA sequences - SRY (sex determining region, Y-chromosome) & TSPY (testis specific protein, Y-chromosome) and an intergenic 1.6 kb region on the X- chromosome was sequenced for 20 guenons and four out-group taxa (Macaca, Mandrillus, Presbytis and Trachypithecus). The two Y-chromosome genes are located in non- recombining portions analyzed together but the X-chromosome separately. The objective of these analyses was to determine if the more terrestrial guenons (E. patas, Chlorocebus spp. and C. lhoesti-group) formed a monophyletic clade and represents a single transition to

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more terrestrial habitat use as opposed to the other mainly arboreal guenons, such as Cercopithecus mitis or C. ascanius. Their results place Allenopithecus and Miopithecus as the basal members of the cercopithecins (both of which utilize arboreal and terrestrial substrates but postcranially resemble the arboreal guenons) and the more terrestrial guenons are a monophyletic group to the exclusion of the other arboreal guenons. Thus, Cercopithecus as defined by Groves (2001a) is paraphyletic (Tosi et al, 2004; see also Van Der Kuyl et al, 1995). Table 3.28 is a distance matrix displaying the genetic distances between genera (provided by A. Tosi to author, from data in Tosi et al, 2003 & 2004). The largest genetic distance between genera is that between Allenopithecus and Cercopithecus (0.0157), while the smallest distance is between Chlorocebus and Erythrocebus (0.0062).

Table 3.28: Inter-generic uncorrected ‘p’ genetic distances between cercopithecin genera from Tosi et al, 2003 & 2004. E. 0 Mean 0.013 Ch. 0.0062 0 Range 0.0095 Cp. 0.0136 0.01462 0 Mi. 0.013 0.0138 0.01375 0 Al. 0.014 0.0149 0.0157 0.0152 0 E. Ch. Cp. Mi. Al.

To summarize, despite cranial (Verheyen, 1962; Schultz, 1970; and this study) and postcranial similarities (Gebo & Sargis, 1994), genetic analyses offer a very clear picture of cercopithecin phylogenetic relationships.

3.8 Discussion and Conclusions: From the varied evidence presented in this chapter, a number of inferences and conclusions are possible. 3.8.1 Question 1: Are the extant Catarrhini genera adequately defined and do they compose an ‘adaptively coherent’ grouping of one or more species which occupy an ‘adaptive zone’, as defined by Wood & Collard (1999a)? Aside from Cercopithecus, the remaining genera (Miopithecus, Allenopithecus, Chlorocebus and Erythrocebus) are adequately defined, each adaptively coherent and occupying different adaptive zones. The type species of Cercopithecus is C. diana (see Stiles & Orleman, 1926). C. diana is highly arboreal and generally occupies the middle to upper forest strata, has an intermembral index of 79% and associates with many other

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primate species (Gautheir-Hion, 1988a; McGraw, 1994; Rowe, 1996; Enstam, & Isbell, 2007). As such, the other members of Cercopithecus should have similar adaptive strategies to the type species (a criterion proposed by Wood & Collard, 1999a). However, this is not the case. For example, C. campbelli is described by Rowe (1996) as “the most terrestrial guenon” (p. 155) and prefers the middle and lower forest strata. C. erythrogaster prefers the lower forest levels and thick undergrowth (Rowe, 1996). C. mona has an intermembral index of 86% and favors lower levels of second growth forests (Rowe, 1996). C. lhoesti is considered semi-terrestrial (Gebo & Sargis, 1994). C. neglectus has an intermembral index of 82% and prefers dense, swamp or bamboo forests, will dig for invertebrates and consumes a high proportion of leaves in its diet (Rowe, 1996) and associates with A. nigroviridis (McGraw, 1994). Furthermore, non-metric characters, such as the robusticity of the nuchal crest or temporal lines also hint at differences in positional behavior, preferred substrate and diet. For instance, in highly arboreal species the neurocranium is more globular and the nuchal crest is less developed (e.g. C. cephus or C. ascanius) while more terrestrial species have a more elongated face and cranium with a stronger developed nuchal crest (e.g. C. lhoesti or C. hamlyni).

Question 2: In terms of cranial morphology, how distinct are genera? In other words, how readily may a genus be determined from cranial dimensions, considered either singular or multiple? With regard to cranial distinctiveness of the cercopithecins, Miopithecus, Allenopithecus and Erythrocebus are highly diagnostic, by either metric or non-metric features. Chlorocebus and Cercopithecus overlap in size and shape. Several cranial dimensions are significantly correlated to body size (Table 3.14 and Figure 3.28). Miopithecus spp.: Talapoins are distinct simply because of their diminutive yet globular cranium with weak muscle markings and little facial projection. Cranial height (bas-br), biporionic and biasterionic breadths (bipor and biast) less than 50 mm; height of face (nas-pros) less than 30 mm (Figures 3.2, 3.3, 3.5 & 3.14). Globular neurocranium with no sagittal cresting in males or females. Cranium lightly built with no strong muscle markings. On average, maxillo-alveolar breadth (biecm) relatively greater than palatal length A (ol-sta; Table 3.10, no. 26), the other cercopithecins were less. Superior facial breadth (bifmt) relatively greater than superior facial height (nas-pros; Table 3.10, no. 37).

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Largest relative proportion of cranial vault length (g-o) to maximum cranium length (pros- o), > 80%, compared to other cercopithecins, < 80% (Table 3.10, no. 51). Allenopithecus nigroviridis: Although, Allenopithecus overlaps in size and limb proportions with Cercopithecus and Chlorocebus, the swamp monkey’s nasal region, palate and dentition (Lang, 1923) are very distinct from the guenons and vervets. In addition, the swamp monkey has relatively long nasal bones (nas-rhi) and palate (ol-sta) in relation to cranial (bas-br) and facial height (nas-pros) (Figures 3.25, 3.26 & 3.27). Sagittal crest common in males. Small length of the parietal sagittal chord (br-lam) and palatal breadth (bienm) compared to Cercopithecus and Chlorocebus (Figures 3.4 & 3.10). Relative length of the zygomatic (zs-zgyi) to palate length (ol-sta) on average the least in Allenopithecus, ~89%, compared to other cercopithecins, >90% (Table 3.10, no. 71). Cercopithecus spp.: The Cercopithecus sample significantly differs from the Chlorocebus sample in the bizygomaxillare superior breadth (bizs; Figure 3.7) and inferior breadth of the nasal bones (inbrnabo; Figure 3.10). The former (bizs) dimension is on average wider in Chlorocebus than Cercopithecus, whereas the latter (inbrnabo) is wider in Cercopithecus than Chlorocebus. Sagittal cresting is not common, although some species exhibit strong temporal lines and developed nuchal crests. Chlorocebus spp.: Sagittal cresting is not common, although some adults, both female and male, can exhibit strong temporal lines. Metrical and proportional overlap with Cercopithecus spp. is common. Some features which can discriminate the two include the sharing of a V-shaped auditory meatus at the inferior margin and angular shaped eye-orbits, not oval as in the guenons (Groves, 1989). Erythrocebus patas: Absolutely largest of the cercopithecins in terms of cranial vault (g-o; Figure 3.9) and zygomatico-maxillary suture (zs-zi; Figure 3.16) lengths and superior facial height (nas-pros; Figures 3.7) with) with sagittal crests and strong nuchal lines in adult males. Relative maxillo-alveolar breadth (biecm) to length of the zygomatic bone (zs-zgyi) is the least in Erythrocebus, ~82%, compared to the other cercopithecins, >90% (Table 3.10, no. 28; Figure 3.19). In addition, Erythrocebus is dissimilar from the other cercopithecins because the nuchal plane and nuchal crest are highly developed and extreme sexual dimorphism.

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Question 3: How much morphological variation is encompassed within a Catarrhine genus? Morphological variation in cercopithecin genera can produce standard deviations and coefficients of variation < 5 to ~ 20. Highly variable measurements are associated with the nasal and facial region. In contrast, some neurocranial variables produce less variation. Intra-generic Euclidean distances based on Ln transformed data were on average 0.88 (Max. - Min., Erythrocebus, 1.06 - Miopithecus, 0.71), while those based on MSV were 0.61 (Max. - Min., Allenopithecus, 0.70 - Miopithecus, 0.51) (Tables 3.16 & 3.20).

Question 4: How much has one genus morphologically and genetically diverged from another? With regard to morphological divergence, measuring size or shape differences, not surprisingly, Miopithecus and Erythrocebus are polar extremes with the other three genera in between. Euclidean distances from Ln transformed data or MSV between Miopithecus and Erythrocebus are the largest (see Tables 3.15, 3.19, 3.21, 3.24, & 3.27). The smallest Euclidean distances are generated between Cercopithecus, Chlorocebus and Allenopithecus. Euclidean distances based on shape (MSV) are generally smaller than those based on size suggesting underlying shape (i.e. proportions) and/or developmental similarities. In other words, while size is an important factor in generic differentiation, shape is more conservative. The morphological distances generated from the cercopithecin datasets is the result of >10 million years of evolution (cercopithecin-papionin estimated divergence; see Page & Goodman, 2001; Raaum et al, 2005; a minimum date would be roughly 6-7 Ma because there are Macaca fossils in the circum-Mediterranean region which is evidence for a phylogenetic divergence between these two tribes; see, Delson, 1980 and Kohler et al, 2000). However, inter-generic genetic distances reveal a close relationship between Erythrocebus and Chlorocebus (0.0062; Table 3.28), whereas the greatest genetic distance is between Allenopithecus and Cercopithecus (0.0157) (Tosi et al, 2003 & 2004). The average inter-generic genetic distance is 0.013 (Range 0.0095).

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Miopithecus spp.: Talapoins occupy dense, well-watered habitats. Associated with this is a cranium that is lightly built with no sagittal crest. There is no indication that the cranium is adapted to withstand or assist in labored mastication requirements such as that seen in Theropithecus or Gorilla, both of which can develop large sagittal crests and strong muscle markings on the neurocranium, maxilla and zygomatic bones. The small size also indicates that this genus is not adapted to open, terrestrial environs, instead preferring dense gallery or inundated forests. Furthermore, the small size, lack of a strongly developed nuchal crest and slight facial projection suggests a well-balanced and easily maneuverable cranium adapted to arboreal movements and dense foliage and a posture that may entail more sitting, leaping and climbing (Gauthier-Horn, 1973) (see also generic description for question 2 above and generic summaries below). Allenopithecus nigroviridis: Allen’s swamp monkey prefers the lower to middle forest strata and as the name implies, favors well-watered or inundated swampy forests. Associated with this is a cranium that is quite robust and macaque-like. The elongated face and well-developed nuchal crest suggest functional adaptations to semiterrestrialism (see also generic description for question 2 above and generic summaries below). Cercopithecus spp.: Guenons occupy tropical forested areas and use various strata of the rainforest. Associated with this is a cranium that is quite varied and versatile and perhaps is best explained as a compromise between the use of arboreal and terrestrial niches. Many differences may be discerned from the nasal region (Verheyen, 1962; Vogel, 1966 & 1968), which could be related to mate recognition (including pelage coloration and vocalizations) and/or epigenetic developmental factors affected by climate, latitude and habitat use (see also generic description for question 2 above and generic summaries below). Chlorocebus spp.: Vervets live in multimale-multifemale groups (Isbell et al, 2002) which exploit both arboreal and terrestrial niches near permanent water sources. Associated with this is a cranium that is similar to species of Cercopithecus and Erythrocebus. Vervets do not develop a sagittal crest but can have deep, robust temporal lines. Like the guenons, the crania of the vervets indicate the ability to use and move either in trees or on the ground (see also generic description for question 2 above and generic summaries below).

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Erythrocebus patas: Patas monkeys enjoy open savanna, but trees and tall bushes are exploited for spotting and evading predators, as well as aiding in group or individual location and cohesion. Associated with this is a cranium that is robust and long. The increase in body size and intermembral index compared to the other cercopithecins is testament to its preference for a terrestrial lifestyle and open environments. However, as McCrossin et al (1998; and Blue et al, 2006) have discussed in relation to the fossil genera Kenyapithecus and Victoriapithecus, it is perhaps more appropriate to argue that an increase in body size (and therefore skull size as well) is a consequence of a terrestrialism not a cause. Selective pressures may include but are not limited to 1) predator avoidance and/or defense; 2) intra-specific male to male aggression or competition and sexual selection; 3) habitat preferences; and 4) troop control or maintenance (see also generic description for question 2 above and generic summaries below).

Question 6: What analogies may be drawn from extant Catarrhini genera for the interpretation of fossil hominin genera? The cercopithecins and their generic arrangement can provide analogies and considerations for the interpretation of fossil hominins. First, size is an important factor in delineating genera. Second, taking into consideration the metrical overlap and similarity of cranial (Verheyen, 1962; Schultz, 1970; and this study) and dental (Plavcan, 1993; Cope, 1993; Plavcan & Cope, 2002) features of some cercopithecin genera (e.g. Allenopithecus, Cercopithecus and Chlorocebus), it would be difficult to discern fossil genera or species from only fragmentary skeletal remains (e.g. Eck & Howell, 1972). Lastly, limb proportions and niche or habitat preference can assist in the delineation of genera, as well as indicating locomotion. Sister species (or genera) can have radically different phenotypes (e.g. Erythrocebus and Chlorocebus).

3.8.2 Generic Summaries: Tribe Cercopithecini - Groves (1989 & 2001a; and Rowell, 1988; see also Gautheir-Hion, 1988a & 1988b) lists several features which define this tribe. Some of these include, 1) Diploid number varied; 2) Male cercopithecins are only weakly integrated into the troop’s social organization; 3) Lack of complex social hierarchies as seen in papionins; 4) Lack of facial communicative gestures, again, as seen in the papionins

(ex. Miopithecus); 5) tribe members generally lack a hypoconulid on M3; 6) smaller and more discrete ischial

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callosities; 7) auditory tubes are transverse in cercopithecins (ex. Miopithecus & Allenopithecus) but postero- laterally in papionins; and 8) reduced palatine foramina. Allenopithecus Lang, 1923. A monotypic genus, A. nigroviridus. 2n = 48. Allenopithecus is a peculiar genus occupied by a very little studied monkey, Allen’s swamp monkey, in part by its choice of habitat and cryptic behavior. Recent genetic analysis by Tosi et al (2002, 2003 & 2004) places Allenopithecus unequivocally as the basal member of the Cercopithecini. This is in agreement with Strasser & Delson’s (1987) cladistic analysis based on skeletal and soft tissue evidence. For example, Allenopithecus displays two symplesiomorphies - ‘molar flare’ and the fusion of the ischial callosities across the midline in males, both more common in Papionini; as well as its stocky-macaque-like body shape (Groves, 2001a). However, in a different analysis, Groves (2000) suggested that Allenopithecus does not belong to the Cercopithecinae. Justification for this rests on cladistic analyses of primarily craniodental features, such as facial elongation and the observation of female sexual swelling (see Lang, 1923; Verheyen, 1962; Hill, 1966). Verheyen (1962) noted that the cranium of Allenopithecus does not have a supra-glabellar fossa; long, sloping, nearly continuous face in lateral view from nasion to prosthion; flat frontal; strong nuchal crest; and a long, narrow cranial vault. Miopithecus I. Geoffroy, 1842. 2 species, M. talapoin Southern talapoin [type species genus]& M. ougouensis Northern talapoin. 2n = 54. The talapoins, or dwarf guenons, represent the smallest Old World monkey. Like Allenopithecus, Miopithecus is also interpreted as a basal member to the Cercopithecini (Tosi et al 2002). Allenopithecus and Miopithecus female species both produce sexual swellings (Hill, 1966), which has been observed in several Old World primate genera (e.g. Macaca, Papio, Procolobus, Trachypithecus, Simias, Hylobates and Pan) and may be interpreted as a primitive trait - perhaps a primitive characteristic for the Catarrhini in general (Davies & Oates, 1994; van Schaik et al, 1999; Groves, 2000 & 2001; Cheyne & Chivers, 2006; Shelmidine et al, 2007). However, in a different analysis, Groves (1989) suggested that Miopithecus does not belong in the Cercopithecini but rather with the Papionini. In addition, the social systems of both genera are between papionins and other cercopithecins. Like papionins, Allenopithecus and Miopithecus individuals live in multimale groups at times but like cercopithecins interaction between male and females are limited to short mating periods (Tosi et al, 2002). Van der Kuyl et al (2000) provide genetic evidence to support the specific separation of M. 111

talapoin and M. ougouensis. Verheyen (1962) noted that the cranium of Miopithecus is lightly built; has no supra-glabellar fossa; weak or light nuchal crest; low temporal lines; and short and triangular nasal bones. Cercopithecus Linnaeus, 1758. A polytypic genus comprising 25 species clustered into eight species-groups based on pelage, vocalizations, behavior, ecology, preferred tree strata and chromosome number; some are sympatric and/or allopatric and largely arboreal but other species are more prone to exploit terrestrial habitats then others (Groves, 2001a); 1 - C. dryas (Dryas or Salongo Monkey), (2 - Diana-group C. diana [type species of genus; Diana Monkey], C. roloway (Roloway Monkey)), (3 - Mitis-group C. nictitans (Greater Spot-nosed Monkey), C. mitis (Blue Monkey), C. doggetti (Silver Monkey), C. kandti (Golden monkey), C. albogularis (Syke’s Monkey)), (4 - Mona-group C. mona (Mona Monkey), C. campbelli (Campbell’s Mona), C. lowei (Lowe’s Mona), C. pogonias Crested Mona), C. wolfi (Wolf’s Mona), C. denti (Dent’s Mona)), (5 - Cephus- group C. petaurista Lesser Spot-nosed Monkey), C. erythrogaster (White-throated Guenon), C. scalteri (Sclater’s Guenon), C. erythrotis (Red-eared Guenon), C. cephus (Mustached Guenon), C. ascanius (Red-tailed Monkey)), ( 6 - Lhoesti-group C. lhoesti (L’Hoest’s Monkey), C. preussi (Preuss’s Monkey), C.. solatus (Sun-tailed Monkey)), 7 - Hamlyni-group C. hamlyni (Hamlyn’s Monkey or Owl-faced Monkey) and 8 - Neglectus- group C. neglectus (De Brazza’s Monkey). 2n has considerable variation, reflecting a complex chromosomal evolution (Dutrillaux et al, 1988; Ruvolo, 1988) - 58 diana-group; 70-72 mitis-group; 66-68 mona-group; 66 cephus-group; 60 lhoesti-group; 64 hamlyni; & 58-62 neglectus-group. In addition, guenon-like fossils have also been allocated to this genus from East Africa (Szalay & Delson, 1979; Leakey, 1988; Pickford & Senut, 1988; Benefit, 2000; Gundling & Hill, 2000). The monophyly of Cercopithecus has been and continues to be questioned for a number of reasons (Purvis & Webster, 1999; Groves, 2000; Tosi et al, 2004). Furthermore, species of this group are known to occasionally hybridize in the wild (e.g. C. ascanius and C. mitis; Struhsaker et al, 1988; Detwiler, 2002). Locomotion and habitat preference varies within this genus and are reflected in postcranial elements (Manaster, 1979; Gebo & Sargis, 1994; Anapol et al, 2005). Chlorocebus Gray, 1870. A polytypic genus comprising 6 species; C. sabaeus, aethiops [type species of genus], C. djamdjamensis, C. tantalus, C. pygerythrus and C.

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cynosuros; C. sabaeus is the type species of this genus. 2n = 60. Vervets are the largest distributed monkey in Africa, present in many sub-Saharan countries, although close to water sources. The genus Chlorocebus was until recently not used and the species of this genus were subsumed under Cercopithecus (e.g. Napier, 1967 & James, 1960); and this classification scheme is still used by some (e.g. Grubb et al, 2003; Bolter & Zihlman, 2003). However, genetic and morphological evidence has shown that the species of Chlorocebus are more closely related to Erythrocebus (Disotel, 2000; Grooves, 2000; Tosi et al, 2002). Recent postcranial analysis by Anapol et al (2005) confirms the terrestriality of Chlorocebus aethiops, which has been independently corroborated by the genetic analysis by Tosi et al (2004) linking the more terrestrial “guenons” - Erythrocebus, Chlorocebus and Cercopithecus lhoesti-group. Chlorocebus spp. do display sexual dimorphism (Turner et al, 1997) but not to the extent seen in Erythrocebus. Erythrocebus Trouessart, 1897. A monotypic genus, E. patas. 2n = 54. The Patas monkey is the fastest ground-running digitigrade primate (Rowe, 1996; other monkeys are palmigrade) and is the most terrestrial among the cercopithecins with very large home- ranges, preferring open woodland-bushland savanna (Hall, 1968). In fact, Gebo (1994) used postcranial data to demonstrate that E. Patas and other genera, such as, Chlorocebus and Cercopithecus lhoesti & C. preussi all exhibit terrestrial adaptations to some extent, particularly with regard to limb elements, but there is overlap. Despite these and other adaptations some do not feel generic separation from Cercopithecus is necessary (e.g. Chism & Rowel, 1988), yet, Stanyon et al (2005) report karyological evidence clearly uniting the patas with Chlorocebus. Verheyen (1962) noted that the cranium of Erythrocebus has a long and narrow palate and cranial vault; adult males develop large nuchal and sagittal crests and very large canines.

113 Chapter 4: Results for genera of the Papionini

Rungwecebus Cercocebus Mandrillus Macaca Lophocebus Papio Theropithecus

Papionini Cercopithecinae Cercopithecidae Cercopithecoidea Catarrhini Figure 4.1: A taxonomic and phylogenetic Simiiformes diagram representing the likely relationships within the Papionini based on biomolecular Haplorrhini and morphological data. Primates

4.1 Introduction: The purpose of this chapter is to report the results for genera of the Papionini. Groves (2001a) taxonomy for the Tribe Papionini (Burnett, 1828) includes six genera; Cercocebus (6 species), Mandrillus (2 species), Macaca (20 species), Lophocebus (3 species), Papio (5 species) and Theropithecus gelada (Figure 4.1). However, in 2005 a new genus and species was described from Tanzania, Rungwecebus kipunji (Jones et al, 2005; Davenport et al, 2006). No large comparative sample exists and further corroboration by other researchers is necessary. The average number of species per papionin genus is 6.2, ranging from one to twenty with a standard deviation of 7.0. However, if Macaca is excluded, average number of species per genus drops to 3.4, ranging from one to six with a standard deviation of 2.0. Furthermore, if the Cercopithecini are considered alongside the

Papionini, as both compromise the Cercopithecinae (eleven genera and 72 species), the average number of species per genus is 6.5, ranging from one to twenty-five with a standard deviation of 8.2. However, if Cercopithecus (25 species) and Macaca (20 species) are excluded (considering only nine genera and 27 species), the average number of species per genus reduces to 3.0, ranging from one to six, with a standard deviation of 2.1. The species of papionin genera range in size from less than 10 kg (e.g. Lophocebus and Cercocebus) to greater than 35 kg (Mandrillus and Papio). Most species of this ‘adaptive array’ (Jolly, 1970) may be characterized by some degree of sexual dimorphism, facial elongation and a diploid number of 42. Degree of facial elongation varies and creates two informal groups; 1) the first group of genera are small to medium sized with moderate facial projection, including, Lophocebus, Cercocebus and Macaca and are more arboreal; and 2) the second group of genera are medium to large with extreme facial projection and are much more terrestrial, including Theropithecus, Papio and Mandrillus. However, despite any terrestriality these genera may exhibit, all require trees, rocky outcrops or cliffs for predator evasion and sleeping sites. In addition, members of this tribe possess cheek pouches (which most likely promoted the adaptive radiation and survival/foraging strategies of the Cercopithecinae; see Lambert, 2005; Buzzard, 2006b) a feature shared with their sister-tribe the Cercopithecini. Previously researchers (e.g. Delson, 1975; Groves, 1989; Whitehead et al, 2005) stated papionins lack enamel on the lingual surface of their lower incisors but Aimi & Nogami (1989) have presented evidence that this is not the case. As such, the Papionini represent the sister tribe to the Cercopithecini and are each grouped together in the subfamily Cercopithecinae, and then grouped with the Colobinae in the family Cercopithecidae of the superfamily Cercopithecoidea Gray, 1821 (Delson, 1975; Andrews, 1981). However, unlike the Cercopithecini, the Papionini are well sampled from the fossil record in Africa and Eurasia (Pan & Jablonski, 1987; Barry, 1987; Ardito & Moturra, 1987; Geraads, 1987; Jablonski & Yuerong, 1988; Ciochon, 1993; Delson et al, 2000; Jablonski, 2002). Indeed, all papionini genera, excluding Mandrillus, have generically allocated fossil material. Tables 4.1 to 4.6 list several key features which characterize the adaptive zone of these genera. Excluding Theropithecus, which is a gramnivore (Dunbar, 1983 & 1993;

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Table 4.1: Cercocebus spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) Average Habitat (,; (range) Min.-Max.) 11019, 6400*; Intermembral Quadrupedal - Frugivore + Terrestrial and Multimale- 4000-13000 82.85 (80-87) above branch, seeds and some arboreal/ Multifemale/ Brachial sitting & animal prey Tropical forest 1.43 -1.61 100.6 (93.2- walking & savanna (C. torquatus) 109.7) Crural 89.1 (85.8-94.1)

Table 4.2: Mandrillus spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) Average Habitat (,; (range) Min.-Max.) 21950, Intermembral Quadrupedal - Frugivore + Terrestrial and Multimale- 10750*; 92 (88-96) above branch, seeds and some arboreal/ Multifemale 10000-45000 Brachial sitting & animal prey Tropical forest with 1 male- 104 (102-108) walking multifemale Crural subgroups/ 85 (81-88) 1.19-1.29 (159.4g adult brain weight - M. sphinx)

Table 4.3: Macaca spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) Average Habitat (,; (range) Min.-Max.) 10553.7, Intermembral Quadrupedal - Frugivore + Arboreal and Multimale- 6829.14*; 92 (84-99) with some Omnivory - Terrestrial/ multifemale, 1- 4000-25000 Brachial leaping, above fruits, seeds, Tropical to 2 male- 97 (90-104) branch sitting animal prey, Temperate multifemale/ Crural & walking plants, flowers, lowland 1.95 (70-110g 88 (81-93) leaves, primary & adult brain secondary, weight) gallery, mangrove, evergreen, montane, coniferous, monsoon & dry forests. Up to 3000m.

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Table 4.4: Lophocebus spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) Average Habitat (,; Min.-Max.) 8092, 5557*; Intermembral Quadrupedal - Frugivore + Arboreal & Multimale – 4500-9000 78 (76-81) above branch, seeds some terrestrial/ multifemale/ Brachial sitting & walking Tropical 1.53-1.73 95.8 (89.6-100) primary and (L.albigena) Crural semideciduous 89 (86-92.1) forest, middle to upper canopy

Table 4.5: Papio spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) Average (range) Habitat (,; Min.-Max.) 23674, Intermembral Quadrupedal, Frugivore + Terrestrial/ 1 male- 13027*; 95 (90-100) ground standing omnivory Tropical forest, multifemale, 12000-40000 Brachial & walking woodland & multimale- 103 (96-113) savanna multifemale/ Crural 1.18-1.69 83 (80-91) (P. anubis)

Table 4.6: Theropithecus gelada Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) Average Habitat (,; Min.-Max.) 18375, 11920; Intermembral Quadrupedal - Herbivore Terrestrial/ Multimale- 10000-30000 97 ground standing Topical multifemale: Brachial & walking highland grass combination 109 savanna of many Crural harems & 93 bachelor groups/ .75-1.04 (131.9g adult brain weight)

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Iwamoto, 1993), all papionins may be characterized as frugivores but flowers, seeds and animal prey (including invertebrates or vertebrates) are also a dietary component. Excluding Lophocebus and some species of Macaca, most papionins exploit terrestrial niches. However, despite any degree of routine terrestriality, all papionins require trees, or at least rocky outcrops or cliffs, for sleeping sites and predator evasion. All genera, excluding Macaca, are restricted to sub-Saharan Africa, although a population of hamadryas baboons does exist along the southern border of the Arabian Peninsula. The genus Macaca has only one relict species (M. sylvanus) in mountainous Northern Africa and Gibraltar while all other macaque species are distributed from Pakistan to China, Japan and Pacific Southeast Asia. In 2005, Sinha et al announced the discovery and description of a new macaque species, M. munzala, in northeastern India at very high altitudes (2000- 3500m), which they believe is a member of the sinica-group. The papionins, like many other primate groups, have a complex taxonomy. Indeed, one example among many is Papio, for which the ICZN had to rule on the priority, suppression and availability of genus and species names (see Groves, 2001a and references therein). With regard to competing classificatory schemes, most disagreements are concerned with the content (i.e. the number of species) of these genera and whether or not some should be accorded subgeneric or subspecies status rather than full generic rank. For example, Goodman et al (1998 & 2001) propose Lophocebus and Theropithecus to be subgenera within Papio based on genetic similarity and their estimated time of origin, <5 Ma. Their work also suggests a similar arrangement for Mandrillus, which they treat as a subgenus of Cercocebus. Another disagreement is the specific and/or subspecific status of the species within Papio because hybrid zones and clinal distributions have been observed between species (Jolly, 1993, 2001 & 2003). Yet, several species populations can be readily discerned based on physical appearance and cranial morphometrics (Frost et al, 2003), as well as striking ecological niche differences (Kamilar, 2006). Macaca spp. are temporally, geographically, phenotypically and behaviorally diverse which has led to questioning of its monophyly and scenarios of dispersal (Delson, 1975; Groves, 1989; Fooden & Albrecht, 1993; Purvis & Webster, 1999; Groves, 2000; Kohler et al, 2000; Abegg & Thierry, 2002; Thierry, 2007). However, macaques are the most conservative and primitive of the papionins (Delson, 1980) and generally lack deep

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facial fossae (Hill, 1974) as observed in species Lophocebus or Mandrillus. Additionally, the ante-orbital drop is less extreme, resembling instead species of Cercocebus and Parapapio (Freedman, 1957; Szalay & Delson, 1979). Similar to Papio, macaque species are also known to hybridize (Van Gelder, 1977; Bernstein & Gordon, 1980; Moore et al, 1999), a process which in one instance is proposed to have produced a new species (e.g. M. arctoides originated from a hybridization process involving M. assamensis/thibetana-like males with M. fascicularis-like females; Tosi et al, 2000 & 2003; Brandon-Jones et al, 2004; Morales & Melnick, 1998). For most of the 20th century species of Lophocebus were subsumed within Cercocebus because of similarities in body size, sympatry, suborbital fossae and attempts at taxonomic stability. However, molecular and morphological research in the late 1970’s (Cronin & Sarich, 1976; Groves, 1978) exposed the unlikelihood of this arrangement. Despite this, Lophocebus spp. remained in Cercocebus until the 1990s until further molecular and morphological research unequivocally demonstrated that the mangabeys of Cercocebus were more closely related to Mandrillus, while Lophocebus is more closely related to Papio and Theropithecus (Disotell, 1994; Fleagle & McGraw, 1999 & 2002; Harris, 2000; Page & Goodman, 2001; McGraw & Fleagle, 2006). In fact, mandrills and drills were once classified under Papio (e.g. Strasser & Delson, 1987) because of their extreme facial elongation, terrestrial lifestyle and sexual dimorphism. More recently, Gilbert’s (2007) reanalysis of the Plio-Pleistocene fossil papionins from southern Africa led to the erection of a new genus, Procercocebus, which is most likely an ancestor of the Cercocebus-Mandrillus clade (see also Jolly, 2007).

4.2 Descriptive Statistics and Univariate Analyses for cranial variables: Tables 4.7 lists the results of the pooled sex descriptive statistics for the thirty-six measurements collected. Due to the extreme sexual dimorphism between adult males and females some variables exhibit very high coefficients of variation, >15-20%, which is appropriate for samples that include more than one species as well as taking into consideration sexual dimorphism (Simpson et al, 1960; Cope & Lacy, 1995). Some of these include sagittal length of the nasal bones (nas-rhi), maximum width of the nasal aperture (maxnawi), palatal height (palhei) or inferior breadth of the nasal bones (inbrnabo).

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Table 4.7: Papionini pooled sex. g-o pros-o biast bipor biaur bizygo bifmo bifmt bizs bizi zs-zi zs-zygi Cercoc. (n=19) Mean 87.4 123.2 49.4 63.5 68.0 82.4 56.0 62.6 33.1 59.9 24.6 40.1 Median 86.0 125.2 49.9 62.4 67.0 81.8 55.1 60.5 33.1 59.3 24.3 39.5 Maximum 97.0 137.0 55.7 73.0 77.6 97.0 65.9 74.1 40.2 68.0 27.9 48.0 Minimum 79.4 108.0 42.0 56.0 61.4 70.0 50.9 55.0 23.1 52.8 21.1 30.7 Range 17.6 29.0 13.7 17.0 16.2 27.0 15.0 19.1 17.1 15.2 6.8 17.3 SD 5.9 10.4 3.6 5.3 4.4 7.6 4.3 5.6 4.8 4.7 1.8 4.6 CV 6.7 8.4 7.4 8.4 6.5 9.2 7.6 8.9 14.5 7.8 7.5 11.5

Mand. (n=22) Mean 103.1 177.2 60.9 77.1 81.6 102.0 63.9 76.5 37.5 72.9 30.2 45.8 Median 97.5 159.8 58.3 69.9 75.5 94.0 63.1 70.9 36.8 69.0 28.0 41.8 Maximum 130.0 240.0 84.0 99.4 99.4 139.0 79.2 116.3 46.1 101.6 47.1 65.0 Minimum 88.6 136.9 51.0 63.1 71.3 82.8 55.6 62.4 31.3 61.0 20.4 34.3 Range 41.4 103.1 33.0 36.2 28.1 56.2 23.7 53.9 14.8 40.6 26.7 30.7 SD 13.0 35.8 8.3 12.5 10.0 19.3 6.7 14.0 3.7 11.4 8.0 9.6 CV 12.6 20.2 13.7 16.2 12.3 18.9 10.5 18.3 9.8 15.7 26.6 21.0

Macaca (n=34) Mean 87.7 122.5 51.3 60.4 67.3 82.1 53.8 63.5 27.5 55.9 25.8 43.1 Median 89.0 121.8 50.9 60.0 66.8 80.5 53.5 64.0 27.6 56.3 25.8 42.7 Maximum 100.0 149.0 63.0 73.0 76.0 99.0 63.0 77.0 40.3 71.0 33.5 55.0 Minimum 73.7 103.0 39.2 47.0 57.0 63.0 46.0 50.3 18.9 44.2 18.3 29.7 Range 26.3 46.1 23.9 26.0 19.0 36.0 17.0 26.7 21.4 26.8 15.2 25.3 SD 6.6 11.3 6.5 6.1 5.3 9.2 4.6 6.2 4.3 6.3 3.8 5.6 CV 7.6 9.2 12.7 10.1 7.8 11.2 8.5 9.7 15.5 11.2 14.5 13.0

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g-o pros-o biast bipor biaur bizygo bifmo bifmt bizs bizi zs-zi zs-zygi Lopho. (n=23) Mean 84.9 117.9 50.8 60.3 67.2 77.1 52.0 59.3 34.3 56.7 23.1 37.4 Median 85.0 114.0 49.8 59.0 67.0 76.0 51.7 58.4 33.7 56.6 23.3 36.3 Maximum 94.0 137.0 60.0 70.0 75.0 90.0 57.0 69.1 44.9 64.7 28.0 43.7 Minimum 79.0 105.0 46.2 51.1 61.0 67.0 48.2 53.5 28.0 50.8 18.5 32.8 Range 15.0 32.0 13.8 18.9 14.0 23.0 8.8 15.6 16.9 13.9 9.6 10.9 SD 4.0 9.2 3.4 4.4 3.5 6.0 2.7 4.3 3.8 3.8 2.7 3.2 CV 4.7 7.8 6.7 7.4 5.2 7.8 5.1 7.3 11.0 6.8 11.6 8.4

Papio (n=31) Mean 111.2 194.3 68.8 84.3 89.5 111.0 69.8 80.0 37.6 75.7 37.0 55.3 Median 111.1 202.1 68.1 84.8 89.6 109.5 70.9 79.2 38.5 77.2 36.1 56.5 Maximum 125.0 235.8 84.0 98.9 100.4 136.5 78.9 94.1 44.5 87.1 45.5 66.6 Minimum 98.0 143.0 52.0 65.0 79.0 85.0 60.6 67.1 30.8 56.7 29.2 39.9 Range 27.0 92.8 32.0 33.9 21.4 51.5 18.3 27.0 13.7 30.4 16.3 26.8 SD 7.1 24.6 6.9 8.6 6.2 12.5 5.2 7.8 3.9 7.1 5.2 6.6 CV 6.4 12.7 10.0 10.2 6.9 11.3 7.4 9.7 10.4 9.4 13.9 11.9

Thero. (n=14) Mean 102.5 151.7 57.2 72.3 77.9 103.6 57.9 64.7 35.4 73.8 32.6 54.8 Median 103.0 152.2 55.6 70.2 77.2 102.2 56.8 64.9 34.6 74.5 31.8 53.4 Maximum 112.0 170.0 68.0 86.0 87.0 118.0 69.5 73.9 46.7 83.4 37.5 63.2 Minimum 96.9 136.9 51.3 64.0 68.6 91.6 49.4 56.6 27.8 64.5 29.1 47.9 Range 15.1 33.1 16.7 22.0 18.4 26.5 20.1 17.3 18.9 19.0 8.4 15.3 SD 4.6 10.7 4.8 7.0 6.3 9.3 5.1 5.4 5.1 5.6 2.6 5.8 CV 4.5 7.0 8.5 9.6 8.1 9.0 8.8 8.3 14.4 7.5 7.9 10.6

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Table 4.7 continued: Papionini pooled sex. bas-pros bas-nas bas-br nas-pros nas-br br-lam lam-opn nas-rhi inbrnabo rhi-ns maxnawi nas-ns Cercoc. (n=19) Mean 86.8 66.3 56.9 52.2 53.3 39.8 35.8 27.2 9.0 21.7 13.0 47.3 Median 86.8 66.2 56.8 52.8 52.0 40.2 36.3 27.0 9.1 21.5 13.0 47.8 Maximum 98.0 74.0 63.0 62.5 62.9 44.6 42.5 32.7 12.8 27.0 16.7 58.8 Minimum 73.8 59.9 52.0 42.4 46.2 34.0 28.5 19.2 7.5 17.8 10.9 35.2 Range 24.2 14.1 11.1 20.2 16.7 10.6 14.0 13.5 5.3 9.3 5.8 23.6 SD 8.3 4.9 2.9 5.8 4.8 3.3 3.4 3.5 1.2 2.4 1.5 6.0 CV 9.6 7.3 5.0 11.1 9.0 8.3 9.4 13.0 13.5 11.1 11.4 12.6

Mand. (n=22) Mean 128.0 74.3 63.7 89.4 60.0 45.4 44.5 57.0 10.7 26.2 18.5 83.2 Median 116.8 72.3 62.5 77.6 58.5 43.7 41.6 50.1 9.9 24.9 17.2 71.7 Maximum 173.3 90.8 72.0 135.8 73.4 63.0 59.3 96.9 14.8 35.6 28.6 127.2 Minimum 97.3 62.9 57.4 63.0 51.2 35.7 35.9 37.4 8.2 19.6 13.1 60.5 Range 76.0 27.9 14.6 72.8 22.1 27.3 23.4 59.5 6.5 16.0 15.4 66.7 SD 27.2 9.2 4.3 23.1 5.9 7.4 7.2 16.5 2.1 4.6 3.9 21.5 CV 21.2 12.4 6.8 25.8 9.9 16.3 16.2 28.9 19.2 17.4 21.3 25.8

Macaca (n=34) Mean 86.1 64.8 52.6 56.6 55.5 38.3 29.3 30.9 9.7 19.9 14.3 50.5 Median 85.0 64.0 53.0 54.6 55.4 38.1 29.2 30.1 9.4 20.0 13.9 49.4 Maximum 105.0 76.0 58.0 76.8 66.1 45.8 35.3 47.0 13.4 25.9 18.5 70.3 Minimum 71.0 57.2 45.6 43.8 45.7 32.8 22.4 16.9 6.0 13.9 9.7 30.7 Range 34.0 18.8 12.4 33.1 20.5 13.0 13.0 30.1 7.4 11.9 8.8 39.6 SD 8.7 4.8 3.2 8.5 4.6 3.6 2.8 7.9 2.1 2.7 2.2 9.6 CV 10.1 7.5 6.1 14.9 8.3 9.4 9.4 25.4 21.9 13.5 15.5 19.0

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bas-pros bas-nas bas-br nas-pros nas-br br-lam lam-opn nas-rhi inbrnabo rhi-ns maxnawi nas-ns Lopho. (n=23) Mean 83.2 64.1 53.2 48.7 51.0 40.5 30.6 25.8 8.0 19.2 12.3 44.3 Median 80.0 62.7 53.0 46.5 50.6 40.0 29.9 25.3 8.0 18.6 12.2 43.5 Maximum 100.0 74.0 59.0 67.0 55.3 50.4 37.6 36.8 10.0 23.4 15.4 55.5 Minimum 74.0 59.0 48.1 37.5 47.3 36.5 23.4 18.2 5.8 15.7 8.6 33.3 Range 26.0 15.0 10.9 29.5 8.0 13.8 14.2 18.6 4.2 7.6 6.7 22.3 SD 8.2 4.4 3.0 7.3 2.2 3.1 3.4 4.4 1.0 2.2 1.5 6.2 CV 9.9 6.8 5.6 14.9 4.3 7.5 11.3 17.1 12.1 11.3 12.1 14.1

Papio (n=31) Mean 140.2 84.4 69.4 113.4 65.2 48.2 44.1 68.4 12.7 37.2 22.0 105.6 Median 150.7 85.0 69.5 115.4 64.9 48.3 43.9 66.9 12.1 35.7 20.9 105.2 Maximum 171.7 97.9 77.1 152.7 73.1 59.7 57.2 96.6 17.1 51.7 28.1 145.5 Minimum 102.0 71.0 59.5 86.3 57.8 32.9 33.7 39.2 9.7 26.1 16.8 66.1 Range 69.7 26.9 17.7 66.3 15.3 26.9 23.5 57.4 7.5 25.6 11.3 79.4 SD 19.9 7.5 4.0 19.0 3.8 4.8 6.0 13.8 2.2 7.2 2.9 20.3 CV 14.2 8.8 5.7 16.8 5.9 10.0 13.7 20.2 17.6 19.3 13.3 19.3

Thero. (n=14) Mean 110.0 77.6 64.5 73.7 63.3 41.7 38.2 39.1 10.7 28.6 15.9 65.3 Median 109.1 78.3 64.2 75.0 60.9 40.7 38.5 39.8 10.4 28.7 15.2 66.5 Maximum 127.0 85.0 71.0 84.0 72.4 54.4 46.0 47.4 13.8 34.0 19.7 75.4 Minimum 96.2 70.0 58.6 64.4 58.0 36.7 32.8 30.4 7.8 23.9 13.9 55.9 Range 30.8 15.0 12.4 19.6 14.4 17.7 13.2 17.1 6.0 10.1 5.8 19.5 SD 9.9 5.3 4.4 6.8 5.1 4.8 4.0 5.0 1.7 3.4 2.0 7.5 CV 9.0 6.9 6.8 9.3 8.0 11.6 10.4 12.7 16.1 11.7 12.5 11.5

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Table 4.7 concluded: Papionini pooled sex. maxalvlen biecm iv-pms i1i2 bicanin bicanex palhei biseptal bienm ol-sta ol-pms bien Cercoc. (n=19) Mean 50.9 39.7 22.3 15.2 19.6 32.6 7.7 24.0 21.1 48.0 34.8 43.6 Median 48.6 39.3 21.9 15.1 19.2 32.3 7.6 23.9 21.1 47.9 34.5 44.0 Maximum 61.0 44.9 26.2 18.4 26.1 39.2 10.0 29.0 24.6 58.0 41.1 50.0 Minimum 44.0 35.7 18.0 12.0 16.7 28.2 6.0 20.6 18.2 39.6 23.7 39.0 Range 17.0 9.2 8.3 6.4 9.4 11.0 4.0 8.4 6.4 18.4 17.5 11.0 SD 5.5 2.7 2.4 1.5 2.1 2.6 1.3 2.4 1.9 5.9 4.7 3.2 CV 10.8 6.9 10.9 10.0 10.5 8.0 16.4 10.0 8.8 12.2 13.6 7.2

Mand. (n=22) Mean 72.9 50.4 39.0 20.6 28.0 45.2 8.1 33.5 27.7 77.1 54.8 48.3 Median 68.9 48.3 36.2 19.7 27.1 41.4 7.9 31.8 27.0 68.4 49.7 47.5 Maximum 94.3 65.3 52.4 30.3 41.0 71.7 13.0 46.2 35.9 112.4 80.2 60.0 Minimum 56.2 42.1 27.4 12.5 18.2 30.6 5.1 21.5 21.5 55.1 38.5 40.7 Range 38.1 23.2 25.0 17.9 22.8 41.1 7.9 24.7 14.4 57.3 41.7 19.3 SD 12.9 6.6 8.2 5.0 6.4 12.0 2.1 7.6 4.1 19.4 13.9 5.8 CV 17.7 13.2 21.0 24.3 22.7 26.5 26.1 22.7 14.9 25.1 25.4 11.9

Macaca (n=34) Mean 49.4 39.8 20.3 13.7 18.7 32.2 8.8 23.1 21.9 46.3 33.2 39.6 Median 49.0 40.1 19.8 13.7 18.4 31.9 9.0 22.9 22.1 45.0 32.2 38.9 Maximum 60.0 48.0 29.4 17.1 25.1 41.9 13.5 30.3 28.4 61.2 45.3 50.0 Minimum 36.0 30.6 13.7 12.0 14.4 24.8 5.0 18.8 16.2 35.8 24.7 30.6 Range 24.0 17.4 15.7 5.1 10.8 17.1 8.5 11.6 12.2 25.4 20.6 19.4 SD 5.5 4.1 3.1 1.2 2.4 4.1 1.7 2.5 2.4 5.8 4.4 5.0 CV 11.1 10.3 15.4 8.9 12.6 12.8 19.7 10.7 11.0 12.5 13.2 12.5

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maxalvlen biecm iv-pms i1i2 bicanin bicanex palhei biseptal bienm ol-sta ol-pms bien Lopho. (n=23) Mean 47.1 36.5 22.5 15.2 19.2 31.5 7.0 24.3 20.1 44.2 34.0 42.3 Median 46.0 36.8 20.9 14.9 19.3 31.1 7.0 24.0 20.1 41.8 32.4 42.8 Maximum 56.0 40.6 29.3 18.0 23.0 39.6 9.8 30.6 23.6 55.2 45.9 46.1 Minimum 41.8 32.4 18.3 12.2 15.7 27.2 5.0 21.3 17.3 36.1 28.0 35.6 Range 14.2 8.2 11.0 5.8 7.3 12.4 4.8 9.3 6.4 19.1 17.8 10.6 SD 3.9 2.2 3.1 1.3 1.7 3.0 1.1 2.4 1.6 6.1 4.8 2.8 CV 8.4 5.9 13.8 8.8 8.9 9.5 16.4 9.8 8.0 13.9 14.2 6.6

Papio (n=31) Mean 78.3 53.4 38.3 21.6 26.7 47.6 11.8 34.1 29.9 82.8 60.8 55.2 Median 80.9 53.5 38.4 21.8 26.7 51.2 12.4 34.5 30.0 87.0 62.6 55.7 Maximum 94.8 64.7 51.8 26.5 35.8 63.6 15.2 40.6 38.6 110.3 82.0 63.8 Minimum 56.0 44.0 27.3 15.8 19.8 30.7 8.4 25.4 22.3 54.9 39.1 44.3 Range 38.8 20.7 24.6 10.7 16.0 32.9 6.7 15.3 16.3 55.4 42.8 19.5 SD 10.0 4.6 6.5 3.1 3.4 8.3 2.1 4.3 3.3 14.8 10.9 4.9 CV 12.8 8.6 17.0 14.2 12.9 17.4 18.1 12.6 11.0 17.9 17.9 8.8

Thero. (n=14) Mean 66.7 44.5 27.2 15.3 17.4 32.8 8.8 23.7 23.1 65.0 46.1 49.3 Median 65.3 44.9 26.6 15.4 17.3 31.9 8.3 23.3 22.9 65.8 47.2 47.1 Maximum 79.0 48.2 31.5 17.4 20.6 40.7 13.0 27.4 26.5 74.9 54.0 62.0 Minimum 54.7 41.3 24.3 12.1 14.6 24.6 5.2 18.4 19.3 53.4 38.7 42.6 Range 24.3 6.9 7.2 5.3 6.0 16.2 7.8 8.9 7.2 21.6 15.3 19.4 SD 8.0 2.5 2.4 1.7 1.6 5.4 2.0 2.6 2.2 7.8 5.2 6.0 CV 12.0 5.6 8.7 11.2 9.4 16.4 22.6 11.0 9.7 12.0 11.2 12.2

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Despite this, other features such as maxillo-alveolar breadth (biecm) or cranial vault length (g-o) possess much less variation. Examination of the box-plots for the Papionini reveals many interesting trends. Figures 4.2 to 4.15 are box-plots which demonstrate these patterns. Genera are arranged by generic mean body weight starting with the least, Lophocebus to the greatest, Papio; with Cercocebus, Macaca, Theropithecus and Mandrillus in between. The box-plots present two patterns. First, some cranial dimensions are intimately related to body size. Secondly, two groups emerge; the first has moderate facial elongation, Lophocebus, Cercocebus and Macaca, while the second, Theropithecus, Mandrillus and Papio, have an extremely elongated naso-dental snout (absolutely and relatively). In particular, maxillo-alveolar (biecm) and palatal (bienm) breadths and sagittal length of the nasal bones (nas-rhi) (Figures 4.6, 4.7 and 4.12) exemplify the first pattern, namely, the increase in body size and its affect on cranial dimensions. In addition, the second pattern, distinction between moderate and extreme facial elongation, can be seen in the superior facial length (bas-pros), bizygomatic breadth (bizygo), cranial vault length (g-o) and palatal length A (ol-sta) (Figures 4.4, 4.10, 4.11 and 4.13). Based on simple cranial dimensions it is perhaps easy to understand why for many years Lophocebus was subsumed within Cercocebus because they do overlap for many measurements. Cercocebus can be diagnosed from Lophocebus by much less concavity of the nasal bones (i.e. lacking a steep anteorbital drop; Groves, 1978; as seen in Papio or Theropithecus), large upper incisors and deep suborbital fossae, although the latter can be present in other papionins, e.g. Mandrillus and Theropithecus (McGraw & Fleagle, 2006). The latter two characteristics have been hypothesized to be adaptations for eating hard food objects (Fleagle, 1998; Singleton, 2004). Groves (1978) also discusses the inferior portion of the nasal bones in Lophocebus as being “upturned”, giving the skull a “snub-nosed” appearance. Furthermore, Cercocebus actually resembles fossil specimens of Parapapio from South Africa (Freedman, 1957). In fact, Delson et al (2000) place some Parapapio fossil material within Cercocebus, although with some reservations. Mandrillus is easily diagnosable from other papionins by the extreme development of inflated or swelled maxillary ridges lateral to the nasal bones (Hill, 1970; Elton & Morgan, 2006).

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Box Plots for cranial variables of the Papionini (L. – Lophocebus, n=23; Cc. – Cercocebus, n=19; Mc. – Macaca, n=34; Th. – Theropithecus, n=14; Mn. – Mandrillus, n=22 & Pp. – Papio, n=31; variables in alphabetical order by abbreviation): Figure 4.3 80 Figure 4.2 Pp. 100 Pp. Th. Mn. Mn. 90 70 Th. Cc. L. 80 Mc. 60 Mc. L. Cc. mm mm 70 50 60

1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: Cranial Height (bas-br) Papionini: Inferior Cranial Length (bas-nas)

Figure 4.4 Figure 4.5 200 90 Pp. 190 180 Mn. Pp. Mn. 170 80 160 Th. 150 70 Mc. 140 Th. L. 130 Cc. mm 60 120 mm Mc. 110 L. Cc. 100 50 90 80 40 70

1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: Superior Facial Length (bas-pros) Papionini: Biasterionic Breadth (biast)

Figure 4.6 Figure 4.7 Pp. 70 40 Mn. Pp. Mn.

60 Mc. 30 Th. Mc Th. L. Cc. 50 . mm Cc. mm L. 20 40

1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: Maxillo-alveolar Breadth (biecm) Papionini: Palatal Breadth (bienm)

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Figure 4.8 Mn. Pp. Figure 4.9 100 Mn. 100 90 Th. Pp. 90 Th. 80 Cc. Mc. 80 L. Mc 70 Cc. . mm mm 70 L. 60 60 50 50 40 1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: Biporionic Breadth (bipor) Papionini: Bizygomaxillare Inferior Breadth (bizi)

Figure 4.11 Figure 4.10 Mn. Mn. 140 Pp. 130 Pp. 130 120 Th 120 Th. . 110 110 Mc. Cc. Mc. Cc.

100 mm L. mm 100 L. 90 90 80 70 80 60 70 1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: Bizygomatic Breadth (bizygo) Papionini: Cranial Vault Length (g-o)

Figure 4.12 Mn. Pp. Figure 4.13 100 120 Mn. Pp. 90 110 80 100 70 90 60 80 Th. mm Mc. Th. mm 70 50 Mc. L L. Cc. 40 . Cc. 60 30 50 20 40 30 1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: Sagittal length of the nasal bones (nas-rhi) Papionini: Palatal Length A (ol-sta)

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Figure 4.14 70 Pp. 50 Figure 4.15 Mn. Th. Mn. Pp. 60 Mc. 40 Th. Mc. 50 Cc. L. 30 L. Cc. mm mm 40

20 30

1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: Max. Length of the Zygomatic (zs-zgyi) Papionini: Zygomatico-maxillary Suture Length (zs-zi)

More recently, Gilbert (2007) has created a new genus of the Cercocebus/Mandrillus clade for fossil material recovered from the paleo-cave deposits of South Africa, Procercocebus. However, interestingly, the maximum length of the zygomatic bone (Figure 4.14, zs-zgyi) revealed a difference between Mandrillus and Papio and Theropithecus, which is also revealed in relative cranial proportions (see below). However, the sample for Mandrillus is slightly unbalanced, with more females than males (n=22, 8, 12) and with more males this difference in medians may not occur with larger samples. Lastly, Theropithecus, which Strasser & Delson (1987) describe as extremely autapomorphic, is airorhynchous (but also observed in a hominoid, e.g. Pongo, see Shea, 1985) compared to the other papionins which are klinorhyncous (Vogel, 1966 & 1968).

4.2.1 Shapiro-Wilk results for cranial variables: Table 4.8 lists the results of subjecting several papionin cranial measurements to Shapiro-Wilk tests. Unlike the cercopithecins, there are many more cranial variables which are not normally distributed but instead are bimodal due to the sexual dimorphism these genera exhibit. However, all cranial dimensions for Cercocebus and Macaca are normally distributed. In addition, Lophocebus and Theropithecus each have only two variables not normally distributed. These include biasterionic breadth (biast) and palatal length a (ol-sta) for Lophocebus and palatal length A (ol-sta) and maximum length of the zygomatic

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Table 4.8: Shapiro-Wilk results for cranial variables of the Papionini; those in bold represent significant departures from normality (variables listed in alphabetical order by abbreviation).

L. Cc. Mc. Th. Mn. Pp. n=23 n=19 n=34 n=14 n=22 n=31 1. Bas-br W: 0.96 0.97 0.96 0.91 0.92 0.97 p(normal): 0.41 0.79 0.19 0.19 0.07 0.57 2. Bas-nas W: 0.92 0.92 0.97 0.93 0.91 0.97 p(normal): 0.06 0.13 0.49 0.27 0.04 0.64 3. Bas-pros W: 0.85 0.92 0.96 0.94 0.85 0.91 p(normal): 0.003 0.11 0.22 0.47 0.003 0.01 4. Biast W: 0.91 0.98 0.97 0.88 0.92 0.98 p(normal): 0.04 0.96 0.42 0.07 0.07 0.85 5. Biecm W: 0.98 0.92 0.98 0.89 0.91 0.98 p(normal): 0.95 0.11 0.88 0.09 0.06 0.74 6. Bienm W: 0.98 0.96 0.97 0.96 0.96 0.98 p(normal): 0.84 0.63 0.51 0.67 0.44 0.87 7. Bipor W: 0.95 0.93 0.99 0.92 0.81 0.95 p(normal): 0.32 0.19 0.92 0.2 0.0008 0.17 8. Bizi W: 0.96 0.94 0.98 0.97 0.88 0.96 p(normal): 0.43 0.26 0.78 0.87 0.01 0.28 9. Bizygo W: 0.94 0.96 0.98 0.91 0.84 0.98 p(normal): 0.17 0.61 0.65 0.17 0.002 0.72 10. G-o W: 0.96 0.93 0.97 0.91 0.83 0.97 p(normal): 0.53 0.15 0.58 0.17 0.002 0.52 11. Nas-rhi W: 0.95 0.97 0.97 0.93 0.89 0.96 p(normal): 0.37 0.85 0.44 0.32 0.02 0.3 12. Ol-sta W: 0.9 0.94 0.96 0.88 0.84 0.93 p(normal): 0.02 0.23 0.3 0.06 0.03 0.05 13. Zs-zgyi W: 0.95 0.97 0.99 0.84 0.86 0.97 p(normal): 0.34 0.84 0.99 0.02 0.004 0.6 14. Zs-zi W: 0.97 0.98 0.97 0.94 0.88 0.94 p(normal): 0.66 0.89 0.55 0.4 0.014 0.07

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(zs-zgyi) for Theropithecus. The Papio sample has three cranial variables that are not normally distributed; superior facial length (bas-pros), palatal length A (ol-sta) and the zygomatico-maxillary suture length (zs-zi). The sample for Mandrillus contains ten variables that are not normally distributed. Interestingly, most of these include measurements were normally distributed for the other papionin genera. Some of these include, inferior cranial length (bas-nas), biporionic (bipor), bizygomaxillare inferior (bizi) and bizygomatic (bizygo) breadths and cranial vault length (g-o). These non-normally distributed cranial dimensions highlight the great size disparity between males and females of Mandrillus, which is greater than the other papionin genera.

4.2.2 Kruskal-Wallis and Mann-Whitney results for cranial variables: Table 4.9 presents the results of applying Kruskal-Wallis to several cranial variables. The results reinforce observations made via examination of box-plots. Several cranial variables for the Lophocebus, Cercocebus and Macaca samples are not significantly different. Some of these include inferior (bas-nas) and superior (bas-pros) cranial lengths and biasterionic (biast) and biporionic (bipor) breadths. Due to their larger size, most variables for Theropithecus, Mandrillus and Papio are significantly different from Lophocebus, Cercocebus and Macaca. However, there are three exceptions. First, palatal breadth (bienm) between Macaca and Theropithecus is not significantly different. Secondly, the maximum length of the zygomatic (zs-zgyi) and the zygomatico-maxillary suture length (zs-zi) between Macaca and Mandrillus are not significantly different; although the latter is approaching significance (p-value, 0.07). Due to their large size, Papio is significantly different from all other papionins for most cranial dimensions. Two exceptions to this are the bizygomaxillare inferior (bizi) and bizygomatic (bizygo) breadths which overlap with those of Theropithecus and Mandrillus and are thus not significantly different.

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Table 4.9: Kruskal-Wallis results for cranial variables of the Papionini with Mann-Whitney pairwise comparisons (p (same)) (variables listed in alphabetical order by abbreviation).

L. Cc. Mc. Th. Mn. Pp. n=23 n=19 n=34 n=14 n=22 n=31 1. Bas-br 0 0.0004 0.71 8.13E-07 2.83E-08 4.74E-10 L. H: 112.60 0 0.00004 0.00004 7.33E-06 4.46E-09 Cc. Hc: 112.70 0 7.16E-08 5.69E-10 4.68E-12 Mc. p(same): 1.14E-22 0 0.70 0.001 Th. 0 0.00002 Mn. 0 Pp. 2. Bas-nas 0 0.15 0.58 1.79E-06 0.00003 7.39E-10 L. H: 87.87 0 0.29 0.00003 0.005 1.22E-08 Cc. Hc: 87.90 0 5.69E-07 0.00005 1.12E-11 Mc. p(same): 1.88E-17 0 0.19 0.005 Th. 0 0.0002 Mn. 0 Pp. 3. Bas-pros 0 0.23 0.11 6.92E-07 1.38E-08 4.74E-10 L. H: 107.10 0 0.63 4.07E-06 5.80E-08 4.20E-09 Cc. Hc: 107.10 0 7.62E-07 1.81E-09 5.63E-12 Mc. p(same): 1.66E-21 0 0.08 0.00004 Th. 0 0.06 Mn. 0 Pp. 4. Biast 0 0.39 0.84 0.00008 1.25E-06 1.03E-09 L. H: 87.64 0 0.32 0.00003 4.83E-07 6.78E-09 Cc. Hc: 87.65 0 0.009 0.00005 6.40E-11 Mc. p(same): 2.10E-17 0 0.12 0.00001 Th. 0 0.00007 Mn. 0 Pp. 5. Biecm 0 0.0006 0.001 5.00E-07 8.55E-09 4.74E-10 L. H: 105.80 0 0.69 0.00009 3.20E-07 6.02E-09 Cc. Hc: 105.80 0 0.0004 2.68E-08 1.02E-11 Mc. p(same): 3.10E-21 0 0.003 1.64E-06 Th. 0 0.04 Mn. 0 Pp. 6. Bienm 0 0.14 0.0002 0.0004 6.12E-08 6.62E-10 L. H: 92.67 0 0.17 0.01 9.48E-04 7.64E-09 Cc. Hc: 92.67 0 0.20 7.46E-07 4.12E-11 Mc. p(same): 1.85E-18 0 0.0007 8.85E-07 Th. 0 0.04 Mn. 0 Pp. 7. Bipor 0 0.06 0.98 0.00001 1.47E-07 8.25E-10 L. H: 92.88 0 0.09 0.001 0.00006 1.22E-08 Cc. Hc: 92.91 0 7.88E-06 1.26E-07 9.32E-12 Mc. p(same): 1.67E-18 0 0.29 0.0003 Th. 0 0.02 Mn. 0 Pp.

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L. Cc. Mc. Th. Mn. Pp. n=23 n=19 n=34 n=14 n=22 n=31 8. Bizi 0 0.02 0.60 5.89E-07 1.01E-07 1.43E-09 L. H: 94.84 0 0.02 5.76E-06 0.00005 1.18E-07 Cc. Hc: 94.84 0 2.47E-07 2.21E-08 6.98E-11 Mc. p(same): 6.45E-19 0 0.34 0.28 Th. 0 0.12 Mn. 0 Pp. 9. Bizygo 0 0.02 0.01 5.00E-07 2.12E-07 6.62E-10 L. H: 91.44 0 0.91 5.28E-06 0.0001 1.74E-08 Cc. Hc: 91.45 0 8.56E-07 0.00003 4.91E-11 Mc. p(same): 3.35E-18 0 0.23 0.50 Th. 0 0.02 Mn. 0 Pp. 10. G-o 0 0.21 0.05 5.00E-07 3.22E-08 4.74E-10 L. H: 101.90 0 0.75 1.67E-06 0.00001 4.20E-09 Cc. Hc: 102.00 0 1.93E-07 7.46E-07 5.13E-12 Mc. p(same): 2.05E-20 0 0.20 0.0002 Th. 0 0.002 Mn. 0 Pp. 11. Nas-rhi 0 0.17 0.008 1.79E-06 9.86E-09 4.74E-10 L. H: 108.6 0 0.08 2.86E-06 5.01E-08 4.20E-09 Cc. Hc: 108.6 0 0.001 6.78E-09 8.13E-12 Mc. p(same): 8.03E-22 0 0.00006 2.80E-07 Th. 0 0.006 Mn. 0 Pp. 12. Ol-sta 0 0.04 0.10 9.54E-07 1.13E-08 5.30E-10 L. H: 106.60 0.30 0.00001 8.97E-08 6.02E-09 Cc. Hc: 106.60 3.54E-07 7.03E-10 6.46E-12 Mc. p(same): 2.17E-21 0.08 0.0009 Th. 0.10 Mn. 0 Pp. 13. Zs-zgyi 0 0.04 0.00003 5.00E-07 0.0003 9.21E-10 L. H: 80.02 0 0.06 1.67E-06 0.08 3.09E-08 Cc. Hc: 80.02 0 3.51E-06 0.85 5.08E-09 Mc. p(same): 8.31E-16 0 0.002 0.65 Th. 0 0.0003 Mn. 0 Pp. 14. Zs-zi 0 0.05 0.005 5.00E-07 0.0006 4.74E-10 L. H: 84.41 0 0.13 1.39E-06 0.006 4.20E-09 Cc. Hc: 84.41 0 1.28E-06 0.07 9.07E-11 Mc. p(same): 1.00E-16 0 0.04 0.01 Th. 0 0.0004 Mn. 0 Pp.

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4.2.3 One-way ANOVA results for cranial variables: Table 4.10 provides the results of subjecting the selected papionin cranial variables to one-way ANOVA. Mandrillus was excluded from a number of analyses because of non- normal distributions. Palatal length A (ol-sta) was not included in the ANOVA analysis because four of the six genera had non-normally distributed samples. For seven variables the majority of variation (>70%) is produced between groups. The remaining variables generate variation between groups ranging from >50% to <69%. Again, results reveal the similar sizes between Lophocebus, Cercocebus and Macaca. In fact, nine variables between these genera were not significantly different. Some of these include inferior (bas-nas) and superior (bas-pros) cranial lengths, and a number of breadth measurements (biecm, bienm, bipor and bizi). Very similar to results for Kruskal-Wallis, Theropithecus, Mandrillus (when included) and Papio are again significantly different from the other papionins. However, there are some exceptions to this. The palatal breadth (bienm) for Cercocebus, Macaca and Theropithecus were not significantly different. There was not a significant difference in cranial height (bas-br) or biasterionic breadth (biast) between Theropithecus and Mandrillus. Lastly, some cranial variables for Mandrillus and Papio overlapped considerably and as such are not significantly different. These include maxillo-alveolar, palatal, inferior bizygomaxillare and bizygomatic breadths (biecm, bienm, bizi and bizygo); although the latter approaches significance (p-value, 0.08).

4.2.4 Summary for cranial variables: In summary, linear dimensions of papionin crania revealed two groups, as mentioned previously. The first, small to medium with moderate facial elongation; and the second, medium to large with extreme facial elongation. Many cranial dimensions increase with body size.

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Table 4.10: One-Way ANOVA results for cranial variables of the Papionini (variables listed in alphabetical order by abbreviation). Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 1. Bas-br p(same): L. Cc. Mc. Th. Mn. Pp. Between F: 107.5 0 .007 0.99 0.00002 0.00002 0.00002 groups: 6553.81 79.42 5 1310.76 105.8 2.89E-45 0.10 df: 55.24 0 .0008 0.00002 0.00002 0.00002 Within groups: 1697.99 20.58 137 12.39 p: 3.49E-27 0 0.00002 0.00002 0.00002 Total: 8251.81 142 0 0.98 0.00003 0 0.00002 0 2. Bas-nas L. Cc. Mc. Th. Pp. Between F: 54.76 0 0.68 0.99 0.0001 0.0001 groups: 8972.32 71.00 4 2243.08 71.01 2.76E-30 0.007 df: 49.24 0 0.90 0.0001 0.0001 Within groups: 3664.12 29.00 116 31.59 p: 1.58E-17 0 0.0001 0.0001 Total: 12636.4 120 0 0.001 ex. Mn. 0 3. Bas-pros L. Cc. Mc. Th. Between F: 25.15 0 0.58 0.73 0.0001 groups: 7294.66 52.83 3 2431.55 32.1 5.10E-14 0.71 df: 38.96 0 0.99 0.0001 Within groups: 6514.17 47.17 86 75.75 p: 3.22E-09 0 0.0001 Total: 13808.6 89 ex. Mn. & Pp. 0 Cc. Mc. Th. Mn. Pp. 4. Biast 0 0.84 0.0006 0.0001 0.0001 Between F: 46.85 0 0.02 0.0002 0.0001 groups: 6653.86 60.59 4 1663.46 44.2 2.01E-22 0.002 df: 50.8 0 0.50 0.0001 Within groups: 4328.44 39.41 115 37.64 p: 1.89E-16 0 0.0002 Total: 10982.3 119 ex. L. 0

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Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. F comparisons sqrs: Var.: df: square: F: p(same): of Var. test: p(same): 5. Biecm p(same): L. Cc. Mc. Th. Mn. Pp. Between F: 77.16 0 0.14 0.11 0.00002 0.00002 0.00002 L. groups: 5898.58 70.78 5 1179.72 66.39 6.53E-35 .000004 df: 57.8 0 1 0.002 0.00002 0.00002 Cc. Within groups: 2434.57 29.22 137 17.77 p: 2.68E-24 0 0.004 0.00002 0.00002 Mc. Total: 8333.15 142 0 0.00006 0.00002 Th. 0 0.16 Mn. 0 Pp. 6. Bienm L. Cc. Mc. Th. Mn. Pp. Between F: 48.76 0 0.86 0.26 0.006 0.00002 0.00002 L. groups: 2032.19 65.76 5 406.44 52.62 3.09E-30 0.001 df: 56.52 0 0.92 0.17 0.00002 0.00002 Cc. Within groups: 1058.17 34.24 137 7.72 p: 2.83E-19 0 0.75 0.00002 0.00002 Mc. Total: 3090.36 142 0 0.00002 0.00002 Th. 0 0.10 Mn. 0 Pp. 7. Bipor L. Cc. Mc. Th. Pp. Between F: 52.28 0 0.52 1 0.0001 0.0001 L. groups: 12179.3 70.76 4 3044.83 70.2 4.43E-30 0.0004 df: 49.13 0 0.53 0.0003 0.0001 Cc. Within groups: 5031.68 29.24 116 43.38 p: 4.14E-17 0 0.0001 0.0001 Mc. Total: 17211 120 0 0.0001 Th. ex. Mn. 0 Pp. 8. Bizi L. Cc. Mc. Th. Pp. Between F: 61.68 0 0.35 0.99 0.0001 0.0001 L. groups: 9314.67 70.33 4 2328.67 68.73 1.04E-29 0.06 df: 49.94 0 0.17 0.0001 0.0001 Cc. Within groups: 3930.15 29.67 116 33.88 p: 1.04E-18 0 0.0001 0.0001 Mc. Total: 13244.8 120 0 0.83 Th. ex. Mn. 0 Pp.

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Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. F comparisons sqrs: Var.: df: square: F: p(same): of Var. test: p(same): 9. Bizygo p(same): L. Cc. Mc. Th. Pp. Between F: 56.4 0 0.36 0.41 0.0001 0.0001 L. groups: 23235 68.96 4 5808.76 64.43 1.40E-28 0.002 df: 49.48 0 1 0.0001 0.0001 Cc. Within groups: 10458.7 31.04 116 90.16 p: 7.89E-18 0 0.0001 0.0001 Mc. Total: 33693.7 120 0 0.08 Th. ex. Mn. 0 Pp. 10. G-o L. Cc. Mc. Th. Pp. Between F: 95.01 0 0.64 0.54 0.0001 0.0001 L. groups: 14208.3 77.09 4 3552.08 97.58 3.49E-36 0.02 df: 50.66 0 0.99 0.0001 0.0001 Cc. Within groups: 4222.69 22.91 116 34.4 p: 6.65E-23 0 0.0001 0.0001 Mc. Total: 18431 120 0 0.0001 Th. ex. Mn. 0 Pp. 11. Nas-rhi L. Cc. Mc. Th. Pp. Between F: 76.3 0 0.98 0.30 0.01 0.0001 L. groups: 35561.7 80.23 4 8890.42 117.7 6.91E-40 1.39E-11 df: 52.12 0 0.62 0.0002 0.0001 Cc. Within groups: 8760.32 19.77 116 75.52 p: 3.80E-21 0 0.02 0.0001 Mc. Total: 44322 120 0 0.0001 Th. ex. Mn. 0 Pp. 12. Zs-zgyi L. Cc. Mc. Pp. Between F: 58.75 0 0.26 0.01 0.0001 L. groups: 5206.26 63.94 3 1735.42 60.87 1.01E-22 0.02 df: 53.34 0 0.20 0.0001 Cc. Within groups: 2936.62 36.06 103 28.51 p: 6.46E-17 0 0.0001 Mc. Total: 8142.88 106 ex. Mn. & Th. 0 Pp. 13. Zs-zi L. Cc. Mc. Th. Between F: 41.46 0 0.34 0.02 0.0002 L. groups: 842.66 52.26 3 280.89 31.38 8.51E-14 0.03 df: 41.74 0 0.58 0.0002 Cc. Within groups: 769.88 47.74 86 p: 1.39E-12 0 0.0002 Mc. Total: 1612.53 89 ex. Mn. & Pp. 0 Th.

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4.3 Cranial Indices: To correct for absolute size differences between genera several cranial indices were examined. Table 4.10 presents the mean and standard deviation (SD) for several cranial indices of the Papionini. The standard deviation for the samples is reduced compared to raw pooled sex data. Figures 4.16 to 4.26 are box-plots which illustrate scaling trends and some distinguishing indices between genera. Two indices in particular reveal shared cranial proportions between closely related genera as revealed by recent genetic analyses which had previously been incorrectly grouped because of supposed morphological features (previous incorrect groupings, Cercocebus (included species of Lophocebus - e.g. Hill, 1974; and ‘close’ relationship between Mandrillus-Papio - e.g. Szalay & Delson, 1979; correct phylogenetic relationships Cercocebus-Mandrillus and Lophocebus-Papio- Theropithecus; Gilbert & Rossie, 2007). These include biasterionic breadth relative to the occipital sagittal chord (biast/lam-opn) and the occipital sagittal chord in proportion to the cranial vault length (lam-opn/g-o) (Figures 4.16 and 4.24). Two interesting distinctions of Theropithecus are the relative proportion of the external bicanine breadth (bicanex) to palatal length (ol-sta; Figure 4.17) and the relative proportion of superior facial breadth (bifmt) to the lower facial breadth (bizi; Figure 4.20).

4.3.1 Shapiro-Wilk results for cranial indices: Table 4.12 lists the results of applying Shapiro-Wilk to several cranial indices. Only two indices were found to be non-normally distributed. The first is external bicanine breadth (bicanex) relative to palatal length A (ol-sta). This index was non-normal for Cercocebus and Mandrillus. The second is palatal length A (ol-sta) in relation to cranial height (bas-br). This index was non-normal for Lophocebus and Mandrillus. The results for all other indices possess a normal distribution. An important feature of the results for these papionin indices is that there are fewer variables with non-normal distribution due to absolute size differences between sexually dimorphic adult males and females. Thus, despite large differences due to absolute size and sexual dimorphism, males and females also share cranial developmental patterns and/or trajectories.

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Table 4.10: Generic means for cranial indices (%; indices listed in alphabetical order by abbreviation). L. Cc. Mc. Th. Mn. Pp. 1. bas-br/bas-nas 83.18 86.00 81.92 83.10 86.31 82.47 SD 3.24 3.58 5.27 2.50 5.69 4.93 2. bas-br/bas-pros 64.27 65.98 61.57 58.77 51.19 50.23 SD 3.76 4.44 5.53 2.64 7.42 5.97 3. bas-br/biast 105.13 115.88 103.72 113.02 105.49 101.64 SD 6.78 10.41 10.51 6.59 8.61 8.00 4. bas-br/biaur 79.22 83.95 78.38 82.90 78.58 77.67 SD 3.03 4.40 4.39 2.83 5.36 4.17 5. bas-br/g-o 62.1 65.3 60.1 62.9 62.0 62.4 SD 2.7 2.7 2.7 2.2 4.2 3.0 6. bas-br/bizygo 69.27 69.45 64.64 62.37 63.71 63.02 SD 3.37 4.53 5.54 2.68 7.80 5.81 7. bas-br/pros-o 45.27 46.38 43.50 42.50 36.81 36.10 SD 2.06 2.50 3.59 1.02 4.83 3.64 8. bas-nas/bas-pros 77.25 76.66 75.16 70.76 59.07 60.79 SD 3.00 2.52 4.75 3.28 5.47 4.70 9. bas-pros/pros-o 70.51 70.38 70.79 72.39 72.07 72.01 SD 1.83 1.98 3.66 1.97 2.11 1.90 10. biast/biaur 75.56 72.78 76.11 73.53 74.68 76.84 SD 3.99 4.93 5.90 4.00 4.20 4.97 11. biast/g-o 59.2 57.4 58.4 55.7 58.9 61.8 SD 3.9 2.7 5.1 2.9 3.2 4.0 12. biast/lam-opn 167.6 139.0 175.3 150.7 135.4 157.3 SD 18.5 11.1 16.0 14.5 10.7 15.0 13. biaur/bizygo 87.45 82.74 82.46 75.25 80.85 81.09 SD 2.93 3.74 4.19 2.55 5.48 5.58 14. biaur/pros-o 57.17 55.32 55.48 51.32 46.70 46.48 SD 2.30 2.94 3.20 1.87 3.82 4.08 15. bicanex/bas-pros 37.83 37.69 37.39 29.68 35.03 33.82 SD 1.84 2.86 2.95 3.09 2.51 2.07 16. bicanex/nas-pros 65.41 62.93 59.18 44.27 50.77 42.13 SD 6.43 6.73 9.73 4.17 5.47 4.65 17. bicanex/ol-sta 71.79 68.33 70.49 50.20 58.51 57.57 SD 5.40 5.65 7.04 3.59 3.07 3.32 18. biecm/bas-br 68.65 69.70 75.57 69.04 78.95 77.09 SD 3.30 4.24 5.82 2.10 6.38 5.60 19. biecm/bas-nas 57.06 59.90 61.82 57.37 67.91 63.43 SD 2.73 3.79 5.17 2.12 4.16 3.95 20. biecm/biast 72.07 80.57 78.10 78.02 83.00 77.96 SD 4.46 6.43 7.43 4.92 6.38 5.24 21. biecm/bicanex 116.47 122.25 124.79 138.12 115.13 114.37 SD 6.01 9.92 14.52 16.57 15.57 13.13 22. biecm/bifmt 61.72 63.56 62.76 68.91 67.21 66.93 SD 2.74 4.25 3.95 3.72 3.64 3.39 23. biecm/bizi 64.53 66.28 71.40 60.42 69.39 70.80 SD 2.34 3.16 5.75 3.85 3.56 3.86

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L. Cc. Mc. Th. Mn. Pp. 24. biecm/bizygo 47.48 48.33 48.63 43.07 49.88 48.32 SD 1.86 3.32 3.26 2.43 3.18 2.52 25. biecm/g-o 43.05 45.45 45.36 43.39 48.92 48.05 SD 2.14 2.83 3.03 1.62 2.89 2.80 26. biecm/ol-sta 83.60 83.29 87.39 68.95 67.15 65.82 SD 7.13 7.38 8.43 6.08 7.98 8.23 27. biecm/pros-o 31.04 32.30 32.75 29.35 28.82 27.70 SD 1.24 2.22 2.20 1.16 2.28 2.04 28. biecm/zs-zgyi 98.05 99.61 92.83 81.58 112.00 97.18 SD 5.74 9.20 7.31 5.21 11.80 6.93 29. bien/bas-br 79.50 76.56 75.24 76.45 75.78 79.66 SD 4.53 4.38 7.68 7.63 6.85 6.01 30. bien/bas-nas 66.07 65.78 61.48 63.46 65.35 65.57 SD 3.62 3.69 5.92 5.82 6.66 4.59 31. bien/bas-pros 51.06 50.43 46.10 44.90 38.67 39.83 SD 3.71 3.33 4.16 4.51 5.72 3.90 32. bien/biast 83.46 88.61 77.48 86.31 79.73 80.56 SD 5.71 8.18 5.92 8.96 7.36 5.73 33. bifmt/biaur 88.14 92.11 94.24 83.12 92.04 89.39 SD 4.23 5.23 4.51 1.89 4.33 5.51 34. bifmt/bizi 104.65 104.57 114.01 87.77 103.47 105.94 SD 3.51 6.33 9.37 4.96 6.75 6.27 35. bifmt/bizygo 77.01 76.10 77.56 62.54 74.28 72.31 SD 3.11 3.32 3.72 2.29 4.22 4.02 36. bifmt/g-o 69.85 71.58 72.34 63.07 72.89 71.91 SD 4.11 2.77 3.42 2.78 4.43 4.64 37. bifmt/nas-pros 123.35 120.91 117.25 88.04 86.56 71.77 SD 12.39 12.79 19.48 5.25 11.34 9.09 38. bifmt/pros-o 50.34 50.86 52.24 42.65 42.89 41.44 SD 2.09 2.38 3.01 1.81 2.85 3.21 39. bipor/g-o 70.4 72.6 68.8 70.5 74.3 75.7 SD 4.8 3.8 3.6 4.0 4.1 5.8 40. bizi/bas-br 106.44 105.33 106.13 114.62 114.12 109.09 SD 5.05 7.26 7.89 6.83 11.66 8.32 41. bizi/biast 111.73 121.59 109.78 129.47 119.91 110.30 SD 6.76 8.36 10.96 9.77 11.35 7.91 42. bizi/bizygo 73.60 72.96 68.39 71.43 71.93 68.37 SD 2.29 4.41 5.47 4.12 3.82 3.84 43. bizi/nas-pros 118.0 115.8 102.9 100.5 84.0 67.9 SD 9.1 11.6 15.2 6.7 12.0 8.9 44. bizs/zs-zgyi 92.08 82.90 64.82 64.73 84.76 68.55 SD 10.91 11.97 13.00 7.45 16.71 9.26 45. bizs/zs-zi 150.32 134.56 109.20 109.12 130.12 98.93 SD 22.75 19.97 26.80 17.16 32.77 22.58 46. bizs/bizi 60.41 55.13 49.54 47.91 52.28 48.05 SD 4.38 6.50 8.42 5.24 7.49 10.87

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L. Cc. Mc. Th. Mn. Pp. 47. bizygo/pros-o 65.41 66.89 67.42 68.21 57.81 57.33 SD 2.40 2.94 3.65 2.09 3.48 3.23 48. br-lam/g-o 46.8 45.6 43.7 40.7 43.9 43.8 SD 3.4 3.0 3.2 4.2 3.6 3.2 49. g-o/biaur 126.35 128.74 130.38 131.98 126.48 124.53 SD 5.25 6.52 5.39 5.43 5.96 6.82 50. g-o/bizygo 110.51 106.44 107.43 99.33 102.28 100.91 SD 6.35 5.56 6.88 5.53 8.55 7.87 51. g-o/pros-o 72.22 71.10 72.30 67.68 59.01 57.78 SD 3.86 3.03 4.21 2.71 4.76 4.78 52. iv-pms/bicanex 71.84 68.60 63.36 84.55 87.44 81.11 SD 7.01 6.86 7.69 12.32 6.95 8.25 53. iv-pms/ol-sta 51.44 46.60 44.28 42.14 51.04 46.53 SD 5.20 3.16 3.69 4.17 3.21 3.89 54. lam-opn/bas-br 57.43 62.94 55.64 59.19 69.63 63.57 SD 5.30 5.46 3.98 3.89 7.22 7.63 55. lam-opn/bas-pros 36.88 41.54 34.18 34.82 35.30 31.75 SD 3.78 4.60 3.18 3.20 3.68 4.05 56. lam-opn/biaur 45.53 52.84 43.55 49.03 54.50 49.30 SD 4.91 5.28 3.18 3.08 4.34 5.84 57. lam-opn/g-o 35.6 41.5 33.4 37.2 43.7 39.6 SD 3.0 3.2 2.6 2.9 3.0 4.3 58. maxnawi/nas-rhi 48.4 48.7 48.3 40.8 33.4 32.8 SD 6.3 9.1 10.8 4.0 5.2 4.6 59. nas-br/g-o 59.5 61.1 63.3 62.4 58.3 58.7 SD 1.8 4.3 3.4 2.8 3.5 3.2 60. nas-pros/biaur 72.08 76.82 82.38 94.67 108.13 126.25 SD 7.51 7.47 13.13 5.08 15.47 15.90 61. nas-pros/bien 114.74 120.01 140.89 150.63 181.67 204.90 SD 12.28 13.15 24.66 15.26 30.62 25.83 62. nas-pros/g-o 57.26 59.75 63.30 71.86 85.65 101.58 SD 7.58 5.88 10.42 5.08 12.70 13.24 63. nas-rhi/nas-pros 53.2 52.3 55.1 53.0 63.4 60.1 SD 5.7 5.5 5.2 3.3 3.4 4.3 64. ol-sta/bas-br 82.75 84.16 87.19 100.67 119.98 119.03 SD 8.65 7.61 10.18 6.95 22.54 17.94 65. ol-sta/bas-nas 68.69 72.21 71.04 83.62 102.51 97.57 SD 6.16 5.19 5.68 5.62 13.67 11.61 66. ol-sta/bas-pros 52.91 55.30 53.25 59.04 59.86 58.81 SD 3.17 3.52 3.61 2.92 2.48 2.84 67. ol-sta/bizygo 57.17 58.18 55.96 62.64 74.97 74.20 SD 5.05 2.97 4.46 2.54 7.13 7.12 68. palhei/ol-sta 15.99 16.13 18.98 13.54 10.72 14.28 SD 2.62 2.32 3.04 2.42 2.13 1.44 69. zs-zgyi/bizi 65.87 67.02 77.23 74.24 62.53 73.10 SD 3.55 6.32 7.28 5.28 6.51 5.08

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L. Cc. Mc. Th. Mn. Pp. 70. zs-zgyi/bizygo 48.48 48.74 52.52 52.84 44.85 49.84 SD 2.34 3.65 3.10 1.71 3.85 2.41 71. zs-zgyi/ol-sta 85.22 83.86 94.31 84.48 60.17 67.66 SD 7.03 6.08 7.81 4.23 6.30 6.15 72. zs-zi/bizi 40.68 41.26 46.14 44.24 40.94 48.81 SD 3.83 3.51 3.97 3.06 6.29 4.26 73. zs-zi/bizygo 29.94 30.03 31.50 31.56 29.33 33.31 SD 2.75 2.20 3.18 2.20 3.91 2.79 74. zs-zi/zs-zgyi 61.70 61.96 59.99 59.74 65.61 66.88 SD 4.17 6.90 5.16 4.10 8.66 5.11

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Box Plots for cranial indices of the Papionini (L. – Lophocebus, n=23; Cc. – Cercocebus, n=19; Mc. – Macaca, n=34; Th. – Theropithecus, n=14; Mn. – Mandrillus, n=22 & Pp. – Papio, n=31; indices in alphabetical order by abbreviation):

Figure 4.16 Figure 4.17 Mc. 90 210 L. Mc. 200 L. Pp 80 190 . Cc. Th. 180 Mn. 170 Cc. 70 Pp. Mn. % 160 % 150 60 Th. 140 130 50 120 110 1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: biast/lam-opn x 100 Papionini: bicanex/ol-sta x 100 Figure 4.18 Figure 4.19 100 Mc. Mn. Cc. Mn. Pp. 130 Cc. 90 120 Th. L. Pp. 110 Mc. % % L. 80 100 Th. 90 70 80 70 1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: biecm/biast x 100 Papionini: biecm/zs-zgyi x 100

Figure 4.20 Figure 4.21 Mc. 180 130 170 Mc. Pp. 160 L. 120 Cc. Mn. 150 Cc. L. 140 110 130 120 % % Mn. 100 Th. 110 Th. 100 Pp. 90 90 80 80 70 60

1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: bifmt/bizi x 100 Papionini: bifmt/nas-pros x 100

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Figure 4.22 Figure 4.23 150 Th. L. Cc. Mc. 140 140 130 Cc. Mc. Mn. Pp. 120 Th. 130 L. 110 Mn. 120 100 Pp % . 110 % 90 100 80 70 90 60 80 50 1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: bizi/biast x 100 Papionini: bizi/nas-pros x 100

Figure 4.24 Figure 4.25 60 Mn. 160 Pp. Mn. Pp. 150 50 Cc. 140 L. Th. 130 Mc. 120 Mc. Th. % 40 % 110 L. Cc. 100 90 30 80 70 60 1 2 3 4 5 6 7 1 2 3 4 5 6 7 Papionini: lam-opn/g-o x 100 Papionini: ol-sta/bas-br x 100

Figure 4.26 Mc. 110 L. 100 Cc. Th. 90 Pp.

% 80 Mn. 70 60 50

1 2 3 4 5 6 7 Papionini: zs-zgyi/ol-sta x 100

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Table 4.12: Shapiro-Wilk results for cranial indices of the Papionini; those in bold represent significant departures from normality (indices listed in alphabetical order by abbreviation).

L. Cc. Mc. Th. Mn. Pp. n=23 n=19 n=34 n=14 n=22 n=31 1. Biast/lam-opn W: 0.97 0.94 0.97 0.96 0.95 0.97 p(normal): 0.75 0.26 0.36 0.68 0.36 0.50 2. Bicanex/ol-sta W: 0.96 0.86 0.99 0.92 0.91 0.98 p(normal): 0.38 0.0009 0.99 0.24 0.05 0.73 3. Biecm/biast W: 0.95 0.97 0.98 0.95 0.95 0.97 p(normal): 0.32 0.86 0.63 0.49 0.31 0.63 4. Biecm/zs-zgyi W: 0.97 0.95 0.97 0.95 0.96 0.97 p(normal): 0.58 0.35 0.48 0.55 0.43 0.62 5. Bifmt/bizi W: 0.97 0.95 0.97 0.98 0.97 0.94 p(normal): 0.72 0.43 0.37 0.97 0.6 0.07 6. Bifmt/nas-pros W: 0.97 0.93 0.97 0.92 0.98 0.97 p(normal): 0.61 0.14 0.45 0.24 0.91 0.41 7. Bizi/biast W: 0.98 0.98 0.97 0.95 0.93 0.96 p(normal): 0.96 0.98 0.41 0.51 0.11 0.29 8. Bizi/nas-pros W: 0.97 0.97 0.96 0.96 0.95 0.96 p(normal): 0.68 0.71 0.27 0.71 0.37 0.33 9. Lam-opn/g-o W: 0.98 0.95 0.98 0.93 0.93 0.97 p(normal): 0.82 0.38 0.65 0.26 0.14 0.47 10. Ol-sta/bas-br W: 0.91 0.95 0.96 0.96 0.88 0.97 p(normal): 0.05 0.39 0.23 0.78 0.01 0.41 11. Zs-zgyi/ol-sta W: 0.97 0.97 0.99 0.97 0.95 0.95 p(normal): 0.63 0.87 0.98 0.88 0.32 0.14

4.3.2 Kruskal-Wallis and Mann-Whitney results for cranial indices: By Applying Kruskal-Wallis to papionin samples many significant differences are apparent but also some revealing similarities (Table 4.13). For example, despite absolute differences in size, Cercocebus and Mandrillus are not significantly different in the relationship between 1) bizygomaxillare inferior and biasterionic breadth (biast) and 2) biasterionic breadth (biast), which is essentially the widest lateral points of the occipital bone, and the occipital sagittal chord (lam-opn). Furthermore, the latter index also shows a similarity between Papio and Theropithecus. Surprisingly Lophocebus and Cercocebus overlap in many relative proportions and are not significantly different (Table 4.13, No. 4, 5, 6, 8, 10 and 11). Similarly, Papio and Mandrillus are not significantly different in the relative proportion of the superior facial breadth to inferior facial breadth (bifmt/bizi); and the length of the palate in relation to cranial height (ol-sta/bas-br) (Table 4.13 No. 5

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Table 4.13: Kruskal-Wallis results for cranial indices of the Papionini with Mann-Whitney pairwise comparisons (p (same)) (indices listed in alphabetical order by abbreviation).

L. Cc. Mc. Th. Mn. Pp. n=23 n=19 n=34 n=14 n=22 n=31 1. Biast/lam-opn 0 3.53E-06 0.13 0.01 2.55E-07 0.05 L. H: 74.91 0 1.18E-08 0.02 0.57 0.0001 Cc. Hc: 74.92 0 0.00006 6.32E-10 0.0001 Mc. p(same): 9.70E-15 0 0.005 0.28 Th. 0 3.82E-06 Mn. 0 Pp. 2. Bicanex/ol-sta 0 0.08 0.14 5.00E-07 2.49E-08 1.03E-09 L. H: 102.90 0 0.80 1.39E-06 0.00001 7.17E-07 Cc. Hc: 102.90 0 7.16E-08 5.55E-09 4.91E-11 Mc. p(same): 1.27E-20 0 1.70E-06 1.29E-06 Th. 0 0.35 Mn. 0 Pp. 3. Biecm/biast 0 2.03E-06 0.001 0.0006 1.68E-08 0.00001 L. H: 47.02 0 0.44 0.39 0.004 0.18 Cc. Hc: 47.02 0 0.96 0.009 0.80 Mc. p(same): 5.63E-09 0 0.001 0.70 Th. 0 0.00007 Mn. 0 Pp. 4. Biecm/zs-zgyi 0 0.74 0.002 8.13E-07 0.00007 0.24 L. H: 67.07 0 0.008 3.41E-06 0.0009 0.24 Cc. Hc: 67.07 0 0.00001 1.45E-07 0.03 Mc. p(same): 4.18E-13 0 7.47E-07 4.13E-07 Th. 0 1.97E-06 Mn. 0 Pp. 5. Bifmt/bizi 0 0.84 0.00006 5.00E-07 0.62 0.62 L. H: 58.43 0 0.0005 2.86E-06 0.89 0.54 Cc. Hc: 58.43 0 7.16E-08 0.0001 0.0004 Mc. p(same): 2.57E-11 0 8.83E-07 1.11E-07 Th. 0 0.45 Mn. 0 Pp.

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L. Cc. Mc. Th. Mn. Pp. n=23 n=19 n=34 n=14 n=22 n=31 6. Bifmt/nas-pros 0 0.32 0.12 5.00E-07 1.13E-08 4.74E-10 L. H: 107.50 0 0.37 1.39E-06 1.04E-07 4.20E-09 Cc. Hc: 107.50 0 1.28E-06 1.20E-07 6.77E-12 Mc. p(same): 1.37E-21 0 0.83 3.39E-06 Th. 0 0.00002 Mn. 0 Pp. 7. Bizi/biast 0 3.32E-06 0.73 2.09E-06 0.001 0.81 L. H: 51.74 0 0.0004 0.004 0.47 0.00001 Cc. Hc: 51.74 0 6.72E-06 0.006 0.95 Mc. p(same): 6.11E-10 0 0.003 1.37E-06 Th. 0 0.002 Mn. 0 Pp. 8. Bizi/nas-pros 0 0.47 0.00002 9.37E-06 1.29E-08 4.74E-10 L. H: 105.00 0 0.002 0.0002 1.04E-07 4.20E-09 Cc. Hc: 105.00 0 0.49 0.00003 2.64E-11 Mc. p(same): 4.76E-21 0 0.00008 1.11E-07 Th. 0 0.00001 Mn. 0 Pp. 9. Lam-opn/g-o 0 8.19E-06 0.009 0.15 3.67E-08 0.0006 L. H: 82.58 0 2.25E-08 0.001 0.04 0.06 Cc. Hc: 82.60 0 0.0004 4.12E-10 4.33E-08 Mc. p(same): 2.42E-16 0 8.76E-06 0.09 Th. 0 0.0002 Mn. 0 Pp. 10. Ol-sta/bas-br 0 0.60 0.07 0.00001 4.17E-08 3.74E-09 L. H: 89.29 0 0.28 0.00002 1.20E-07 1.74E-08 Cc. Hc: 89.29 0 0.0002 8.32E-08 1.72E-09 Mc. p(same): 9.49E-18 0 0.02 0.002 Th. 0 0.96 Mn. 0 Pp. 11. Zs-zgyi/ol-sta 0 0.58 0.0001 0.55 1.44E-08 7.24E-09 L. H: 110.30 0 0.00001 0.76 5.01E-08 1.55E-08 Cc. Hc: 110.30 0 0.00005 3.70E-10 6.77E-12 Mc. p(same): 3.47E-22 0 6.32E-07 1.11E-07 Th. 0 0.0003 Mn. 0 Pp.

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and 10). Likewise, Macaca is not significantly different from Lophocebus and Cercocebus in the relative proportion of the superior facial breadth to superior facial height (bifmt/nas- pros) (Table 4.13 No. 6).

4.3.3 One-way ANOVA results for cranial indices: Results from one-way ANOVA support those of the Kruskal-Wallis, although with an increase in statistical power. Only four indices have percentages of variation between groups greater than 70% (Table 4.14, No. 2, 6, 8 and 11) and the remaining indices have results ranging from ~30% to <69% of variation between groups. With regard to the maxillo-alveolar breadth (biecm) and its proportional relationship with biasterionic breadth (biast), Papio is very similar to Cercocebus, and Macaca and Theropithecus and are not significantly different. As box-plot Figure 4.24 suggested, the relative proportion of biasterionic breadth (biast) to the occipital sagittal chord (lam-opn) between Cercocebus

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Table 4.14: One-Way ANOVA results for cranial indices of the Papionini (indices listed in alphabetical order by abbreviation). Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 1. Biast/lam-opn L. Cc. Mc. Th. Mn. Pp. Between F: 32.66 0 0.00002 0.53 0.002 0.00002 0.19 groups: 30989.1 50.70 5 6197.83 28.18 1.48E-19 0.06 df: 56.71 0 0.00002 0.10 0.97 0.0007 Within groups: 30136.1 49.30 137 219.97 p: 1.59E-15 0 0.00002 0.00002 0.0008 Total: 61125.3 142 0 0.008 0.69 0 0.00003 0 2. Bicanex/ ol-sta L. Mc. Th. Pp. Between F: 109.7 0 0.45 0.0001 0.0001 groups: 6460.1 75.27 3 2153.37 99.4 1.30E-29 0.07 df: 44.14 0 0.0001 0.0001 Within groups: 2123.03 24.73 98 21.66 p: 1.74E-20 0 0.0001 Total: 8583.13 101 ex. Mn. & Cc. 0

3. Biecm/biast L. Cc. Mc. Th. Mn. Pp. Between F: 29.09 0 0.00004 0.003 0.003 0.00002 0.004 groups: 1631.19 29.78 5 326.24 11.62 2.27E-09 0.0009 df: 56.32 0 0.88 0.86 0.10 0.84 Within groups: 3845.72 70.22 137 28.07 p: 1.85E-14 0 1 0.004 1 Total: 5476.91 142 0 0.003 1 0 0.003 0

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Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. F comparisons sqrs: Var.: df: square: F: p(same): of Var. test: p(same): 4. Biecm/zs-zgyi L. Cc. Mc. Th. Mn. Pp. Between F: 29.64 0 0.98 0.11 0.00002 0.00002 0.94 L. groups: 9227.14 54.30 5 1845.43 32.56 9.01E-22 0.0005 df: 55.41 0 0.02 0.00002 0.00002 0.59 Cc. Within groups: 7765.2 45.70 137 56.68 p: 1.62E-14 0 0.00006 0.00002 0.60 Mc. Total: 16992.3 142 0 0.00002 0.00002 Th. 0 0.00002 Mn. 0 Pp. 5. Bifmt/bizi L. Cc. Mc. Th. Mn. Pp. Between F: 36.56 0 1 .00007 0.00002 1 0.99 L. groups: 6928.9 53.31 5 1385.78 31.28 3.84E-21 0.0001 df: 56.82 0 .00006 0.00002 1 0.98 Cc. Within groups: 6069.44 46.69 137 44.3 p: 1.44E-16 0 0.00002 0.00004 0.0009 Mc. Total: 12998.3 142 0 0.00002 0.00002 Th. 0 0.97 Mn. 0 Pp. 6. Bifmt/nas-pros L. Cc. Mc. Th. Mn. Pp. Between F: 102.8 0 0.94 0.44 0.00002 0.00002 0.00002 L. groups: 62657.8 72.76 5 12531.6 73.19 5.61E-37 2.40E-07 df: 59.57 0 0.94 0.00002 0.00002 0.00002 Cc. Within groups: 23457 27.24 137 171.22 p: 5.56E-28 0 0.00002 0.00002 0.00002 Mc. Total: 86114.8 142 0 0.99 0.0006 Th. 0 0.003 Mn. 0 Pp.

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Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. F comparisons sqrs: Var.: df: square: F: p(same): of Var. test: p(same): 7. Bizi/biast L. Cc. Mc. Th. Mn. Pp. Between F: 17.38 0 0.002 0.99 0.00002 0.06 0.99 L. groups: 5670.89 37.74 5 1134.18 16.61 8.42E-13 0.0003 df: 56.37 0 0.0003 0.005 0.91 0.0006 Cc. Within groups: 9355.23 62.26 137 68.29 p: 2.25E-10 0 0.00002 0.01 0.99 Mc. Total: 15026.1 142 0 0.00002 0.00002 Th. 0 0.03 Mn. 0 Pp. L. Cc. Mc. Th. Mn. Pp. 8. Bizi/nas-pros 0 0.99 0.002 0.00003 0.00002 0.00002 L. Between F: 101.5 0 0.003 0.0002 0.00002 0.00002 Cc. groups: 48156.1 72.97 5 9631.21 73.98 3.31E-37 0.02 df: 58.03 0 0.98 0.00002 0.00002 Mc. Within groups: 17836.5 27.03 137 130.19 p: 2.17E-27 0 0.00004 0.00002 Th. Total: 65992.6 142 0 0.00006 Mn. 0 Pp. L. Cc. Mc. Th. Mn. Pp. 9. Lam-opn/g-o 0 0.00002 0.25 0.56 0.00002 0.0007 L. Between F: 42.65 0 0.00002 0.0003 0.21 0.43 Cc. groups: 1867.56 56.10 5 373.51 35.01 3.08E-20 0.20 df: 55.95 0 0.002 0.00002 0.00002 Mc. Within groups: 1461.69 43.90 137 10.67 p: 6.96E-18 0 0.00002 0.15 Th. Total: 3329.24 142 0 0.0006 Mn. 0 Pp.

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Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 10. Ol-sta/bas-br Cc. Mc. Th. Pp. Between F: 38.56 0 0.86 0.0003 0.0001 Cc. groups: 21388.9 59.19 3 7129.63 45.44 3.08E-18 8.26E-06 df: 46.36 0 0.0004 0.0001 Mc. Within groups: 14749.2 40.81 94 156.91 p: 1.19E-12 0 0.0002 Th. Total: 36138.1 97 0 Pp. ex. L. & Mn.

L. Cc. Mc. Th. Mn. Pp. 11. Zs-zgyi/ol-sta 0 0.99 0.00007 0.99 0.00002 0.00002 L. Between F: 94.76 0 0.00003 1 0.00002 0.00002 Cc. groups: 21555 79.07 5 4311 102.8 1.99E-44 0.10 df: 58.03 0 0.00002 0.00002 0.00002 Mc. Within groups: 5705.61 20.93 136 41.95 p: 1.29E-26 0 0.00002 0.00002 Th. Total: 27260.6 141 0 0.002 Mn. 0 Pp.

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and Mandrillus is not significantly different, yet Mandrillus remains significantly different to every other genus (Table 4.14, No. 1). This index reveals proportional similarities between Lophocebus-Papio-Theropithecus on the one hand and Cercocebus-Mandrillus on the other. More recently, Gilbert & Rossie (2007) also presented results from allometrically corrected data which is congruent with molecular findings (e.g. Cronin & Sarich, 1976; Distotell, 1994; Page & Goodman, 2001). 4.3.4 Summary for cranial indices: In summary, like the cercopithecins, using raw data to generate indices greatly reduces the size differences between males and females. Furthermore, some indices show similarities between closely related genera that have been identified via genetic analyses (Figures 4.16 and 4.24). Lastly, indices were also helpful in revealing the distinctiveness of Theropithecus (Figures 4.17 and 4.20). 4.4 Bivariate Results: To examine the relationship between cranial variables and body size the Ln transformed generic mean for particular variables were plotted against Ln transformed mean generic body weights (g). Table 4.15 presents the results of linear regression between the generic means of cranial variables and body weight. Very much like the cercopithecins, several cranial variables for the papionins are also significantly correlated with body size. Only five variables were not statistically significant. These include external bicanine breadth (bicanex), interentoglenoid breadth (bien), bizygomaxillare superior breadth (bizs), palatal height (palhei) and maximum length of the zygomatic (zs-zgyi). Figure 4.27 is a bivariate plot of generic mean body weight plotted against the generic means of palatal length A (ol-sta). Only one figure is presented because despite which cranial variable is plotted against body weight, the pattern and distribution of genera remains very similar. This figure, like the one for the cercopithecins, again illustrates how these genera are distributed as well as the informal two groups, mentioned previously and illustrated with box-plots (Figures 4.2-4.15); 1) the first group of genera are small to medium sized with moderate facial projection and vary from arboreal to semiterrestrial, including, Lophocebus, Cercocebus and Macaca; and 2) the second group of genera are medium to large with extreme facial projection and are much more terrestrial, including Theropithecus, Papio and Mandrillus.

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Table 4.15: Results of body weight (Ln (g)) plotted against the generic mean for particular cranial measurements (Ln (mm)) (cranial variables listed in alphabetical order by abbreviation; those in bold represent cranial variables with an adjusted R squared value >0.70 and a p-value <0.05).

P-value/ Mean Generic Body Weight (g) Adjusted Significance (Napier, 1981; Rowe, 1996; Ln (mm) R squared F Delson et al, 2000): bas-br 0.80 0.01 bas-nas 0.79 0.01 Lophocebus spp. - 6825 bas-pros 0.92 0.002 Cercocebus spp. - 8710 biast 0.80 0.01 Macaca spp. - 9020 biaur 0.88 0.004 Theropithecus gelada - 15148 Mandrillus spp. - 18750 bicanex 0.63 0.04 Papio spp. - 19016 biecm 0.93 0.001 bien 0.66 0.03 bienm 0.83 0.007 bifmo 0.77 0.01 bifmt 0.76 0.01 bipor 0.88 0.004 bizi 0.86 0.005 bizs 0.22 0.19 bizygo 0.94 0.0009 br-lam 0.62 0.04 g-o 0.93 0.001 lam-opn 0.73 0.02 nas-br 0.86 0.005 nas-pros 0.90 0.002 nas-rhi 0.89 0.003 ol-sta 0.94 0.001 ol-pms 0.89 0.003 palhei 0.44 0.09 pros-o 0.91 0.002 zs-zgyi 0.67 0.03 zs-zi 0.84 0.007

4.50 Figure 4.27: Adjusted R2: 0.94 Pp. 4.40 Significance F & P-value 0.0001 Mn. 4.30 y = 0.6076x - 1.6334 Th. 4.20 4.10 4.00 Cc. 3.90 L. Mc. 3.80 Ln Palatal Length (ol-sta) Ln Palatal 3.70 3.60 8.60 8.80 9.00 9.20 9.40 9.60 9.80 10.00 Ln Body Weight (g) 154

4.5 Multivariate Statistics and Morphological Distances for the Papionini: To consider and evaluate the entire multivariate datasets simultaneously PCA and CVA were utilized. 4.5.1 PCA results for Ln transformed data: To identify the underlying structure (i.e. the cranial variables which account for most of the variation between genera) of the Papionini multivariate datasets based on Ln transformed generic means, PCA was employed. By doing so, PCA based on the variance- covariance matrix results in the first (Eigenvalue - 1.31; 91.77%), second (Eigenvalue - 0.07; 4.92%) and third (Eigenvalue - 0.03; 2.28%) PCs accounting for 98.97% of the variation (Figure 4.28). One hundred percent of the variation is explained by five PCs. Table 4.18 lists the low to moderate variable loadings for the papionin cranial measurements. The variable loadings for the first PC are all positive and can be interpreted as size differences between genera; the remaining PCs have positive and negative variable loadings. Evidence to substantiate this can be seen in the mean PCA object scores per genus. Lophocebus, one of the smallest papionins, achieves the least with 19.53 (succeeded by Macaca at 19.79) and Papio, one of the largest papionins, scores the greatest with 22.32 (followed by Mandrillus at 21.67). The first PC is governed by nasofacial lengths. These include, the sagittal length of the nasal bones (nas-rhi), nasal height (nas-ns), superior facial height (nas-pros), all three palatal lengths (ol-sta, ol-pms and iv-pms), sagittal length of the nasal aperture (rhi-ns) and superior facial length (bas-pros) with positive variable loadings of 0.35, 0.30, 0.29, 0.24, 0.24, 0.23, 0.21 and 0.20, correspondingly (Table 4.18). The second PC’s highest positive variable loadings dental and palatal measurements such as, internal bicanine (bicanin) and biseptal (biseptal) breadths, I1I2 alveolar length (i1i2), distance between incisivion and the palatomaxillary suture (iv-pms) and external bicanine breadth (bicanex) with 0.45, 0.30, 0.27, 0.26 and 0.24, respectively. In contrast to these dimensions are negatively loaded features such as, the maximum length of the zygomatic bone (zs-zgyi), sagittal length of the nasal aperture (rhi-ns), the zygomatico- maxillary suture length (zs-zi) and palatal height at M2 (palhei) with -0.36, -0.27, -0.25 and -0.25, correspondingly.

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Figure 4.28: Papionini scatterplot of PC 1 & 2 and PC 2 & 3 for Ln transformed generic means. Cc. - Cercocebus; Mn. - Mandrillus; Mc. - Macaca; L. - Lophocebus; Pp. - Papio; and Th. - Theropithecus. Bizygomaxillare -1 Canine & Incisor Superior & Breadths with -1.1 Th. Occipital Length Palatal lengths 3.7 -1.2 Mn. -1.3 3.6 Cc. L. Mn. -1.4 L.

PC 2: -1.5 PC 3: Cc. Increase in body size 3.5 4.92% 2.28% -1.6 Pp. Mc. -1.7 3.4 Pp.

-1.8 Length of Canine & Incisor -1.9 3.3 Zygomatic bone Breadths with -2 Th. Palatal lengths Nasal & Facial lengths Mc. 3.2

Length of 20 21 22 -1.9 -1.8 -1.7 -1.6 -1.5 -1.4 -1.3 -1.2 -1.1 -1 Zygomatic bone PC 1: 91.77% PC 2: 4.92% Palatal height & Nasal length

% of PCA PC Eigenvalue Var. Cum. %. scores Axis 1 Axis 2 Axis 3 1 1.31 91.77 91.77 Cc. 19.82 -1.50 3.61 2 0.07 4.92 96.68 Mn. 21.67 -1.21 3.60 3 0.03 2.28 98.97 Mc. 19.79 -1.67 3.23 L. 19.53 -1.37 3.62 Table 4.16: Inter-generic Euclidean distances based on PCA scores from PC 1-5 per genus from Ln transformed generic means. Pp. 22.32 -1.61 3.41 Th. 20.88 -1.98 3.73 Cc. 0 Mean 1.51 Mean 20.67 -1.56 3.53 Mn. 1.89 0 Range 2.43 Max. 22.32 -1.98 3.73

Mc. 0.46 1.97 0 Min. 19.53 -1.21 3.23

L. 0.38 2.16 0.58 0 Range 2.79 0.77 0.50

Pp. 2.51 0.83 2.55 2.81 0 SD 1.14 0.26 0.18 0 Th. 1.19 1.11 1.24 1.49 1.53 CV 5.54 17.04 5.11 Cc. Mn. Mc. L. Pp. Th.

Mean Max Min Range Cc. 0.85 1.42 0.41 1.01 Table 4.17: Intra-generic Mn. 1.41 3.11 0.27 2.85 Euclidean distances based on Ln transformed data. Mc. 1.07 3.00 0.42 2.58 L. 0.79 1.72 0.24 1.48 Pp. 1.04 2.70 0.27 2.43 Th. 0.87 1.59 0.25 1.34

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Table 4.18: Variable loadings for Papionini Ln transformed generic means. PC loadings PC 1 PC 2 PC 3 bizs 0.07 zs-zygi -0.36 palhei -0.42 br-lam 0.07 rhi-ns -0.27 nas-rhi -0.24 nas-br 0.08 zs-zi -0.25 maxnawi -0.23 bas-nas 0.09 palhei -0.25 inbrnabo -0.18 bas-br 0.09 inbrnabo -0.18 bicanin -0.17 bifmo 0.09 nas-br -0.16 nas-ns -0.16 g-o 0.10 bizygo -0.15 bicanex -0.15 bien 0.10 bas-nas -0.13 bienm -0.15 bifmt 0.10 bizi -0.10 bifmt -0.12 biaur 0.10 g-o -0.08 nas-pros -0.12 biast 0.11 bas-br -0.08 biseptal -0.07 zs-zygi 0.11 maxalvlen -0.07 biast -0.05 bizi 0.11 bien -0.07 bifmo -0.04 bipor 0.12 nas-pros -0.07 biecm -0.03 palhei 0.12 maxnawi -0.04 zs-zi -0.02 bizygo 0.13 biaur -0.03 nas-br 0.01 inbrnabo 0.13 bipor -0.02 zs-zygi 0.01 biecm 0.13 biast -0.02 biaur 0.04 bienm 0.14 nas-ns -0.02 br-lam 0.04 bicanin 0.14 ol-sta -0.02 pros-o 0.04 biseptal 0.14 bas-pros 0.00 i1i2 0.05 lam-opn 0.14 biecm 0.01 g-o 0.05 zs-zi 0.15 pros-o 0.01 bas-pros 0.08 i1i2 0.15 ol-pms 0.02 bipor 0.08 bicanex 0.15 bifmo 0.03 bas-nas 0.08 pros-o 0.19 br-lam 0.07 rhi-ns 0.09 maxnawi 0.19 bienm 0.07 bizygo 0.11 maxalvlen 0.19 nas-rhi 0.07 bas-br 0.15 bas-pros 0.20 lam-opn 0.08 maxalvlen 0.15 rhi-ns 0.21 bifmt 0.08 bien 0.17 ol-pms 0.23 bizs 0.10 ol-sta 0.17 iv-pms 0.24 bicanex 0.24 ol-pms 0.18 ol-sta 0.24 iv-pms 0.26 iv-pms 0.22 nas-pros 0.29 i1i2 0.27 bizi 0.25 nas-ns 0.30 biseptal 0.30 lam-opn 0.29 nas-rhi 0.35 bicanin 0.45 bizs 0.39

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The third PC’s largest positive loaded measurements are the bizygomaxillare superior breadth (bizs), occipital sagittal chord (lam-opn), bizygomaxillare inferior breadth (bizi) and length between incisivion and the palatomaxillary suture (iv-pms) achieving variable loadings of 0.39, 0.29, 0.25 and 0.22, respectively. Contrasting with these dimensions are cranial measurements such as palatal height at M2 (palhei), sagittal length of the nasal bones (nas-rhi) and the maximum width of the nasal aperture (maxnawi) with negative variable loadings of -0.42, -0.24 and -0.23, correspondingly. Table 4.16 provides the inter-generic Euclidean distance matrix produced from PCA object scores per genus from the first to fifth principal components based on Ln transformed generic means. The mean distance between papionin genera is 1.51 (Range 2.43). The largest distance produced is between Papio and Lophocebus at 2.81 while the smallest distance is between Lophocebus and Cercocebus with 0.38 succeeded by Cercocebus and Macaca with 0.46. For comparison, Table 4.17 provides intra-generic Euclidean distances based on Ln transformed data. The largest mean intra-generic Euclidean distance was produced by Mandrillus with 1.41 (next followed by Macaca and Papio with 1.07 and 1.04). This is due to the extreme sexual dimorphism between males and females. The smallest mean intra-generic Euclidean distance was produced by Lophocebus with 0.79 (succeeded by Cercocebus, 0.85). In summary, the major distinguishing features between papionin genera identified by PCA, like the cercopithecins, involves the dimensions of the naso-facial and palatal regions. In particular, the sagittal length of the nasal bones (nas-rhi), nasal height (nas-ns), superior facial height (nas-pros) and palatal length A (ol-sta) were always amongst the highest loaded variables (Table 4.18). Again, like the cercopithecins, neurocranial dimensions did not figure prominently in distinguishing genera but the occipital sagittal chord (lam-opn) did receive a relatively large loading on the third PC.

4.5.2 PCA results for MSV: PCA based on the variance-covariance matrix results in the first (Eigenvalue – 0.36, 80.57%) and second (Eigenvalue – 0.05, 10.71%) PCs accounting for 91.29% of variation in the tribe (5.39% less variation explained than PCs 1 & 2 for Ln transformed generic means but 84.09% more variation explained then PCs 2 & 3 for Ln transformed generic

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means; Figure 4.29). One hundred percent of the variation is explained by five PCs. The variable loadings for the first PC are not all positive (Tables 4.19) and again vary from low to moderate; the second PC also has mixed values. The extent to which size has been removed can be seen in the PCA object scores per genus. PCA object scores for PC 1 have mixed values. Lophocebus, Cercocebus and Macaca and have positive values, 0.31, 0.24 and 0.11; but Theropithecus, Mandrillus and Papio have negative values, -0.25, -0.83 and - 1.13. The largest positively loaded MSV for the first PC is cranial vault length (g-o) with 0.23 (Table 4.19). Contrasting with this dimension are negatively loaded cranial and facial length MSV. Some of these include, superior facial height (nas-pros), nasal height (nas-ns), sagittal length of the nasal bones (nas-rhi), palatal length A (ol-sta), maximum cranial length (pros-o) and superior facial length (bas-pros) with negative variable loadings of - 0.44, -0.43, -0.36, -0.22, -0.21 and -0.20, respectively. The positively loaded cranial MSV of the second PC are breadth measurements, which include external and internal bicanine breadths (bicanex and bicanin), superior facial breadth (bifmt) and biseptal breadth (biseptal) with variable loadings of 0.27, 0.25, 0.24 and 0.24, correspondingly. Contrasting with these MSV are length and breadth dimensions such as the maximum length of the zygomatic (zs-zgyi), bizygomatic (bizygo) and bizygomaxillare inferior (bizi) breadths and maxillo-alveolar length (pros-dm3) with variable loadings of -0.42, -0.40, -0.25 and -0.21, respectively. Table 4.20 provides the inter-generic Euclidean distance matrix produced from PCA object scores per genus from the first to fifth principal components based on generic mean MSV. The average distance between papionin genera is 0.87 (Range 1.22). Again, the largest distance produced is between Papio and Lophocebus at 1.46 while the smallest distance is between Lophocebus and Cercocebus with 0.24 followed by Cercocebus and Macaca with 0.40. For comparison, Table 4.21 provides intra-generic Euclidean distances based on MSV. The largest mean intra-generic Euclidean distance was again produced by Mandrillus at 0.78 (again, followed by Macaca and Papio with 0.77 and 0.71). The smallest mean intra-generic Euclidean distance was produced by Theropithecus at 0.54 (succeeded closely by Lophocebus, 0.55).

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Figure 4.29: Papionini scatterplot of PC 1 & 2 for generic mean MSV. Cc. - Cercocebus; Mn. - Mandrillus; Mc. - Macaca; L. - Lophocebus; Pp. - Papio; and Th. - Theropithecus. Table 4.19: Variable loadings for -2 Papionini generic mean MSV. Canine breadths -2.1 PC loadings

-2.2 PC 1 PC 2 nas-pros -0.44 zs-zygi -0.42 -2.3 nas-ns -0.43 bizygo -0.40 Mn. L. -2.4 nas-rhi -0.36 bizi -0.25 PC 2: Cc. ol-sta -0.22 maxalvlen -0.21 -2.5 Pp. 10.71% Mc. pros-o -0.21 nas-br -0.18 -2.6 bas-pros -0.20 zs-zi -0.18

-2.7 ol-pms -0.14 bas-nas -0.17 iv-pms -0.11 ol-sta -0.17 -2.8 maxalvlen -0.09 rhi-ns -0.16 -2.9 rhi-ns -0.06 nas-pros -0.16 Nasal & Facial lengths Th. Cranial length maxnawi -0.03 g-o -0.15 Length & breadth -1.1 -1 -0.9-0.8-0.7-0.6-0.5-0.4-0.3-0.2 -0.1 0 0.1 0.2 0.3 0.4 bicanex -0.01 bas-pros -0.09 of Zygomatic PC 1: 80.57% i1i2 0.00 bas-br -0.08 biseptal 0.01 ol-pms -0.07 bicanin 0.01 pros-o -0.05 % of inbrnabo 0.01 bien -0.04 PC Eigenvalue Var. Cum. %. palhei 0.01 inbrnabo -0.03 1 0.36 80.57 80.57 zs-zi 0.01 palhei -0.02 2 0.05 10.71 91.29 bienm 0.02 bipor -0.01

3 0.03 5.69 96.97 lam-opn 0.02 maxnawi -0.01

biecm 0.04 nas-ns 0.00 PCA zs-zygi 0.09 biaur 0.01 scores Axis 1 Axis 2 biast 0.10 biecm 0.03 Cc. 0.24 -2.43 bipor 0.10 lam-opn 0.04 Mn. -0.83 -2.36 bien 0.11 biast 0.05 Mc. 0.11 -2.50 bizygo 0.12 nas-rhi 0.06 L. 0.31 -2.36 bizi 0.12 bienm 0.07 Pp. -1.13 -2.47 bizs 0.12 bizs 0.08 Th. -0.25 -2.94 bifmo 0.14 iv-pms 0.12 Mean -0.26 -2.51 br-lam 0.14 i1i2 0.12 Max. -1.13 -2.94 bifmt 0.15 br-lam 0.14 Min. 0.31 -2.36 bas-br 0.15 bifmo 0.15 Range 1.44 0.58 biaur 0.16 biseptal 0.24 SD 0.60 0.22 nas-br 0.19 bifmt 0.24 CV 232.97 8.70 bas-nas 0.19 bicanin 0.25 g-o 0.23 bicanex 0.27

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Table 4.20: Inter-generic Euclidean distances based on PCA scores from PC 1-5 per genus from generic mean MSV. Cc. 0 Mean 0.87 Mn. 1.10 0 Range 1.22 Mc. 0.40 1.04 0 L. 0.24 1.16 0.45 0 Pp. 1.39 0.50 1.26 1.46 0 Th. 0.72 0.83 0.67 0.81 1.03 0 Th. Cc. Mn. Mc. L. Pp.

Table 4.21: Intra-generic Euclidean distances based on MSV.

Mean Max Min Range Cc. 0.61 0.85 0.34 0.51 Mn. 0.78 1.57 0.27 1.30

Mc. 0.77 1.42 0.36 1.06 L. 0.55 0.99 0.23 0.76 Pp. 0.71 1.49 0.25 1.24 Th. 0.54 0.77 0.22 0.56

In summary, the major distinguishing MSV of the papionin genera identified by PCA again include naso-facial lengths and breadths (nas-pros, nas-ns, ol-sta bicanex, bicanin and bifmt) along with one neurocranial measurements, such as cranial vault length (g-o). Other measurements which receive large variable loadings comprise the maximum length of the zygomatic (zs-zgyi) and bizygomaxillare inferior breadth (bizi).

4.5.3 CVA results for Ln transformed data: By subjecting the entire Ln transformed pooled sex multivariate dataset to MANOVA and CVA the first (Eigenvalue - 17.61; 50.20%) and second (Eigenvalue - 9.07; 25.84%) canonical variate axes account for 76.04% of the variation within the tribe (Figure 4.30). Table 4.24 lists the CVA object scores per genus and Table 4.22 lists the variable loadings for the cranial measurements. Similar to PCA, the mean CVA object scores for the first canonical variate axis per genus also seem to be sized related. Lophocebus has the least, 9.03 (Min. - Max., 8.64 - 9.39), with Papio receiving the highest, 10.35 (Min. - Max., 9.73 - 10.84). To test for significance of results, CVA object scores per genus (for both Ln transformed and MSV multivariate datasets) from the first ten canonical variate axes were subject to a two-sample multivariate t-test using StatistiXL (version 1.5).

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Figure 4.30: Papionini scatterplot of canonical variate axes 1 & 2 for Ln transformed pooled sex data with 95% confidence ellipses. Cercocebus - black ; Mandrillus - cross +; Macaca - square ; Lophocebus - x; Papio - circle o; and Theropithecus - diamond . Superior Facial breadth 2.6 Table 4.22: Variable loadings for Papionini 2.5 Ln transformed pooled sex data. 2.4 CVA Eigen- Eigen- 2.3 CV vectors Axis 1 vectors Axis 2 Axis 2: 2.2 zs-zygi -0.28 bifmt -0.44 25.84% bas-nas -0.26 pros-o -0.24 2.1 bicanex -0.23 bicanex -0.23 2 bizygo -0.14 bicanin -0.22 palhei -0.13 nas-ns -0.21 1. 9 zs-zi -0.10 bifmo -0.20 1. 8 Max. Cranial & bifmt -0.08 bienm -0.17 Zygomatic & Inferior Nasal lengths biseptal -0.04 biseptal -0.15 1. 7 Inferior cranial lengths ol-pms 0.00 biast -0.10 Facial 9 10 11 biast 0.01 palhei -0.07 breadth CV Axis 1: 50.20% inbrnabo 0.02 biecm -0.07 bizs 0.02 nas-rhi -0.06 g-o 0.02 maxnawi -0.05 MANOVA bipor 0.03 iv-pms -0.05 Wilks's Pillai br-lam 0.03 nas-br -0.03 lambda: 0.00 trace: 3.86 bienm 0.04 inbrnabo 0.00 df1: 180.00 df1: 180.00 rhi-ns 0.04 biaur 0.02 df2: 511.30 df2: 530.00 bien 0.06 br-lam 0.02 F: 14.26 F: 9.99 ol-sta 0.06 i1i2 0.02 p(same): 0.00 p(same): 0.00 bizi 0.07 bas-pros 0.04 CVA nas-br 0.07 bas-nas 0.07 Eigenvalue 1: 17.61 Percent: 50.20 bicanin 0.11 nas-pros 0.10 Eigenvalue 2: 9.07 Percent: 25.84 i1i2 0.15 bipor 0.11 Total %: 76.04 maxnawi 0.15 maxalvlen 0.11 nas-ns 0.16 ol-pms 0.14 bifmo 0.17 bien 0.14

Table 4.23: Inter-generic Euclidean distances based on mean biaur 0.18 zs-zygi 0.15 CVA scores from axes 1-5 per genus from Ln transformed data. biecm 0.18 zs-zi 0.15 bas-pros 0.21 bas-br 0.15 Cc. 0 Mean 0.94 nas-rhi 0.22 bizs 0.18 Mn. 1.09 0 Range 1.39 iv-pms 0.24 bizygo 0.19 Mc. 0.39 1.17 0 lam-opn 0.25 lam-opn 0.20 L. 0.26 1.22 0.45 0 maxalvlen 0.26 g-o 0.21 Pp. 1.49 0.61 1.51 1.65 0 bas-br 0.27 rhi-ns 0.23 Th. 0.81 0.72 0.85 0.98 0.89 0 nas-pros 0.29 ol-sta 0.25 Cc. Mn. Mc. L. Pp. Th. pros-o 0.33 bizi 0.26

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Table 4.24: Cercocebus spp. (n=19; 10, 9). Lophocebus spp. (n=23; 11, 12). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean 9.17 2.10 Mean 9.03 2.03 Max. 9.38 2.25 Max. 9.39 2.16 Min. 8.93 2.00 Min. 8.64 1.94 Range 0.45 0.24 Range 0.75 0.22 SD 0.15 0.07 SD 0.20 0.06 CV 1.62 3.26 CV 2.21 2.96 Mandrillus spp. (n=22; 8, 14). Papio spp. (n=31; 19, 12). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean 10.17 2.05 Mean 10.35 2.16 Max. 10.75 2.16 Max. 10.84 2.28 Min. 9.70 1.94 Min. 9.73 2.07 Range 1.06 0.22 Range 1.11 0.22 SD 0.38 0.06 SD 0.27 0.05 CV 3.72 2.87 CV 2.62 2.46 Macaca spp. (n=34; 15, 19). Theropithecus gelada (n=14; 7, 7). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean 9.04 1.93 Mean 9.63 2.46 Max. 9.61 2.14 Max. 9.92 2.55 Min. 8.38 1.72 Min. 9.45 2.36 Range 1.23 0.42 Range 0.47 0.19 SD 0.25 0.07 SD 0.16 0.05 CV 2.80 3.78 CV 1.66 1.89

For each pair wise generic comparison, the null hypothesis that there is no significant difference was rejected (p < 0.0001). The maximum length of the cranium (pros-o) followed by superior facial height (nas-pros) and cranial height (bas-br) are positively correlated with the first CV axis possessing variable loadings of 0.33, 0.29 and 0.27, respectively. Juxtaposed against these dimensions are negatively correlated measurements such as the maximum length of the zygomatic bone (zs-zgyi), inferior facial length (bas-nas) and the external bicanine breadth (bicanex) with variable loadings of -0.28, -0.26, and -0.23, correspondingly. The second CV axis has an interesting assortment of positively and negatively loaded dimensions. Measurements which are positively correlated with CV axis 2 include bizygomaxillare inferior breadth (bizi), palatal length A (ol-sta) and sagittal height of the nasal aperture (rhi-ns), with variable loadings of 0.26, 0.25 and 0.23, respectively. Working against these dimensions are negatively loaded dimensions which include the superior facial breadth (bifmt), maximum cranial length (pros-o) and the external and internal 163

bicanine breadth (bicanex and bicanin) with variable loadings of -0.44, -0.24, -0.23 and - 0.22, correspondingly. The scatterplot for CV axes 1 & 2 of pooled sex Ln transformed data reveal the distinctiveness of Theropithecus but Cercocebus, Lophocebus and Macaca noticeably overlap, as well as, Papio and Mandrillus (Figure 4.30). Table 4.23 provides the inter-generic Euclidean distance matrix produced from mean CVA object scores per genus from the first to fifth canonical variate axes based on Ln transformed pooled sex data. The average distance between papionin genera was 0.94 (Range 1.39). The largest distance produced was between Papio and Lophocebus at 1.65 (followed by Papio and Cercocebus with 1.49) whereas the smallest distance was between Lophocebus and Cercocebus with 0.26 succeeded by Cercocebus and Macaca with 0.39. In summary, the major distinguishing features identified by CVA which achieve relatively large variable loadings comprise both neuro- and viscerocranial measurements (Table 4.22). Specifically, dimensions with large variable loadings include maximum cranium length (pros-o), superior facial breadth (bifmt), palatal length A (ol-sta), bizygomaxillare inferior breadth (bizi), cranial vault length (g-o), occipital sagittal chord (lam-opn) and external and internal bicanine breadths (bicanex and bicanin).

4.5.4 CVA results for MSV: After converting the raw data of the Papionini into MSV and subjecting the entire multivariate dataset to MANOVA and CVA, the first (Eigenvalue - 13.69; 43.75%) and second (Eigenvalue - 9.40; 30.05%) canonical variate axes account for 73.8% of the variation (Figure 4.31; only 2.24% less variation explained than Ln transformed pooled sex data). Table 4.25 lists the low to moderate variable loadings for the cranial measurements. CVA object scores for the first CV axis are positive while those for the second axis are negative (Table 4.27). Interestingly, despite correction for size, Papio received on average the largest for CV axis 1, 1.03 (Max. - Min., 1.36 - 0.74; followed by Mandrillus at 1.0) and Lophocebus the smallest, 0.21 (Max. - Min., 0.43 - 0.0; succeeded by Macaca at 0.25). The first CV axis is very similar to the results for Ln transformed generic means. Cranial MSV which are positively correlated with CV axis 1 include the maximum length of the cranium (pros-o), superior facial height (nas-pros),

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Figure 4.31: Papionini scatterplot of canonical variate axes 1 & 2 for pooled sex MSV with 95% confidence ellipses. Cercocebus - black ; Mandrillus - cross +; Macaca - square ; Lophocebus - x; Papio - circle o; and Theropithecus - diamond .

Palatal length and Inferior -0.6

Facial -0.7 breadth Table 4.25: Variable loadings for -0.8 Papionini pooled sex MSV: -0.9 CVA CV Eigen- Eigen- -1 Axis 2: vectors Axis 1 vectors Axis 2 30.05% -1.1 bas-nas -0.43 ol-sta -0.32 zs-zygi -0.35 bizi -0.27 -1.2 palhei -0.25 lam-opn -0.23 -1.3 bicanex -0.20 rhi-ns -0.23

-1.4 bizygo -0.19 bizygo -0.18 Inferior cranial & zs-zi -0.18 maxalvlen -0.18 Max. Cranial & zygomatic lengths g-o -0.16 ol-pms -0.17 Nasal lengths nas-br -0.14 bizs -0.14 Superior Facial 0 1 2 br-lam -0.10 bas-br -0.14 breadth bizs -0.09 g-o -0.13 biast -0.08 zs-zi -0.13 rhi-ns -0.07 nas-pros -0.13 MANOVA ol-pms -0.05 bien -0.12 Wilks's Pillai inbrnabo -0.03 bipor -0.11 lambda: 0.00 trace: 3.84 biaur -0.03 bas-pros -0.10 df1: 180.00 df1: 180.00 bien -0.03 i1i2 -0.09 df2: 511.30 df2: 530.00 biseptal -0.02 zs-zygi -0.07 F: 13.37 F: 9.72 bienm -0.01 inbrnabo -0.01 p(same): 0.00 p(same): 0.00 bizi -0.01 maxnawi 0.00 CVA bifmt -0.01 br-lam 0.01 Eigenvalue 1: 13.69 Percent: 43.75 bipor 0.00 bas-nas 0.01 Eigenvalue 2: 9.40 Percent: 30.05 bas-br 0.04 iv-pms 0.04 Total %: 73.80 i1i2 0.06 biaur 0.04 ol-sta 0.06 nas-rhi 0.07 bifmo 0.08 pros-o 0.07 bicanin 0.10 nas-br 0.08 Table 4.26: Inter-generic Euclidean distances based on mean CVA scores from axes 1-5 per genus from MSV. biecm 0.12 palhei 0.11 bas-pros 0.14 biast 0.11 Cc. 0 Mean 0.69 iv-pms 0.14 biecm 0.12 Mn. 0.83 0 Range 0.90 maxnawi 0.14 nas-ns 0.13 Mc. 0.35 0.80 0 lam-opn 0.17 biseptal 0.14 L. 0.20 0.87 0.37 0 nas-rhi 0.18 bienm 0.15 Pp. 1.07 0.42 0.96 1.10 0 nas-ns 0.21 bicanin 0.19 Th. 0.60 0.69 0.57 0.67 0.82 0 maxalvlen 0.23 bifmo 0.23 Cc. Mn. Mc. L. Pp. Th. nas-pros 0.24 bicanex 0.25 pros-o 0.35 bifmt 0.46

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Table 4.27: Cercocebus spp. (n=19; 10, 9). Lophocebus spp. (n=23; 11, 12). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean 0.27 -0.98 Mean 0.21 -0.92 Max. 0.46 -1.08 Max. 0.43 -1.04 Min. 0.13 -0.91 Min. 0.00 -0.82 Range 0.33 0.18 Range 0.43 0.22 SD 0.08 0.05 SD 0.11 0.05 CV 30.60 4.63 CV 51.66 5.94 Mandrillus spp. (n=22; 8, 14). Papio spp. (n=31; 19, 12). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean 1.00 -1.00 Mean 1.03 -1.02 Max. 1.36 -1.11 Max. 1.36 -1.15 Min. 0.72 -0.89 Min. 0.74 -0.95 Range 0.63 0.22 Range 0.62 0.20 SD 0.19 0.05 SD 0.18 0.05 CV 19.31 4.91 CV 17.33 5.12 Macaca spp. (n=34; 15, 19). Theropithecus gelada (n=14; 7, 7). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean 0.25 -0.82 Mean 0.43 -1.34 Max. 0.53 -0.93 Max. 0.55 -1.44 Min. -0.09 -0.66 Min. 0.33 -1.23 Range 0.61 0.27 Range 0.22 0.21 SD 0.16 0.06 SD 0.07 0.05 CV 64.06 7.15 CV 15.62 4.06 the maxillo-alveolar length (maxalvlen) and nasal height (nas-ns) with variable loadings of 0.35, 0.24, 0.23 and 0.21, respectively. Juxtaposed against these MSV are negatively loaded dimensions such as inferior cranial length (bas-nas), maximum length of the zygomatic bone (zs-zgyi), palatal height (palhei) and external bicanine breadth (or intercanine breadth, bicanex) with variable loadings of -0.43, -0.35, -0.25 and -0.20, correspondingly. The second CV axis has positively loaded breadth MSV juxtaposed against negatively loaded length and breadth dimensions. The positively loaded MSV include bifrontomalartemporale (bifmt), external bicanine (bicanex) and bifrontomalarorbitale (bifmo) breadths with variable loadings of 0.46, 0.25 and 0.23, respectively. The negatively loaded MSV include palatal length A (ol-sta), bizygomaxillare inferior breadth (bizi), occipital sagittal chord (lam-opn) and the sagittal height of the nasal aperture (rhi-ns) with variable loadings of -0.32, -0.27, -0.23 and -0.23, correspondingly. The scatterplot for CV axes 1 & 2 of pooled sex MSV is very similar to the scatterplot for Ln transformed data, just orientated differently. Again, the scatterplot reveals 166

the distinctiveness of Theropithecus but Cercocebus, Lophocebus and Macaca overlap, as well as, Papio and Mandrillus (Figure 4.31). Table 4.26 provides the inter-generic Euclidean distance matrix produced from mean CVA object scores per genus from the first to fifth canonical variate axes based on pooled sex MSV. The average distance between papionin genera is 0.69 (Range 0.90). Again, the largest distance produced is between Papio and Lophocebus at 1.10 while the smallest distance is between Lophocebus and Cercocebus with 0.20 followed by Cercocebus and Macaca with 0.35. For comparison, Table 4.28 provides Cherry et al’s (1978) M distance between papionin genera. The average M distance between genera is 1.38 (Range 1.61). The largest morphological divergence is achieved by Papio and Lophocebus, 2.09 (followed by Papio and Cercocebus at 1.88); the smallest is between, Lophocebus and Cercocebus, 0.48 (succeeded by Cercocebus and Macaca at 0.66). In summary, the measurements identified by CVA are very similar to those for Ln transformed data (Tables 4.22 and 4.25).

Table 4.28: Inter-generic Cherry et al’s M distance between papionin genera. Cc. 0 Mean 1.38 Mn. 1.59 0 Range 1.61 Mc. 0.66 1.6 0 L. 0.48 1.65 0.8 0 Pp. 1.88 0.74 1.68 2.09 0 Th. 1.43 1.45 1.23 1.85 1.54 0 Cc. Mn. Mc. L. Pp. Th.

4.6 Limb Proportions: Published data on postcranial elements and limb proportions for the papionins (Tables 4.1-4.6) was gathered from Napier & Napier (1967), Jolly (1970), Hill (1970 & 1974), Napier (1981), Strasser & Delson (1987), Strasser (1988, 1992 & 1994), Groves (1989 & 2001a), Krentz (1993), Ciochon (1993), Nakatsukasa (1994 & 1996) and Whitehead et al (2005). Limb proportions of the papionins are very informative (Tables 4.1-4.6). Lophocebus, which is highly arboreal (Horn, 1987a & 1987b), and has a intermembral index to match, 78-79%; and is significantly different from Cercocebus, based on metric analyses and gross anatomical features of the fore- and hindlimbs (Nakatsukasa, 1994 & 1996; Fleagle & McGraw, 1999 & 2002; Yirga, 2002). Further

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relative proportions of papionin manus and pedal elements also reveal striking differences related to substrate preference (Schultz, 1963; Napier & Napier, 1967; Jolly, 1970; Watkins, 2003). Of particular note is the large range of variation seen in the limb indices for Macaca. All three indices have ranges >10, which certainly suggests varied body proportions related to different locomotion and substrate preferences when compared to the other papionin and catarrhine genera (ex. Cercopithecus). For example, the intermembral index for M. nigra is 84% while for M. ochreata it is 100% (Fleagle, 1998).

4.7 Genetics of the Papionini: Genetic research of the Papionini has greatly elucidated the phylogenetic relationships of genera and their subsequent adaptive radiation. Prior to robust comparative genetic analyses many classification schemes were suggested based on morphological features. These include a close relationship between mandrills, drills and baboons on the one hand and another between the African tropical forest mangabeys of Cercocebus and Lophocebus. Both of these arrangements are highly unlikely (Disotell, 1994; Harris, 2000; Page & Goodman, 2001). Instead, there is strong evidence for close genetic relationships between Cercocebus and Mandrillus on the one hand and another between Lophocebus, Papio and Theropithecus. As to the latter relationship, more work is needed to clarify which genus diverged first and which two genera are more closely related to the exclusion of the first. This is very similar to the ‘trichotomy’ problem of the African Apes and their phylogenetic relationships during the 1980s and early 90s (see Rogers, 1993) but with further research it has been demonstrated that Homo and Pan are more closely related to each other than either is to the Gorilla (e.g. Ruvolo, 1997a; Satta et al, 2000; Page & Goodman, 2001). With regard to intra-generic relationships, the only seriously questionable papionin genus is Macaca. Recent studies have highlighted the fact that some macaque species are more closely related to each other than any are to other macaque species. One of the most striking examples of this phenomenon is the origin and phyletic placement of M. arctoides, which Fooden (1990) describes as one of the most ‘distinctive’ species of the genus (see also, Eudey, 1980 and Tosi et al, 2003b). Prior to mtDNA and nuclear DNA comparisons, male genitalia anatomy, pelage, tail-length and osteological morphologies formed the bases

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of inferring phylogenetic relationships of macaque species and larger species-groups (e.g. Fooden, 1976; Delson, 1980). Fooden (1976) recognized three species-groups while Delson (1980) recognized four. However, this thesis is employing the taxonomy proposed by Groves (2001a) and his analysis recognizes six species-groups, which include; 1) M. sylvanus; 2) M. nemestrina group; 3) Sulawesi group; 4) M. fascicularis group; 5) M. mulatta group; and 6) M. sinica group. By and large genetic analyses have supported these arrangements but the evolutionary history of M. arctoides and M. mulatta are quite complicated. First, genetic evidence shows that M. arctoides originated from a hybridization process involving M. assamensis/thibetana-like males with M. fascicularis- like females (Tosi et al, 2000 & 2003a & b; Brandon-Jones et al, 2004; Morales & Melnick, 1998). Second, M. mulatta, which has the largest geographic range of macaque species (from Afghanistan and Pakistan to Eastern China) and is known to hybridize with M. fascicularis (Tosi et al, 2003b; Li & Zhang, 2005; Thierry, 2007). In addition, Froehlich & Supriatna (1996) also report hybridizations between macaque species on the island of Sulawesi. Furthermore, the presence or absence of sexual skin swellings for some macaque species also suggests intra-generic relationships based on phylogeny and sexual behavior differences (van Schaik et al, 1999). Table 4.29 provides inter-generic genetic distances between papionin genera from Page et al, 1999.

Table 4.29: Inter-generic Uncorrected ‘p’ genetic distances between papionin genera from Page et al, 1999.

Cc. 0 Mean 0.02 Mn. 0.018 0 Range 0.014 Mc. 0.025 0.024 0 L. 0.025 0.021 0.0193 0 Pp. 0.026 0.0225 0.023 0.014 0 Th. 0.024 0.0215 0.0216 0.012 0.012 0

Cc. Mn. Mc. L. Pp. Th.

Lastly, yet remarkably, the diploid number of all papionins is 42. This uniformity is unusual when compared to their sister-tribe the Cercopithecini or the Lesser apes of the Hylobatidae which have varied chromosome numbers from >38 up to 72 (Groves, 2001a). However, colobines (diploid number ranges from 44 to 48) and Great apes (diploid number

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ranges from 46 to 48) have experienced only slight chromosome number increases and/or decreases.

4.8 Discussion and Conclusion: From the forgoing analysis many sound inferences and conclusions may be drawn. Like the cercopithecins, size, particularly the length of the face or nasal bones, is an important feature which distinguishes papionin genera. There have been varying degrees of conservatism, plasticity (or versatility?) and evolutionary convergence within the papionins related to similarities in social organization and substrate use (Collard & Wood, 2001a; Fleagle & McGraw, 2002; Singleton, 2002). All having certainly been affected by allometry and scaling trends (Gilbert & Rossie, 2007).

4.8.1 Question 1: Are the extant Catarrhini genera adequately defined and do they compose an ‘adaptively coherent’ grouping of one or more species which occupy an ‘adaptive zone’, as defined by Wood & Collard (1999a)? Aside from Macaca, the remaining genera (Cercocebus, Mandrillus, Lophocebus, Papio and Theropithecus) are adequately defined, each adaptively coherent and occupying different adaptive zones. The type species of Macaca is M. sylvanus and on average has an intermembral index of ~ 86% (calculated from data presented in Strasser, 1992 and Ciochon, 1993). M. sylvanus is restricted to mountainous regions of Northern Africa (Morocco and Algeria) with a small, probably introduced, population in Gibraltar. M. sylvanus is a relict species, genetically, skeletally and behaviorally distinct from the other macaques in Eastern and Southeast Asia; and is perhaps more closely related to fossil forms recovered from circum-Mediterranean deposits than to species in East and Southeast Asia (Delson, 1980; Stanyon et al, 1980; Cronin et al, 1980; Fa, 1989; Zhang & Shi, 1993; Hoelzer & Melnick, 1996; Morales & Melnick, 1998; Harris, 2000; Thierry et al, 2000; Groves, 2001a; Deinard & Smith, 2001; Pan et al, 2003; Pan & Oxnard, 2004; Li & Zhang, 2005; Thierry, 2007). Furthermore, genetics and limb indices for macaques of Eastern and Southeast Asia also suggest complex evolutionary histories involving hybridizations and locomotor differences. As such, there are some species within Macaca that are more closely related to some species but not others. Subgenera would certainly better organize the species of Macaca. Metrical overlap may occur between Lophocebus and Cercocebus 170

but allometric analysis and 3-D morphometrics can separate these genera (Singleton, 2002, 2004a & 2004b; Gilbert, 2007).

Question 2: In terms of cranial morphology, how distinct are genera? In other words, how readily may a genus be determined from cranial dimensions, considered either singular or multiple? The crania of papionins are noticeably distinct from one another based simply on general appearance alone (Jolly, 2003). Several cranial dimensions are significantly correlated with body size (Table 4.15 and Figure 4.27). Cercocebus spp.: Overall, very small compared to other papionins with extreme facial elongation. Species may or may not develop deep sub-orbital fossae and cranial height (bas-br) and palatal length (ol-sta) on average less than 60 mm (Figures 4.2 and 4.13). The cranium is rather primitive with little ante-orbital drop, lacking major sub-orbital and facial fossae as seen in Lophocebus or Theropithecus and sagittal crests are not common; widely placed anterior temporal lines, upturned nuchal line morphology (both shared with Mandrillus) and no sagittal crest (Hill, 1974; Gilbert, 2007). Inferior cranial length between Cercocebus, Macaca and Lophocebus very similar (Figure 4.3). Two indices revealed similarities between Cercocebus and Mandrillus and different from the other papionins. These include biasterionic breadth relative to the occipital sagittal chord (biast/lam-opn) and the occipital sagittal chord in proportion to the cranial vault length (lam-opn/g-o) (Figures 4.16 and 4.24). Mandrillus spp.: Mandrills and drills are noted for extreme sexual dimorphism and paranasal swelling of the maxilla (i.e. lateral to the nasal bones) along the dental snout or muzzle (Elton & Morgan, 2006), both the consequence of intense male-to-male competition, sexual selection and predator and/or intraspecific defense. The Mandrill is one of the largest of the cercopithecines; adult males can reach up to ~40 kg. Adult males can develop a sagittal crests but it is placed more posteriorly on the neurocranium. Adult male Mandrills and baboons can have palatal lengths (ol-sta) greater than 100 mm (Figure 4.13). Macaca spp.: Macaques species have a generalized, primitive cranium (although some species are highly derived, e.g. M. nigra) that lacks dramatic facial and mandibular fossae, and extreme facial elongation although some species can be very sexually dimorphic. The more primitive macaque species (e.g. M. sylvanus, M. mulatta, M.

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fascicularis and M. nemestrina) most likely resemble the ancestral papionin cranium form (Delson, 1975 & 1980). There is a large range of variation for some Macaca indices, such as the relative proportion of the maxillo-alveolar breadth (biecm) to biasterionic breadth (biast); as well as the relative proportion of bizygomaxillare inferior breadth (bizi) to biasterionic breadth (biast) (Figures 4.18 and 4.22). Lophocebus spp.: Pseudo-mangabeys are highly arboreal and are the smallest if the papionins. Like Cercocebus, cranial height (bas-br) and palatal length (ol-sta) on average less than 60 mm. The cranium has short nasal bones, pinched or convergent anterior temporal lines, downturned nuchal line (both shared with Papio and Theropithecus) and deep sub-orbital fossae (Hill, 1974; Groves, 1978; Gilbert, 2007). Papio spp.: Like mandrills and drills, intense male-to-male competition and sexual selection have resulted in extreme sexual dimorphism between males and females, although survival requirements during the Plio-Pleistocene to the present and a savanna lifestyle must have also played a vital component. Unlike geladas, the skull of baboons and other papionins is klinorhyncous (Vogel, 1968). Similar to Mandrills, baboons are some of the largest of the cercopithecines; adult males >30 kg. Several dimensions overlap with those of Mandrillus, such as superior facial length, sagittal length of the nasal bones palatal breadth (bas-pros, nas-rhi and bienm; Figures 4.4, 4.7 and 4.12). However, relative percentages revealed some distinctions. In particular, the relative proportion of the maxillo- alveolar breadth (biecm) to the maximum length of the zygomatic bone (zs-zgyi) for the Papio sample was on average less (~97%) than the Mandrillus sample (112%) (Figure 4.19; as well as the indices mentioned above for Cercocebus; see Figures 4.16 and 4.24). Theropithecus gelada: The skull of geladas is very distinct and airorhynchous with a strong, continuous supra-orbital torus and marked muscle markings. Two indices which distinguish the gelada from other papionins included the relative percentage of external bicanine breadth (bicanex) to the length of the palate (ol-sta) (~50%) and the relative difference of the superior facial breadth (bifmt) to bizygomaxillare inferior breadth (bizi) (~88%) (Figures 4.17 and 4.20). Sagittal crests are common for adult males but the crest begins much more anteriorly compared to Papio or Mandrillus. In addition, the complex occlusal morphology of gelada molars is highly distinct from other papionins (Delson, 1975).

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Question 3: How much morphological variation is encompassed within a Catarrhine genus? Ranges or variation for the papionins are larger than the cercopithecins, certainly the result of body size increase and extreme sexual dimorphism. Morphological variation, either singular variables or indices, for papionin genera can produce coefficients of variation less than 5% to greater than 20% (Table 4.7). Highly variable measurements are associated with the nasal and facial region. In contrast, some neurocranial variables produce much less variation. Intra-generic Euclidean distances based on Ln transformed data are on average 1.0 (Max. - Min., Mandrillus, 1.41 - Lophocebus, 0.79) while those based on MSV were 0.66 (Max. - Min., Mandrillus, 0.78 - Theropithecus, 0.54) (Tables 4.17 and 4.21).

Question 4: How much has one genus morphologically and genetically diverged from another? With regard to morphological divergence, by measuring size or shape differences, Papio and Lophocebus produce the largest morphological distances (Euclidean distances and Cherry et al‘s M) while the smallest distances are between Lophocebus and Cercocebus (Tables 4.16, 4.20, 4.23, 4.26, & 4.28). The morphological distances generated from the papionin datasets are very similar those of the cercopithecins. Again, these distances are the result of at least 7 to 10 million years of evolution and half of this period is documented in the fossil record. However, inter-generic genetic distances are the least between Papio, Theropithecus and Lophocebus (0.012; Table 4.29), while the greatest genetic distance is between Papio and Cercocebus (0.026) (Page et al, 1999). The average inter-generic genetic distance is 0.02 (Range 0.014).

Question 5: Do cranial morphometric similarities and/or differences reflect adaptive zones? Cranial morphometrics and/or dispositions which may indicate the adaptive zones of these genera include the following. Cercocebus spp.: Mangabeys are small to medium African tropical forest papionins which, like Mandrillus, exploit the forest floor and lower strata of the forest (Rowe, 1996; Jolly, 2007). Mangabeys often associate with other primate species (guenons and colobines) for predator spotting and avoidance (McGraw, 1994 & 1998) (see also

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generic description for question 2 above and generic summaries below). Associated with this is a cranium with moderate facial projection and lacks a sagittal crest. The smaller size of the cranium, similar to Lophocebus and Macaca (compared to Papio or Mandrillus) also indicates use of, and compromise for, arboreal niches and/or resources. Mandrillus spp.: The Mandrill and Drill are geographically restricted to tropical forests of Western Africa. Like mangabeys, mandrills and drills forage for food along the forest floor (Jolly, 2007) (see also generic description for question 2 above and generic summaries below). The cranium of mandrills and drills are highly distinct. The larger size of the cranium and large canines, similar Papio and Theropithecus, indicates a more terrestrial lifestyle, which entails aggressive behavior between males and predator defense (see also generic description for question 2 above and generic summaries below). Macaca spp.: Macaques are the most widely distributed primates after humans. Macaque crania are generalized and probably approximate the basal papionin morphotype. Sagittal crests are present in large males. Crania in lateral profile do not have as steep ante- orbital drop seen in Papio or Theropithecus. Crania generally lack major facial and mandibular fossae (see also generic description for question 2 above and generic summaries below). Lophocebus spp.: Pseudo-mangabeys are highly arboreal papionins that prefer the middle to upper strata and rarely come down to the forest floor to forage (Rowe, 1996). Their smaller cranium size compared to genetically close genera (i.e. Papio and Theropithecus; see Disotell, 1994; Harris, 2000) indicates, like Macaca or Cercocebus, adaptations to exploiting arboreal niches and resources (see also generic description for question 2 above and generic summaries below). Papio spp.: Savanna baboons occupy many different habitats throughout sub- Saharan Africa (Kamilar, 2006) and baboons populations are genetically diverse (Rogers, 2000) The larger size of baboon crania ( similar to Mandrillus) is a evidence to their adaptation to and use of terrestrial habitats (see also generic description for question 2 above and generic summaries below). Theropithecus gelada: Geladas are highly derived, exclusively consume grass and are restricted to the mountainous highlands of Ethiopia but were once widespread throughout Africa and Eurasia during the Pliocene and Pleistocene. Associated with is a

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highly diagnostic cranium with strong muscle markings and sagittal crests. Aside from cranial morphology, the dentition of geladas is the best indicator of this genus’ adaptive zone (see also generic description for question 2 above and generic summaries below).

Question 6: What analogies may be drawn from extant Catarrhini genera for the interpretation of fossil hominin genera? Several analogies and considerations have been and continue to be drawn upon from the extant papionins and their generic arrangement for the interpretation of species- concepts, fossil hominin systematics and socioecology (Jolly, 1993 & 2001; Henzi & Barrett, 2005; Elton, 2006). First, perhaps the most striking phenomenon observed within the papionins is the degree of morphological convergence (or homoplasy; Collard & Wood, 2001a), which has important implications for the interpretation of fossil hominin species. Papio and Mandrillus both have converged on a similar design because of sexual and social selection and terrestrialism yet occurred independently of one another. Likewise, the species of Paranthropus, or paranthropines, are generally thought to comprise a monophyletic group (e.g. Strait et al, 1997). However, because most of the features which unite the paranthropines are restricted to mastication and craniodental features (i.e. all of these apparently derived traits are related to one functional complex), there is the possibility that these are convergences bought about via similar selective pressures and diet (see McCollum, 1999). Second, terrestrialism and the selective pressures it entails for survival results in increases to body size and facial elongation for the papionins, similar to the cercopithecins although not as large or extreme. Third, sexual selection and habitat preference are major components affecting papionin cranial morphology. Fourth, sister species (or genera) can have radically different phenotypes (e.g. Cercocebus and Mandrillus or Papio and Lophocebus).

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4.8.2 Generic Summaries: Tribe Papionini - 2n = 42. Based on genetic data Goodman et al (1998) suggest extant lineages emerged ~ 7 Ma. Groves (1978) lists several features which define this tribe. Some of these include, 1) Sexual swelling of female; 2) Facial elongation; 3) Molar flare; 4) M3 has a hypoconulid in all but smallest forms; 5) Females has incisiform canines; and 6) Facial mimicry (see also Groves, 1989 & 2001a). Cercocebus E. Geoffroy, 1812; Mangabeys. A polytypic genus comprising 6 species; C. atys [type species of genus; Sooty mangabey], C. torquatus (Collared or White- throated mangabey), C. agilis (Agile mangabey), C. chrysogaster (Gold-bellied mangabey), C. galeritus (Tana River mangabey) & C. sanjei (Sanje mangabey). In addition, mangabey- like fossil species have also been allocated to this genus or to another closely allied fossil genus and species from East and South Africa, particularly Parapapio and Procercocebus (Freedman, 1957; Leakey, 1988; Benefit, 2000; Delson et al, 2000; Gundling & Hill, 2000; Gilbert, 2007). Based on genetic data Goodman et al (1998) suggest Cercocebus lineages emerged ~ 4 Ma. Cercocebus and Mandrillus both specialize foraging on the forest floor and/or lower strata as well as the ability to process and consume hard fruits, seeds and/or nuts (McGraw & Fleagle, 2006; Jolly, 2007). Mandrillus Ritgen, 1824; Mandrill and Drill. 2 species, M. sphinx [type species of genus] & M. leucophaeus. The mandrill and drill are geographically restricted to equatorial- tropical West Africa and have no fossil record. In the past mandrills and drills were sometimes included within Papio (e.g. Wolfheim, 1983; Strasser & Delson, 1987) based largely on muzzle similarities and extreme sexual dimorphism. However, this classification scheme is not followed anymore due to the genetic and morphological evidence which clearly unites Mandrillus instead with Cercocebus (Groves, 1978; Disotell et al, 1992; Disotell, 1994; Fleagle & McGraw, 1999). A phylogenetic relationship so close and having diverged so recently that Goodman et al (1998) proposed Mandrillus should be a subgenus of Cercocebus. Recent genetic analysis by Telfer et al (2003) reveals a ‘deep’ genetic divergence reaching into the Early-Middle Pleistocene between particular geographical populations of Mandrillus sphinx. Macaca Lacepede, 1799; Macaques. A polytypic and versatile genus comprising 20 species clustered into six species-groups based largely on male genitalia morphology, ecology, pelage, vocalizations, (paleo)biogeography and behavior (see Groves, 2001a and references therein); 1 - M. sylvanus [type species of genus; Barbary macaque], (2 -

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Nemestrina-group - M. silenus (Lion-tailed macaque), M. nemestrina (Pig-tailed macaque), M. leonine (Northern pig-tailed macaque), M. pagensis (Mentawai macaque),) (3 - Sulawesi group - M. maura (Moor macaque), M. ochreata (Booted macaque), M. tonkeana (Tonkean macaque), M. hecki (Heck’s macaque), M. nigrescens (Gorontalo macaque), M. nigra (Crested black macaque),) (4 - Fascicularis-group - M. fascicularis (Long-tailed or Crab-eating macaque), M. arctoides (Stump-tailed or Bear macaque),) (5 - Mulatta-group - M. mulatta (Rhesus macaque), M. cylopis (Formosan rock macaque), M. fuscata (Japanese macaque),) (6 - Sinica-group - M. sinica (Toque macaque), M. radiata (Bonnet macaque), M. assamensis (Assamese macaque) & M. thibetana (Tibetan or Milne- Edward’s macaque)). Additionally, fossil species have also been allocated to this genus (Delson, 1980; Aimi, 1981; Pan & Yanzhang, 1995; Kohler et al, 2000). Genetic evidence shows that M. arctoides originated from a hybridization process involving M. assamensis/thibetana-like males with M. fascicularis-like females (Tosi et al, 2000 & 2003; Brandon-Jones et al, 2004; Morales & Melnick, 1998). Perhaps this should not be surprising because from India to the Eastern coast of China up to eight species of Macaca may overlap (Fooden, 1982), yet are divided by ecogeographic barriers minimizing competition. However, the monophyly of Macaca has been questioned (Groves, 1989 & 2000) and supported (Delson, 1985; Fa, 1989; Tosi et al, 2003). Macaque species have been described as a ‘weed’ species (Burney, 1995) referring to their adaptability and versatility to inhabit and colonize and different environments. Lophocebus Palmer, 1903; Pseudo-mangabeys. A polytypic genus comprising 4 species; L. albigena [type species of genus; Gray-cheeked pseudo-mangabey], L. aterrimus (Black pseudo-mangabey) & L. opdenboschi (Opdenbosch’s pseudo-mangabey). Not included in Groves taxonomy was the recently described L. kipunji (Jones et al, 2005), which was later placed in a new genus Rungwecebus (Davenport et al, 2006). Lophocebus was not used until quite recently but eventually removed from Cercocebus because of genetic and morphological data (Cronin & Sarich, 1976; Groves, 1978; Disotell, 1994 & 2000; Fleagle & Mcgraw, 1999 & 2002; Page & Goodman, 2001). In addition, fossil species have also been allocated to this genus by Harrison & Harris (1996), though their exact age is not known.

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Papio Erxleben, 1777; Savanna baboons. A polytypic genus comprising 5 species; P. hamadryas (Sacred baboon), P. papio [Guinea baboon; type species of Papio as ruled by the ICZN, see Groves 2001a and references therein;], P. anubis (Olive baboon), P. cynocephalus (Yellow baboon) & P. ursinus (Chacma baboon). Baboons are probably one of the most studied nonhuman primate species next to humans and macaques (either in 1) the laboratory; 2,) natural field observations - beginning in the early 1900s by Eugene Marais in South Africa, though not correctly published until 1969; or 3, the fossil record) and continue to produce invaluable information to the study of primate evolution (Cronin & Meikle, 1982; Jolly, 1993 & 2001; Rogers, 2000; Frost et al, 2003). Several baboon fossil species have also been allocated to this genus or to another closely allied fossil genus and species from Eastern and Southern Africa during the Plio-Pleistocene, especially Parapapio (Freedman, 1957; Szalay & Delson, 1979; Leakey, 1988; Pickford & Senut, 1988; McKee, 1993; McKee & Keyser, 1994; Harrison & Harris, 1996; Gundling & Hill, 2000; Delson et al, 2000; Jablonski, 2002). In addition, recently Liang et al (2006) reported the discovery of a fossil baboon mandibular fragment from the Yunnan Province, China, which would be the first occurrence outside sub-Saharan Africa and the Arabian Peninsula for Papio. Based on genetic data Goodman et al (1998) suggest Papio lineages emerged some 4 Ma. In another analysis, Newman et al (2004) demonstrated using mtDNA that Papio and Theropithecus diverged ~ 3.75 Ma. Then within Papio, P. ursinus diverged first ~ 1.79 Ma; next, P. papio ~ 1.37 Ma; followed by, P. hamadryas ~ .62 Ma; and finally P. anubis and P. cynocephalus ~ .16 Ma. Furthermore, Rogers (2000) has shown that the genetically (DNA and mtDNA) Papio is highly variable, both within and between populations. The cranial morphology of Papio spp. is largely the consequence of social behavior and adaptations for intra-sexual sexual competition and survival in terrestrial lifestyles. Baboons occupy a diverse range of habitat types with varying concomitant social structures (Henzi & Barrett, 2002 & 2005). Theropithecus I. Geoffroy, 1843; Gelada baboon. At present, a monotypic genus, T. gelada but many well documented fossil gelada species have also been allocated to this genus (which contains three subgenera; Theropithecus, Simopithecus and Omopithecus) from localities in Africa & Eurasia during the Plio-Pleistocene, including three subgenera (Jolly, 1970 & 1972; Leakey, 1988; Eck, 1993; Jablonski, 1993 & 2002; Delson, 1993;

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Gibert et al, 1995; Benefit, 1999 & 2000; Gundling & Hill, 2000; Delson et al, 2000; Elton, 2001; Jablonski et al, 2002; Rook et al, 2004). Oddly though, at present the gelada is restricted to the Ethiopian highland plateau and is the only extant primate species to eat grass habitually, which is reflected in their adaptive thick-enameled and complex molar occlusal morphology. In addition, the species’ pollex and high thumb index is quite different to other papionins, no doubt due to its adaptation to carefully plucking blades of grass to eat (Napier & Napier, 1967; Jolly, 1970 & 1972; Jablonski, 1993; Delson, 1993; Rowe, 1996). Despite these adaptations, Goodman et al (1998) proposed Theropithecus and Lophocebus be subgenera within Papio because of their recency of estimated genetic divergence (~ 4 Ma) and the adoption of time-ranked classifications. Of particular import are the observations and data presented by Jolly et al (1997; discussed earlier by Van Gelder, 1977) of natural intergeneric hybridization by Theropithecus gelada and Papio hamadryas, despite having been distinct genetic and paleontological lineages for nearly three to four million years (Jablonski, 2002; Newman et al, 2004).

179 Chapter 5: Results for genera of the Colobinae

Kasi Presbyticus Co. Pi. Pro. Se. Tr. Pre. Py. R. Si. N.

Presbytina Colobina Nasalina

Colobinae Cercopithecidae Cercopithecoidea Figure 5.1: A taxonomic and phylogenetic Catarrhini diagram representing the likely relationships Simiiformes within the Colobinae based on biomolecular Haplorrhini and morphological data. Primates

5.1 Introduction: The purpose of this chapter is to report the results for the Subfamily Colobinae (Jerdon, 1867). Although colobine monkeys were once described as the “forgotten leaf- eaters” (Groves, 1970), this is no longer the case. More importantly, many studies have shown that colobines have quite diverse diets and are not exclusively dependent on leaves to meet all nutrition requirements, as well as being ecologically and behaviorally flexible (Oates, 1994; Kirkpatrick, 2007). Where sympatry between colobine species does occur, competition is minimized by ecological niche separation as well as vegetation type and seasonality (e.g. African colobines see Clutton-Brock, 1974; Davies et al, 1999; Fashing, 2007; Asian langurs and leaf-monkeys see Hladik, 1977; Curtin, 1976; Kirkpatrick, 2007). In addition, these different dietary strategies and niche separation between colobine genera and species generally also translate into particular anatomical and limb proportion

distinctions (Fleagle, 1977; Schultz, 1986; Strasser, 1992). Tables 5.1 to 5.10 provide the key attributes describing the adaptive zones of the Colobinae genera.

Table 5.1: Colobus spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) (,; Average (range) Habitat Min.-Max.) 10870, Intermembral Above branch Folivore, can Arboreal & Multimale- 8123*; 79 (76-83) Quadrupedalism, subsist on some terrestrial Multifemale 6000-15500 Brachial with leaping & mature foliage behaviors/ & 1 male- 92 (88-98) some Rain, evergreen multifemale/ Crural semibrachiation & riverine 1.08 (C. 87 (85-93) forests guereza) (73.5-82.3g adult brain weight)

Table 5.2: Piliocolobus spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) (,; Average (range) Habitat Min.-Max.) 8692, Intermembral Above branch Folivore, food Arboreal & Multimale- 8210*; 86 (79-89) Quadrupedalism widely some terrestrial Multifemale & 5000-13500 Brachial with leaping & dispersed and behaviors/ 1 male- 94 (91-99) some diverse Primary & multifemale/ Crural semibrachiation secondary rain 1.17 (P. 86 (79-87) forest, dry preussi) (73.8g deciduous & adult brain riverine forests, weight - P. savanna badius) woodland – prefers middle to upper strata

Table 5.3: Procolobus verus Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) Average (range) Habitat (,; Min.-Max.) 4700, Intermembral Above branch Folivore, Arboreal/ 1 or 2 male- 4200; 80 Quadrupedalism selective feeder Swamps, rain multifemale/ 4000-6000 Brachial with leaping & forest and .97-1.04 92 (89-97) some semideciduous (57.8g adult Crural semibrachiation forest – prefers brain 91 lower to middle weight) strata and dense undergrowth

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Table 5.4: Semnopithecus spp. Body Weight Limb Indices: Locomotion Diet Substrate/ SS/EQ (g) (,; Average (range) Habitat Min.-Max.) 13000, Intermembral Above Branch Herbivore, Terrestrial & 1 male- 9890*; 79 (73-83) Quadrupedalism influenced by arboreal/ multifemale 6000-21000 Brachial with leaping seasonality Tropical, & 100 (97-104) subtropical, moist multimale- Crural or dry deciduous multifemale/ 86 (83-89) & alpine forests. .83 (135.2g adult brain weight - S. entellus)

Table 5.5: Trachypithecus (including Kasi) spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) (,; Average (range) Habitat Min.-Max.) 8813, Intermembral Above Branch Folivore, Arboreal & 1 male- 7505*; 82 (78-87) Quadrupedalism can subsist some multifemale 6000-20000 Kasi - 76 (75-79) with leaping of mature terrestrial/ & Brachial foliage; Tropical & multimale- 94 (88-99) influenced subtropical multifemale/ Kasi - 101 (98-104) by primary & .62-.92 (64- Crural seasonality secondary 85g* adult 86 (82-89) coastal forests, brain Kasi - 89 (88-90) moist weight) evergreen, mangrove, bamboo, karst, montane & deciduous forests up to 3500m asl

Table 5.6: Presbytis spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) (,; Average (range) Habitat Min.-Max.) 6310, Intermembral Above Branch Frugivore & Arboreal & Monogamous, 6244*; 76 (73-79) Quadrupedalism Folivore – limited 1 male- 5000-8000 Brachial with leaping seeds large part terrestrial/ multifemale 109 (99-115) of diet; Tropical, & multimale- Crural influenced by primary & multifemale/ 89 (86-94) seasonality secondary, 1.03-1.06, swamp, 1.2-1.22 (80- mangrove, 92.7g* adult montane forests brain weight)

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Table 5.7: Pygathrix spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) (,; Average (range) Habitat Min.-Max.) 11000, Intermembral Above branch Folivore Arboreal/ 1 male- 8440*; 89 (85-91) Quadrupedalism Tropical, multifemale 8000-12000 Brachial and some evergreen, & 100 (98-103) brachiation moist deciduous multimale- Crural (Byron & forests, high multifemale/ 89 (86-92) Covert, 2004) canopy .91-1.15 (108.5g adult brain weight - P. nemaeus)

Table 5.8: Rhinopithecus (including Rhinopithecus & Presbyticus) spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) (,; Min.- Average (range) Habitat Max.) 15675, Intermembral Above branch Folivore; Terrestrial & 1 male- 10020*; 95 (94-97) Quadrupedalism influenced by Arboreal/ multifemale 8000-35000 Brachial with leaping and seasonality Temperate to & bachelor 99 (96-103) semibrachiation Subtropical groups/ Crural primary, .69-.94 85 (83-90) deciduous, (121.5g bamboo, adult brain coniferous, weight - R. montane, broad- roxellana) leaf forests up to 4500m asl.

Table 5.9: Nasalis larvatus Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) (,; Average (range) Habitat Min.-Max.) 20400, Intermembral Above branch Folivore Arboreal & 1 male- 9820; 93 (89-97) Quadrupedalism Terrestrial multifemale 9000-30000 Brachial with leaping and behaviors/ & bachelor 100 (96-104) climbing Tropical forest, groups/ Crural mangrove 1.24 (94.2g 86 (83-91) adult brain weight)

Table 5.10: Simias concolor Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) (,; Average Habitat Min.-Max.) 9150, Intermembral Above branch Folivore Arboreal & Varied, 6800; 98 Quadrupedalism Terrestrial/ monogamous, 6000-11000 Brachial Tropical forest, multimale- 104 evergreen & multifemale; Crural deciduous 1 male- 83 forests multifemale/ ?

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Unfortunately, many colobines are at present critically endangered. The extant subfamily Colobinae encompasses ten genera, Colobus, Piliocolobus, Procolobus, Semnopithecus, Trachypithecus, Presbytis, Pygathrix, Rhinopithecus, Nasalis and Simias, which together comprise 59 described species (Groves, 2001a); and together with the Cercopithecinae, form the family Cercopithecidae, which in turn along with Victoriapithecidae compromise the Cercopithecoidea (Delson, 1975; Andrews, 1981; Benefit, 2000; Benefit & McCrossin, 1997 & 2002). Please note, however, that Groves (1989) and Jablonski (2002) argued that the colobines should be given familial status, Colobidae. The colobines are the largest comparative grouping within this study. The average number of species per colobine genus is 5.9, ranging from one to 17 with a standard deviation of 5.2. However, when broken down into subtribes the average number of species per genus changes. The average number of species per genus for Colobina is 5.0, ranging from one to nine with a standard deviation of 4.0. The average number of species per genus for Presbytina is 11.7, ranging from seven to seventeen with a standard deviation of 5.0 (however, if Trachypithecus is excluded, the average number of species for Semnopithecus and Presbytis is 9, ranging from seven to eleven with a standard deviation of 2.8). Finally, the average number of species per genus for Nasalina is 2.3, ranging from one to four with a standard deviation of 1.5. The species of colobine genera range in size from less than five kilograms (Procolobus verus, the smallest living colobine) up to 20-35 kg (Nasalis larvatus and Rhinopithecus spp.). Delson (1975 & 1994) suggested different subtribes for the African and East Asian genera, Colobina and Presbytina, which are useful taxonomic divisions and are adopted here for comparative boundaries, although another subtribe may prove useful based on genetic data and soft-tissue nasal anatomy, Nasalina (Groves, 2001a; Sterner et al, 2006; Whittaker et al, 2006). Thus, the Colobinae genera can be grouped into three subtribes based on geography, soft-tissue nasal anatomy, morphology and genetics (Figure 5.1). The subtribes include, Colobina for the African colobines – Colobus (Black-and- White colobines), Piliocolobus (Red colobines) and Procolobus verus (Olive colobine); Presbytina for the Asian langurs and leaf monkeys – Semnopithecus (Hanuman langurs), Trachypithecus (Lutungs) and Presbytis (Surilis); and Nasalina for the odd-nosed colobines

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- Pygathrix (Douc langurs), Rhinopithecus (Snub-nosed monkeys), Nasalis larvatus (Proboscis monkey) and Simias concolor (Simakobu or Pig-tailed langur). The Colobinae is an ancient primate lineage that diverged from the Cercopithecinae some ~14 Ma during the Miocene (Delson, 1994). In fact, colobines have been found alongside some of the earliest hominin species (White et al, 2006; Hlusko, 2006). Colobine monkeys are unique because of their multi-chambered stomach containing bacterial flora which detoxifies foliage and facilitates the digestion of leaves. This dietary adaptation permits colobines to exploit a resource which is widespread and in some regions, readily available year-round. Of course, many other primates do occasionally consume leaves but in lower quantities, with other food items, and in general is a fall-back resource exploited during seasons when invertebrates, fruits, flowers and/or seeds are scarce. In addition to their digestive adaptations, the teeth of colobines have high-pointed cusps with shearing crests which aid in the breaking-down of leaf vegetation (Lucas & Teaford, 1994). Due to their dietary affinity towards leaves, it is not surprising that many colobines are largely arboreal and restricted to tropical-equatorial evergreen habitats, but this has not always been the case and there are exceptions to these generalizations. For most of the Plio- Pleistocene colobines enjoyed a much wider geographic range extending from Europe to North and South Africa, some exhibiting terrestrial adaptations reflected in the postcranium skeleton (e.g. Cercopithecoides from East and South Africa). Furthermore, associated with their arboreal habitats, many colobines also display some degree of thumb shortening and foot lengthening, both of which aid travel in the canopy, involving above branch quadrupedalism, leaping and some semibrachiation. Lastly, most species of the Colobinae have 44 chromosomes but Nasalis, which is cranially one of the most distinct and sexually dimorphic of the colobines, has 48 (Bigoni et al, 2003). With regard to possible taxonomic schemes, each subtribe may be arranged differently. First, Groves (2001a) recognizes three genera within the Colobina and is followed here. Researchers have abandoned earlier classificatory schemes in which Colobus encompassed all the African colobines (e.g. Wolfheim, 1983); as well as the subgeneric placement of Piliocolobus within Colobus (e.g. Napier, 1985). Strasser & Delson’s (1987) cladistic analysis of the Cercopithecidae revealed a closer relationship between the Olive and Red colobines due to shared features such as a four-chambered

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stomach (as opposed to three in the Black-and-white colobines), unique male perineal organ morphology and discontinuous ischial callosities (Black-and-white colobines also have an inter-related enlarged larynx and sub-hyoid sac). Thus, some place both the Olive and Red colobines within Procolobus but separated subgenerically (e.g. Delson et al, 2000) but still, primitive traits and behaviors of the former and species diversity of the latter persuades many researchers to stress subgeneric allocations (Oates et al, 1994; Oates, 1994). Even so, evidence presented by O’Higgins & Pan (2004) demonstrates similar facial growth patterns between Colobus and Piliocolobus but this could be the result of similarities in body size. Secondly, the Presbytina of South Asia and Southeast Asia have had a complicated taxonomic history. For most of the 20th century, the genus Presbytis encompassed all non- odd-nosed langurs divided subgenerically (e.g. Szalay & Delson, 1979). This has since been abandoned by most researchers but can still be found in recent literature (e.g. “Presbytis entellus”, Ziegler et al, 2000) despite phenetic and phylogenetic evidence suggesting otherwise. If this generic classification scheme is upheld using Groves’ taxonomy (2001a), “Presbytis” would contain 35 species. Some important cranial differences between species of Presbytis and Trachypithecus include (see Hooijer, 1962, p. 21 - citing Lyon, 1907): In Presbytis species the anterior nares contracted to a point antero-inferiorly whereas those of Trachypithecus have nares that gradually taper to a point antero-inferiorly; Presbytis species have weak superciliary ridge whereas those of Trachypithecus are well marked; Presbytis species have a well-marked arch under the malo-maxillary suture whereas those of Trachypithecus do not; Presbytis species exhibit less post-orbital constriction whereas those of Trachypithecus have considerable post-orbital constriction; Presbytis species have a well-marked swelling of the braincase just inferior to the lambdoid suture whereas those of Trachypithecus do not; Presbytis species have a short palate but those of Trachypithecus have a much longer palate;

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Presbytis species have a less pronounced rostrum (i.e. very little facial projection) whereas that of Trachypithecus is more pronounced (i.e. greater facial projection); and lastly, The mandibular ramus of Presbytis species is shallow and the angular process is not enlarged whereas those of Trachypithecus have a deep mandibular ramus and an enlarged angular process. Further distinctions between these genera have been researched and reported by Weitzel et al (1988). The genus Trachypithecus contains two species given generic or subgeneric status by some researchers, Kasi vetulus and K. johnii (e.g. Hill, 1934 & 1936; Fleagle, 1998). Brandon-Jones (e.g. 1984 & 1996) instead places all species of Trachypithecus within Semnopithecus due to a lack of derived traits which the former shares to the exclusion of the other, as well as hybridizations between species in southern India (Brandon-Jones, 2004), and are thus lumped together as sister groups in relation to Presbytis, the more apomorphic of the three (Groves, 1989 & 2001a). However, this would result in a genus with 24 species (using Groves, 2001a; one less than Cercopithecus and three more than Macaca and both genera which are in need of revision) and genera with this many species are dubious. Furthermore, taking into consideration the number of species already within Trachypithecus (n=17; again, using Groves, 2001a taxonomy) as well as the genetic evidence (discussed below), this genus is most likely paraphyletic as arranged by Groves (2001a) and is in need of revision. Finally, the odd-nosed colobines of Nasalina located in East Asia and Southeast Asia, like the other colobines, also have had a confusing taxonomic history and there are still differing opinions as how they should be classified. Previously, Groves (1989) placed Nasalis (including Simias) in its own subfamily but has since amended this taxonomic scheme (2001a). Due to the strong influence of cladistic theory and phylogenetic classifications, evidence based on soft-tissue nasal anatomy and cranial morphology, species of Rhinopithecus are sometimes grouped within Pygathrix, while the Simakobu, Simias concolor, is subsumed within Nasalis (e.g. Delson et al, 2000). Furthermore, R. avunculus is sometimes given subgeneric status, Presbyticus, because of features such as relatively long thumbs and tail and habitat preference (e.g. Zhang & Ryder, 1998; Chaplin & Jablonski, 1998; Jablonski, 1998; Boonratana & Canh, 1998).

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5.2 Descriptive Statistics and Univariate Analyses for cranial variables: Table 5.11 provides the descriptive statistics for Colobinae genera. Like the previous chapters the nasal region and its configuration generate the largest coefficients of variation; in particular, inferior breadth of the nasal bones (inbrnabo) and maximum width of the nasal aperture (maxnawi), sagittal height of the nasal aperture (rhi-ns) and palatal height (palhei). These measurements have large CVs because of large absolute size differences between males and females (i.e. sexual dimorphism). Despite this, other dimensions such as cranial height (bas-br), biasterionic breadth (biast), inferior cranial height (bas-nas) or cranial vault length (g-o) show lower variation despite sexual dimorphism and inter-specific samples. The box-plots for the Colobinae, similar to the results for the cercopithecins and papionins, display many remarkable scaling trends related to body size. Figures 5.2 to 5.16 are box-plots which illustrate these trends. Genera are arranged from smallest to largest generic mean body weight per subtribe (Procolobus, Piliocolobus, Colobus; Presbytis, Trachypithecus, Semnopithecus; and Simias, Pygathrix, Rhinopithecus, Nasalis; same format for cranial indices as well, see below). In particular, bizygomatic breadth (bizygo), cranial vault length (g-o) and palatal length A (ol-sta) exhibit clear relationships with body size (Figures 5.10, 5.11 and 5.14). Variables which noticeably highlight marked differences between genera include the long face (nas-pros), lengthy nasal bones (nas-rhi) and narrow upper face (bifmt) of Simias and Nasalis (Figures 5.12, 5.13 and 5.7). In contrast, the latter measurement also demonstrates the rather wide upper face of Pygathrix, Rhinopithecus and Semnopithecus. In addition, the bizygomaxillare superior breadth (bizs) for Pygathrix is extremely wide compared to the other colobines (Figure 5.9). Furthermore, this dimension (bizs) also highlights a dramatic difference for Simias, which is much narrower than other colobines (however, the sample size (n=7) is small and only two males so, as mentioned previously, results for this genus should be considered approximates and will be investigated with larger samples).

5.2.1 Shapiro-Wilk results for cranial variables: Table 5.12 lists the results of applying Shapiro-Wilk to several cranial dimensions of the Colobinae. Most cranial variables for the Colobina and Presbytina may be assumed to be normally distributed. On the other hand, several variables for the Nasalina were not

188

Table 5.11: Colobinae pooled sex. g-o pros-o biast bipor biaur bizygo bifmo bifmt bizs bizi zs-zi zs-zgyi Colobus (n=26) Mean 81.9 112.1 46.7 58.9 64.7 79.9 55.2 60.9 41.0 53.9 14.1 37.0 Median 82.4 112.3 48.0 59.1 65.2 80.5 54.6 60.0 40.4 53.9 13.9 37.1 Maximum 89.0 127.1 54.7 66.0 70.0 88.0 61.8 69.5 51.0 62.0 16.3 42.6 Minimum 75.0 99.9 38.0 53.0 58.0 71.0 48.4 49.4 34.8 45.0 11.3 31.8 Range 14.0 27.2 16.7 13.0 12.0 17.0 13.5 20.1 16.2 17.0 5.0 10.8 SD 4.0 7.3 4.2 3.5 3.6 5.2 3.5 5.7 4.0 4.1 1.4 3.2 CV 4.9 6.5 8.9 5.9 5.6 6.5 6.4 9.3 9.7 7.6 9.9 8.6

Pilio. (n=28) Mean 79.3 102.9 44.4 55.8 62.6 79.4 52.6 61.6 32.8 49.8 15.6 37.2 Median 79.5 103.0 43.8 56.5 63.0 80.0 52.4 61.2 33.2 50.0 15.8 38.0 Maximum 89.0 116.0 52.0 62.0 72.0 93.0 57.0 70.4 37.3 58.0 20.0 43.3 Minimum 72.0 91.0 38.0 46.0 54.0 70.0 45.2 52.7 26.5 39.0 12.4 31.0 Range 17.0 25.0 14.0 16.0 18.0 23.0 11.8 17.7 10.8 19.0 7.6 12.3 SD 4.7 7.0 3.2 4.2 4.2 6.8 2.6 4.9 3.3 4.6 2.2 3.6 CV 5.9 6.8 7.2 7.6 6.7 8.5 5.0 7.9 10.2 9.2 13.8 9.5

Proc. (n=5) Mean 72.6 89.0 40.0 50.4 56.8 70.8 47.2 54.4 32.5 44.4 13.2 34.0 Median 71.0 89.0 39.5 50.0 58.0 72.0 46.8 54.9 32.6 44.0 13.2 33.6 Maximum 77.0 92.0 42.8 53.0 60.0 77.0 50.2 60.1 35.6 48.1 13.9 38.9 Minimum 69.0 83.0 37.8 47.0 54.0 66.0 44.9 50.2 30.0 42.7 12.5 31.3 Range 8.0 9.0 5.0 6.0 6.0 11.0 5.3 9.9 5.6 5.4 1.4 7.6 SD 3.6 3.7 1.9 2.3 2.7 4.8 1.9 4.1 2.2 2.2 0.5 3.0 CV 5.0 4.1 4.8 4.6 4.7 6.7 4.0 7.4 6.8 5.0 3.9 8.8

189

g-o pros-o biast bipor biaur bizygo bifmo bifmt bizs bizi zs-zi zs-zgyi Semno. (n=9) Mean 89.7 116.4 46.5 65.3 73.2 92.3 60.6 66.2 35.7 63.2 19.7 40.1 Median 95.0 117.0 47.3 65.0 75.0 96.0 61.2 65.2 36.6 65.4 19.1 39.7 Maximum 98.0 133.0 50.0 73.0 81.0 101.0 65.7 75.1 40.3 70.5 26.0 48.0 Minimum 77.0 98.0 40.7 54.0 61.0 80.0 54.9 57.4 29.5 53.5 12.2 29.5 Range 21.0 35.0 9.3 19.0 20.0 21.0 10.8 17.7 10.9 16.9 13.8 18.5 SD 8.0 11.9 3.0 7.4 7.5 7.6 4.1 5.4 3.3 6.9 5.4 7.3 CV 8.9 10.2 6.5 11.4 10.3 8.2 6.7 8.1 9.2 10.9 27.4 18.1

Trachy. (n=33) Mean 76.7 96.1 41.9 55.1 62.3 74.6 52.4 57.1 34.7 50.2 14.2 30.4 Median 76.0 96.0 41.0 54.0 62.0 74.0 50.9 57.0 34.1 50.1 13.8 30.0 Maximum 88.0 109.0 50.0 65.2 73.0 84.0 63.0 70.0 49.1 59.1 20.8 39.2 Minimum 68.4 85.9 35.6 49.0 54.1 65.9 46.2 47.2 25.9 44.8 11.5 25.7 Range 19.6 23.1 14.4 16.2 18.9 18.1 16.8 22.8 23.2 14.4 9.4 13.5 SD 4.9 6.5 4.2 4.5 4.3 4.7 4.3 4.5 5.7 4.1 1.9 2.9 CV 6.3 6.7 10.0 8.1 6.8 6.4 8.3 7.9 16.3 8.2 13.4 9.6

Pres. (n=18) Mean 74.3 90.7 43.8 53.2 60.2 69.7 54.2 58.0 34.5 48.4 14.7 31.7 Median 73.5 91.2 44.6 53.0 59.9 68.8 54.6 58.4 34.0 48.2 14.5 31.4 Maximum 80.0 99.0 50.0 59.0 66.0 77.0 58.2 63.2 42.2 54.0 20.4 38.6 Minimum 69.3 85.0 36.5 49.0 56.0 65.0 49.0 51.3 28.6 42.0 10.8 24.4 Range 10.7 14.0 13.5 10.0 10.0 12.0 9.2 11.9 13.5 12.0 9.6 14.2 SD 3.5 3.8 3.4 2.4 2.5 3.4 2.2 3.8 4.5 3.0 2.9 3.4 CV 4.7 4.2 7.8 4.4 4.2 5.0 4.1 6.5 13.0 6.3 19.9 10.7

190

g-o pros-o biast bipor biaur bizygo bifmo bifmt bizs bizi zs-zi zs-zgyi Pyga. (n=25) Mean 83.5 107.1 50.7 59.2 66.7 77.8 63.4 69.6 44.3 52.7 14.3 31.7 Median 84.0 106.0 50.0 60.0 67.0 80.0 63.9 70.3 44.2 52.7 14.5 31.6 Maximum 89.0 119.0 57.2 70.0 75.0 84.0 70.2 77.0 50.2 58.3 16.6 35.1 Minimum 72.7 91.0 41.2 49.0 57.0 64.0 54.8 60.0 35.2 45.3 11.7 28.4 Range 16.3 28.0 16.0 21.0 18.0 20.0 15.4 17.0 15.0 13.0 5.0 6.7 SD 4.1 7.7 3.7 4.7 3.8 5.8 3.8 4.1 4.0 3.1 1.1 1.9 CV 4.9 7.2 7.4 8.0 5.7 7.5 6.1 6.0 9.1 5.8 7.6 5.9

Rhino. (n=17) Mean 90.2 109.5 54.4 62.6 72.5 85.2 65.5 72.4 41.2 58.2 17.7 37.4 Median 90.0 110.0 54.3 62.0 72.0 86.0 66.1 73.0 41.8 58.9 17.5 36.5 Maximum 99.0 126.0 65.0 71.0 79.0 100.0 71.7 80.0 45.4 64.2 21.3 44.9 Minimum 80.5 96.4 46.0 53.6 64.9 70.8 58.1 61.8 34.0 47.1 13.3 31.1 Range 18.5 29.6 19.0 17.4 14.1 29.2 13.6 18.2 11.3 17.1 8.0 13.7 SD 4.2 7.0 4.6 4.1 3.7 7.4 3.3 4.6 2.8 3.9 1.9 3.4 CV 4.7 6.4 8.5 6.6 5.1 8.7 5.1 6.3 6.9 6.7 10.7 9.2

191

g-o pros-o biast bipor biaur bizygo bifmo bifmt bizs bizi zs-zi zs-zgyi Nasalis (n=28) Mean 87.6 118.6 49.5 61.6 69.2 83.9 55.8 61.8 31.4 54.8 20.5 38.8 Median 88.6 123.3 50.0 63.0 70.3 85.5 55.6 61.4 30.6 54.1 20.1 39.9 Maximum 94.0 132.0 56.6 69.0 77.0 95.0 60.5 68.4 38.1 62.8 25.7 45.1 Minimum 79.0 100.0 42.4 50.0 58.0 69.0 49.6 52.9 25.5 47.5 16.3 32.2 Range 15.0 32.0 14.2 19.0 19.0 26.0 10.9 15.5 12.6 15.3 9.4 12.9 SD 4.7 10.7 3.8 4.9 5.0 7.7 3.1 4.7 3.2 4.1 2.9 4.2 CV 5.4 9.0 7.7 7.9 7.2 9.2 5.6 7.7 10.3 7.5 14.0 10.9

Simias (n=7) Mean 73.9 95.3 42.8 49.1 57.7 68.3 50.7 55.4 23.5 47.6 18.5 33.3 Median 74.0 94.0 44.0 48.0 57.0 69.0 51.6 55.3 23.8 47.2 18.7 32.7 Maximum 78.0 106.0 44.2 54.0 62.0 74.0 52.6 60.5 25.0 50.8 21.0 37.8 Minimum 70.0 88.0 39.0 45.0 54.0 61.0 48.0 50.5 21.0 44.3 16.5 29.9 Range 8.0 18.0 5.2 9.0 8.0 13.0 4.6 10.0 4.0 6.5 4.5 7.9 SD 2.4 6.4 1.9 3.2 2.9 4.3 1.8 3.5 1.4 2.6 1.5 2.7 CV 3.3 6.7 4.4 6.6 5.0 6.4 3.6 6.2 6.1 5.4 8.3 8.2

192

Table 5.11 continued: Colobinae pooled sex. bas-pros bas-nas bas-br nas-pros nas-br br-lam lam-opn nas-rhi inbrnabo rhi-ns maxnawi nas-ns Colobus (n=26) Mean 87.0 68.9 50.8 41.1 50.0 37.9 26.7 13.7 8.5 21.8 12.4 35.5 Median 86.0 69.1 50.7 39.9 49.1 37.5 26.7 14.0 8.3 21.7 12.2 35.9 Maximum 100.7 77.6 58.0 49.4 57.9 45.8 30.3 17.1 11.4 28.3 17.3 41.0 Minimum 77.3 61.0 44.5 34.4 45.7 30.3 23.4 9.6 6.5 18.1 10.0 29.2 Range 23.4 16.6 13.5 15.1 12.3 15.6 6.9 7.5 4.9 10.2 7.3 11.9 SD 6.8 4.2 2.3 4.0 3.3 4.1 2.3 1.8 1.3 2.2 1.9 2.8 CV 7.9 6.1 4.6 9.8 6.7 10.7 8.5 13.3 15.0 10.3 15.1 7.9

Pilio. (n=28) Mean 75.3 62.6 50.7 38.8 50.2 35.4 27.7 14.4 7.3 19.5 10.2 33.6 Median 75.5 63.0 51.0 38.9 50.4 35.4 28.0 14.2 7.3 19.3 10.6 33.7 Maximum 88.0 68.0 60.0 44.4 55.3 44.3 33.2 19.0 9.1 22.3 13.7 41.0 Minimum 64.0 54.0 45.0 31.2 45.3 26.3 22.4 9.9 5.6 16.0 7.2 28.0 Range 24.0 14.0 15.0 13.2 10.0 18.0 10.8 9.1 3.5 6.3 6.5 13.0 SD 5.3 3.4 3.5 3.3 2.7 4.6 2.6 2.5 0.9 1.5 1.6 3.2 CV 7.1 5.5 7.0 8.5 5.4 13.0 9.4 17.7 12.0 7.7 15.8 9.6

Proc. (n=5) Mean 61.4 56.0 45.0 32.2 46.3 33.9 24.6 11.9 6.9 16.3 8.7 27.7 Median 62.0 57.0 45.0 32.0 46.1 34.5 24.7 12.5 6.9 16.5 9.0 27.7 Maximum 64.0 59.0 47.0 35.4 51.4 36.4 27.7 13.4 8.2 17.3 10.1 29.2 Minimum 58.0 53.0 43.0 28.7 42.0 30.0 22.4 9.8 5.6 14.9 7.3 26.0 Range 6.0 6.0 4.0 6.7 9.3 6.4 5.3 3.6 2.6 2.5 2.9 3.2 SD 2.4 2.4 1.6 2.4 3.5 2.6 2.1 1.4 1.0 0.9 1.1 1.1 CV 3.9 4.4 3.5 7.5 7.5 7.8 8.7 12.0 13.9 5.7 12.1 4.1

193

bas-pros bas-nas bas-br nas-pros nas-br br-lam lam-opn nas-rhi inbrnabo rhi-ns maxnawi nas-ns Semno. (n=9) Mean 82.2 68.6 55.6 43.1 57.5 37.8 32.5 14.8 7.3 22.7 10.1 37.9 Median 83.0 71.0 56.0 42.3 57.9 37.8 32.3 14.2 7.4 22.5 10.3 35.2 Maximum 91.0 74.0 61.0 56.0 64.3 44.0 36.2 20.6 8.8 26.9 11.9 50.0 Minimum 68.0 59.0 51.0 34.7 48.0 31.5 29.5 11.6 5.7 18.0 8.5 30.6 Range 23.0 15.0 10.0 21.3 16.3 12.5 6.7 8.9 3.1 8.9 3.4 19.4 SD 8.5 5.6 3.7 6.6 5.8 4.3 2.4 3.0 0.9 3.0 1.1 6.3 CV 10.4 8.2 6.6 15.4 10.1 11.3 7.4 20.1 11.8 13.3 11.1 16.7

Trachy. (n=33) Mean 69.6 59.9 49.4 32.1 50.9 33.2 27.4 10.7 6.4 17.3 8.6 27.8 Median 70.0 60.0 49.0 33.0 50.0 33.4 26.9 10.8 6.4 17.5 8.7 28.0 Maximum 79.0 67.0 56.0 38.4 63.2 42.0 34.6 15.0 9.5 21.5 11.2 31.8 Minimum 61.0 53.6 43.7 26.4 44.2 24.3 22.3 6.0 4.0 14.2 7.0 23.5 Range 18.1 13.4 12.3 12.0 18.9 17.7 12.3 8.9 5.5 7.3 4.2 8.4 SD 4.5 3.3 3.6 2.7 4.5 3.6 3.0 2.0 1.3 1.8 1.0 2.2 CV 6.4 5.5 7.3 8.5 8.8 10.8 10.8 18.5 20.1 10.7 11.7 7.8

Pres. (n=18) Mean 62.5 58.0 48.4 27.3 51.3 33.4 28.4 10.0 6.6 14.9 9.8 24.7 Median 62.2 58.1 48.3 26.7 51.6 33.4 28.2 10.0 6.6 14.7 9.8 24.6 Maximum 71.0 62.0 52.0 30.5 55.0 39.1 33.0 11.8 7.6 17.9 12.0 27.3 Minimum 58.0 52.0 45.0 24.7 48.4 27.9 24.7 8.5 5.6 13.1 7.7 23.1 Range 13.0 10.0 7.0 5.8 6.6 11.2 8.3 3.3 2.0 4.8 4.3 4.2 SD 3.4 2.8 1.8 1.6 2.0 3.1 2.1 0.9 0.6 1.2 0.9 1.2 CV 5.4 4.9 3.8 6.0 4.0 9.4 7.3 8.7 9.3 7.7 9.3 4.9

194

bas-pros bas-nas bas-br nas-pros nas-br br-lam lam-opn nas-rhi inbrnabo rhi-ns maxnawi nas-ns Pyga. (n=25) Mean 76.8 64.0 52.7 37.8 56.0 36.0 29.9 13.3 5.5 20.1 11.8 32.9 Median 77.0 64.0 52.0 37.1 55.6 35.9 29.9 13.3 5.8 20.2 11.7 32.5 Maximum 87.0 71.0 58.0 46.1 61.5 40.1 33.4 17.5 8.0 23.6 14.3 39.2 Minimum 66.7 56.5 47.0 28.3 49.0 29.4 27.1 8.2 2.6 15.0 8.0 27.9 Range 20.3 14.5 11.0 17.8 12.5 10.7 6.3 9.3 5.4 8.6 6.3 11.2 SD 6.6 4.2 3.1 4.4 3.1 2.7 1.6 2.3 1.3 1.8 1.5 2.7 CV 8.6 6.5 5.9 11.7 5.6 7.5 5.4 17.2 23.5 9.1 12.4 8.3

Rhino. (n=17) Mean 75.4 65.8 53.7 34.9 63.0 36.6 30.5 9.9 5.3 20.8 13.9 29.2 Median 76.0 65.0 54.0 34.0 63.4 36.8 31.1 8.9 5.0 21.0 13.8 29.0 Maximum 89.0 75.0 58.0 44.7 67.7 41.1 34.4 16.4 8.6 23.1 16.5 37.6 Minimum 63.0 58.1 50.7 29.0 54.7 33.1 23.1 5.0 2.9 18.7 11.0 24.1 Range 26.0 16.9 7.4 15.7 12.9 8.0 11.3 11.4 5.7 4.5 5.5 13.5 SD 6.5 4.2 1.7 4.9 2.7 2.1 2.9 2.9 1.5 1.2 1.4 3.5 CV 8.7 6.4 3.1 14.1 4.3 5.7 9.4 29.0 27.7 5.6 10.1 12.0

195

bas-pros bas-nas bas-br nas-pros nas-br br-lam lam-opn nas-rhi inbrnabo rhi-ns maxnawi nas-ns Nasalis (n=28) Mean 85.5 67.8 51.8 48.5 59.5 33.7 28.1 22.6 7.3 21.2 13.3 43.9 Median 89.6 70.0 52.0 50.7 60.0 33.3 28.1 22.4 7.2 22.2 13.4 45.1 Maximum 98.0 76.0 57.0 59.4 64.2 37.8 32.9 30.0 11.0 26.7 16.7 53.7 Minimum 71.0 59.0 48.0 36.2 54.1 30.3 23.3 17.1 4.4 14.9 9.8 33.1 Range 27.0 17.0 9.0 23.3 10.1 7.5 9.6 12.9 6.6 11.8 6.8 20.5 SD 9.0 5.2 2.3 7.0 2.7 1.9 2.4 3.7 1.7 3.6 1.9 6.2 CV 10.6 7.7 4.4 14.4 4.5 5.7 8.5 16.4 23.6 16.8 14.2 14.1

Simias (n=7) Mean 67.6 57.0 44.1 37.5 49.3 24.5 27.1 17.2 5.6 17.0 10.6 34.2 Median 66.0 56.0 44.0 36.8 49.6 24.2 27.9 18.0 5.7 16.5 10.4 33.3 Maximum 76.0 62.0 46.0 44.3 51.9 28.5 28.5 20.0 7.1 20.2 11.6 39.2 Minimum 61.0 54.0 42.0 31.1 46.6 21.8 25.1 14.3 4.7 14.6 9.2 29.9 Range 15.0 8.0 4.0 13.2 5.3 6.7 3.4 5.7 2.4 5.5 2.5 9.3 SD 5.7 3.5 1.7 4.7 1.6 2.5 1.2 2.1 0.8 2.1 0.9 3.7 CV 8.4 6.2 3.8 12.6 3.3 10.0 4.6 12.1 14.3 12.3 8.5 10.9

196

Table 5.11 concluded: Colobinae pooled sex. maxalvlen biecm iv-pms i1i2 bicanin bicanex palhei biseptal bienm ol-sta ol-pms bien Colobus (n=26) Mean 47.0 36.5 20.6 11.7 17.4 32.1 7.8 20.1 19.8 43.9 31.3 41.7 Median 46.8 37.0 20.2 11.7 16.8 32.3 8.0 19.6 19.6 42.5 30.7 41.8 Maximum 56.0 39.7 24.4 16.1 22.3 38.8 10.0 23.4 22.9 55.2 37.9 46.1 Minimum 40.0 30.0 17.9 9.8 13.0 24.6 5.5 17.5 15.6 36.7 25.7 37.9 Range 16.0 9.7 6.5 6.3 9.3 14.2 4.5 5.9 7.3 18.5 12.2 8.2 SD 4.0 2.2 1.9 1.4 2.4 4.0 1.0 1.7 1.8 4.3 3.2 2.3 CV 8.5 6.1 9.2 11.6 14.1 12.5 13.4 8.4 9.0 9.9 10.2 5.5

Pilio. (n=28) Mean 41.3 32.3 16.4 10.6 15.7 28.8 6.4 18.7 17.3 35.2 26.8 38.3 Median 42.0 32.0 16.2 10.6 15.7 29.1 6.3 18.7 17.0 36.0 26.9 38.5 Maximum 48.0 36.7 20.4 12.1 20.6 35.7 9.0 21.7 20.0 40.0 31.8 45.0 Minimum 35.0 29.0 13.5 8.9 10.0 22.7 4.0 15.9 14.3 30.5 23.0 30.0 Range 13.0 7.7 6.9 3.3 10.5 13.0 5.0 5.8 5.7 9.5 8.8 15.0 SD 3.2 1.9 1.6 0.9 2.2 4.1 1.2 1.9 1.6 3.1 2.2 3.6 CV 7.7 5.9 9.7 8.5 14.2 14.2 18.2 10.0 9.4 8.8 8.4 9.4

Proc. (n=5) Mean 33.0 28.4 12.5 9.0 13.6 25.7 5.7 16.1 15.5 27.0 20.1 35.3 Median 33.0 28.4 13.0 9.2 13.9 27.2 6.0 16.6 16.0 27.9 20.3 35.4 Maximum 34.0 29.1 13.4 9.4 15.4 27.9 7.0 16.6 16.6 28.2 22.2 38.4 Minimum 32.0 28.1 11.5 8.4 11.4 22.3 4.5 15.2 14.3 24.5 18.7 31.8 Range 2.0 1.0 1.9 0.9 4.0 5.6 2.5 1.4 2.2 3.7 3.4 6.6 SD 0.7 0.4 0.9 0.4 1.7 2.6 1.0 0.6 1.0 1.6 1.3 2.4 CV 2.1 1.4 7.2 3.9 12.6 10.3 17.1 4.0 6.7 5.9 6.7 6.7

197

maxalvlen biecm iv-pms i1i2 bicanin bicanex palhei biseptal bienm ol-sta ol-pms bien Semno. (n=9) Mean 47.7 40.9 19.4 11.8 17.7 32.5 7.7 21.3 21.6 41.9 29.5 50.0 Median 47.0 40.6 18.8 12.3 18.5 33.7 7.0 22.3 21.4 41.1 30.4 51.1 Maximum 55.0 46.2 24.0 13.4 20.7 38.4 10.0 23.4 24.6 49.7 36.7 56.1 Minimum 39.0 35.9 16.8 9.7 13.9 24.0 6.0 17.3 19.1 34.2 24.7 43.4 Range 16.0 10.3 7.2 3.8 6.8 14.4 4.0 6.0 5.5 15.4 12.0 12.7 SD 5.8 4.5 2.3 1.3 2.4 4.7 1.5 2.4 1.9 5.4 3.9 4.4 CV 12.1 10.9 12.0 11.0 13.4 14.5 19.6 11.2 8.8 13.0 13.1 8.8

Trachy. (n=33) Mean 37.3 34.0 15.6 9.7 14.2 26.2 6.5 17.2 18.2 30.9 23.2 40.1 Median 37.5 33.7 15.6 9.8 14.0 26.2 6.5 17.3 18.1 31.2 23.0 40.0 Maximum 41.0 39.4 18.6 11.3 17.4 32.5 9.0 20.2 21.9 35.1 28.6 45.8 Minimum 32.0 30.9 12.9 8.0 12.0 21.2 4.0 13.6 15.5 26.2 19.1 33.0 Range 9.0 8.5 5.7 3.3 5.4 11.3 5.0 6.6 6.4 8.8 9.5 12.8 SD 2.4 2.3 1.6 0.9 1.6 3.1 1.1 1.9 1.6 2.4 2.1 3.3 CV 6.6 6.7 9.9 9.5 11.4 11.8 17.4 10.9 8.9 7.9 8.9 8.3

Pres. (n=18) Mean 32.7 30.3 12.8 9.3 13.8 24.3 5.4 16.2 17.0 25.9 19.0 37.7 Median 32.1 30.2 12.6 9.3 13.6 24.2 5.3 16.2 16.9 25.5 18.8 37.7 Maximum 38.0 34.1 16.9 10.8 16.8 27.0 7.0 18.9 19.4 30.9 21.1 43.0 Minimum 29.6 26.6 10.6 7.4 11.2 21.0 4.0 14.0 14.5 22.7 17.4 32.5 Range 8.4 7.4 6.3 3.4 5.6 5.9 3.0 4.9 5.0 8.2 3.7 10.5 SD 2.3 2.0 1.5 0.8 1.3 1.7 0.7 1.3 1.1 2.3 1.2 2.6 CV 7.1 6.6 11.5 8.5 9.7 7.0 12.7 7.8 6.8 8.8 6.1 6.8

198

maxalvlen biecm iv-pms i1i2 bicanin bicanex palhei biseptal bienm ol-sta ol-pms bien Pyga. (n=25) Mean 40.6 36.5 16.1 10.9 15.2 28.0 6.1 19.0 20.6 32.9 24.9 41.1 Median 41.0 36.0 15.9 10.8 15.0 29.0 6.0 19.0 20.5 32.8 24.4 41.9 Maximum 46.0 40.4 19.7 13.1 17.8 32.1 7.6 22.5 23.6 39.4 30.5 45.0 Minimum 32.0 31.3 14.0 8.8 11.5 22.5 3.0 15.6 17.2 23.9 19.1 35.7 Range 14.0 9.2 5.7 4.3 6.3 9.7 4.6 6.8 6.4 15.5 11.3 9.3 SD 3.7 2.8 1.5 1.0 1.2 3.4 1.1 2.0 1.8 3.5 2.6 2.5 CV 9.1 7.8 9.1 9.5 7.9 12.1 18.5 10.3 8.9 10.6 10.5 6.0

Rhino. (n=17) Mean 43.5 37.7 17.0 11.3 16.2 29.3 5.2 20.2 18.1 36.7 24.9 46.7 Median 43.0 38.1 17.0 11.2 16.4 28.3 5.0 20.3 17.7 36.9 25.0 46.0 Maximum 50.0 41.5 24.3 12.6 18.9 37.6 7.2 22.7 22.0 43.2 29.3 51.6 Minimum 38.7 32.8 11.8 9.1 12.4 24.1 3.8 17.0 15.4 28.3 19.5 42.9 Range 11.3 8.7 12.6 3.5 6.6 13.5 3.4 5.7 6.6 14.9 9.8 8.7 SD 2.9 2.5 2.8 0.8 1.8 3.7 0.9 1.4 1.9 3.3 2.3 2.4 CV 6.8 6.7 16.8 7.0 10.9 12.6 16.7 6.8 10.3 9.0 9.4 5.2

199

maxalvlen biecm iv-pms i1i2 bicanin bicanex palhei biseptal bienm ol-sta ol-pms bien Nasalis (n=28) Mean 44.4 36.4 19.7 11.3 16.8 30.8 6.5 20.6 20.1 36.7 29.0 44.5 Median 45.9 37.0 19.4 11.3 17.0 31.5 7.0 20.5 20.4 38.2 30.2 43.9 Maximum 50.0 41.5 23.0 12.9 20.2 37.2 10.0 24.0 23.6 42.6 33.7 51.5 Minimum 38.0 31.6 15.3 9.6 13.6 23.7 4.0 17.6 17.5 29.5 23.0 35.0 Range 12.0 9.9 7.7 3.3 6.6 13.5 6.0 6.4 6.1 13.1 10.7 16.5 SD 3.8 2.3 2.0 1.0 1.8 4.0 1.6 2.0 1.7 4.4 3.6 3.8 CV 8.6 6.2 10.1 8.9 10.6 13.0 24.2 9.5 8.2 11.9 12.3 8.5

Simias (n=7) Mean 37.0 33.5 15.6 9.2 14.3 23.9 6.6 17.2 17.2 29.5 22.9 33.1 Median 36.0 32.5 15.5 9.1 13.1 23.3 7.0 17.6 17.3 28.5 22.8 33.0 Maximum 42.0 39.3 18.2 10.5 18.7 29.0 9.0 18.7 18.5 33.7 25.9 36.5 Minimum 34.0 30.3 13.2 8.5 12.3 19.9 4.0 15.5 16.0 26.3 20.2 29.0 Range 8.0 9.0 4.9 2.0 6.5 9.1 5.0 3.2 2.5 7.4 5.7 7.5 SD 2.6 2.9 1.5 0.6 2.3 3.5 1.5 1.3 1.0 3.0 2.2 2.7 CV 7.2 8.7 9.5 6.7 16.2 14.7 23.3 7.5 5.6 10.1 9.6 8.2

200

Box Plots for cranial variables of the Colobinae (Colobina: Pro. – Procolobus, n=5; Pi. – Piliocolobus, n=28; Co. – Colobus, n=26; Presbytina: Pre. – Presbytis, n=18; Tr. – Trachypithecus, n=33; Se. – Semnopithecus, n=9; & Nasalina: Si. – Simias, n=7; Py. - Pygathrix, n=25; R. – Rhinopithecus, n=17; & N. – Nasalis, n=28; variables in alphabetical order by abbreviation):

80 Figure 5.2 Co. N. R. Se. Py. 70 Pi. Tr. mm Pre. Si. 60 Pro.

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Inferior Cranial Length (bas-nas)

Figure 5.3 Co. 100 N.

Se. 90 Pi. Py. R.

80 Tr.

mm Si. Pre. 70 Pro.

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Superior Facial Length (bas-pros)

201

Figure 5.4 70 R.

60 Py. N. Co. Pi. Pre. Tr. Se. 50 mm Si. Pro. 40

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Biasterionic Breadth (biast)

50 Figure 5.5 49 48 47 Se. 46 45 44 43 R. N. 42 Py. 41 40 Co. Tr. 39 38 Pi. mm 37 36 35 Pre. Si. 34 33 32 31 30 Pro. 29 28 27 26

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Maxillo-alveolar Breadth (biecm)

202

30 Figure 5.6 29 28 27 26 Se. 25 Py. 24 Co. N. 23 Tr. R. 22 21 Pi. 20 Pre. mm 19 Si. 18 17 Pro. 16 15 14 13 12 11

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Palatal Breadth (bienm)

R. 80 Figure 5.7 Py. Se.

Pi. Tr. 70 Co. N.

Pre. Pro. Si. 60 mm

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Bifrontomalartemporale Breadth (bifmt)

203

Figure 5.8 Se. 70

R. Co. N. Tr. 60 Pi. Py. mm Pre. Si. 50 Pro.

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Bizygomaxillare Inferior Breadth (bizi)

60 Figure 5.9

Py. 50 Co. Tr. N. Pre. Se. 40 R. mm Pi. Pro.

30 Si.

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Bizygomaxillare Superior Breadth (bizs)

204

Figure 5.10 Se. N. 100 R. Pi. 90 Co. Tr. Py.

mm 80 Pro. Pre. Si.

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Bizygomatic Breadth (bizygo)

Figure 5.11 R. 100 Se. N.

Pi. Co. Py. 90 Tr.

mm Pre. 80 Si Pro. .

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Cranial Vault Length (g-o)

205

60 Figure 5.12 N. Se.

50 Co. Py. Pi. Si. R.

40 Tr. mm Pro.

Pre. 30

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Superior Facial Height (nas-pros)

N. 30 Figure 5.13

Se. Si. 20 Pi. Co. Py. R. Tr.

mm Pro. Pre.

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Sagittal Length of the Nasal Bones (nas-rhi)

206

60 Figure 5.14

Co.

Se. 50

R. N. Pi. Py. 40 mm

Tr. Si. Pre. 30 Pro.

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Palatal Length A (ol-sta)

50 Figure 5.15 Se.

R. N. Pi. Co.

40 Pro. Tr. Pre. Si. Py. mm

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Maximum Length of the Zygomatic (zs-zgyi)

207

30 Figure 5.16 29 28 27 Se. 26 N. 25 24 23 22 Si. R. Pre. Tr. 21 Pi. 20 mm 19 18 17 Co. Py. 16 15 Pro. 14 13 12 11

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: Zygomatico-maxillary Suture Length (zs-zi)

208

Table 5.12: Shapiro-Wilk results for cranial variables of the Colobinae; those in bold represent significant departures from normality (variables listed in alphabetical order by abbreviation).

Pro. Pi. Co. Pre. Tr. Se. n=5 n=28 n=26 n=18 n=33 n=9 1. Bas-nas W: 0.93 0.96 0.97 0.94 0.98 0.85 p(normal): 0.56 0.28 0.7 0.25 0.67 0.08 2. Bas-pros W: 0.96 0.99 0.94 0.94 0.97 0.88 p(normal): 0.79 0.99 0.1 0.32 0.62 0.16 3. Biast W: 0.97 0.99 0.97 0.97 0.94 0.88 p(normal): 0.88 0.95 0.61 0.76 0.09 0.14 4. Biecm W: 0.85 0.98 0.93 0.97 0.93 0.83 p(normal): 0.19 0.74 0.09 0.79 0.03 0.05 5. Bienm W: 0.83 0.95 0.95 0.98 0.97 0.95 p(normal): 0.14 0.16 0.29 0.93 0.53 0.64 6. Bifmt W: 0.94 0.97 0.94 0.93 0.97 0.98 p(normal): 0.63 0.58 0.15 0.21 0.54 0.97 7. Bizi W: 0.82 0.98 0.98 0.95 0.94 0.87 p(normal): 0.11 0.79 0.88 0.48 0.06 0.13 8. Bizs W: 0.96 0.93 0.96 0.93 0.93 0.96 p(normal): 0.83 0.06 0.41 0.2 0.03 0.84 9. Bizygo W: 0.89 0.93 0.95 0.94 0.97 0.89 p(normal): 0.37 0.08 0.23 0.27 0.43 0.2 10. G-o W: 0.86 0.96 0.96 0.93 0.96 0.84 p(normal): 0.23 0.35 0.48 0.19 0.24 0.06 11. Nas-pros W: 0.97 0.98 0.95 0.91 0.98 0.95 p(normal): 0.87 0.86 0.29 0.08 0.73 0.66 12. Nas-rhi W: 0.93 0.97 0.98 0.95 0.97 0.91 p(normal): 0.58 0.69 0.91 0.51 0.47 0.29 13. Ol-sta W: 0.84 0.92 0.95 0.91 0.97 0.93 p(normal): 0.17 0.04 0.25 0.094 0.64 0.48 14. Zs-zgyi W: 0.87 0.95 0.97 0.98 0.99 0.89 p(normal): 0.25 0.2 0.32 0.93 0.96 0.19 15. Zs-zi W: 0.98 0.95 0.96 0.95 0.96 0.88 p(normal): 0.93 0.2 0.37 0.36 0.3 0.17

209

Si. Py. R. N. n=7 n=25 n=17 n=28 1. Bas-nas W: 0.77 0.96 0.99 0.91 p(normal): 0.02 0.4 0.99 0.02 2. Bas-pros W: 0.85 0.94 0.98 0.87 p(normal): 0.12 0.12 0.98 0.002 3. Biast W: 0.76 0.96 0.96 0.96 p(normal): 0.02 0.51 0.57 0.43 4. Biecm W: 0.88 0.93 0.96 0.98 p(normal): 0.21 0.07 0.6 0.83 5. Bienm W: 0.94 0.97 0.96 0.94 p(normal): 0.62 0.57 0.69 0.15 6. Bifmt W: 0.97 0.96 0.97 0.94 p(normal): 0.88 0.39 0.82 0.09 7. Bizi W: 0.9 0.96 0.88 0.97 p(normal): 0.33 0.51 0.03 0.49 8. Bizs W: 0.89 0.96 0.93 0.89 p(normal): 0.3 0.42 0.25 0.89 9. Bizygo W: 0.97 0.9 0.98 0.92 p(normal): 0.89 0.02 0.96 0.04 10. G-o W: 0.94 0.92 0.91 0.91 p(normal): 0.66 0.04 0.02 0.11 11. Nas-pros W: 0.94 0.97 0.91 0.92 p(normal): 0.65 0.72 0.09 0.03 12. Nas-rhi W: 0.94 0.97 0.97 0.96 p(normal): 0.67 0.73 0.92 0.34 13. Ol-sta W: 0.9 0.96 0.96 0.88 p(normal): 0.32 0.49 0.56 0.004 14. Zs-zgyi W: 0.95 0.97 0.97 0.93 p(normal): 0.75 0.61 0.77 0.07 15. Zs-zi W: 0.97 0.96 0.96 0.94 p(normal): 0.92 0.49 0.64 0.09

210

normally distributed, although mostly attributable to Nasalis, which is one of the largest and most sexually dimorphic of the extant colobines. All cranial variables examined for Procolobus, Colobus and Presbytis were normally distributed. Palatal length A (ol-sta) was the only variable not normally distributed for Piliocolobus, while maxillo-alveolar breadth (biecm) was the only variable not normally distributed for Semnopithecus. Trachypithecus, Simias, Pygathrix and Rhinopithecus each had two variables which were not normally distributed. These include, maxillo-alveolar (biecm) and bizygomaxillare superior (bizs) breadths for Trachypithecus; inferior cranial length (bas-nas) and biasterionic breadth (biast) for Simias; bizygomatic breadth (bizygo) and cranial vault length (g-o) for Pygathrix; and lastly, bizygomaxillare inferior breadth (bizi) and cranial vault length (g-o) for Rhinopithecus. There were five variables for Nasalis that were not normally distributed. These include inferior cranial length (bas-nas), superior facial length (bas-pros), bizygomatic breadth (bizygo), superior facial length (nas-pros) and palatal length A (ol- sta).

5.2.2 Kruskal-Wallis and Mann-Whitney results for cranial variables: By applying Kruskal-Wallis and Mann-Whitney tests to selected variables many significant differences between sample medians were evident. In fact, there were more size- related differences than similarities. Table 5.13 provides the results of Kruskal-Wallis and Mann-Whitney for the Colobinae. Because of its small size, cranial measurements for Procolobus were nearly always significantly different; although not when compared to other similar sized colobines. For example, the bizygomatic breadth (bizygo) of Procolobus is not significantly from Presbytis, Trachypithecus or Simias (Table 5.13, no. 9). Furthermore, because of their larger size, cranial dimensions for Nasalis and Rhinopithecus were also in general significantly different from smaller genera. However, the latter shares some dimensions with Pygathrix and is thus not significantly different [e.g. inferior cranial length (bas-nas) and maxillo-alveolar and palatal breadths (biecm and bienm)]. Despite the unusually narrow superior facial breadth (bifmt) of Nasalis (Groves, 1970), Piliocolobus and Colobus overlap and are similar in size and thus not significantly different (Table 5.13, no. 6), although the length of the nasal bones (nas-rhi, no. 12) and superior facial height (no. 11) exceeds that of all other colobines with statistical significance, even Simias, the

211

Table 5.13: Kruskal-Wallis results for cranial variables of the Colobinae with Mann-Whitney pairwise comparisons (p (same)) (variables listed in alphabetical order by abbreviation).

Pro. Pi. Co. Pre. Tr. Se. Si. Py. R. N. n=5 n=28 n=26 n=18 n=33 n=9 n=7 n=25 n=17 n=28 1. Bas-nas 0 0.002 0.0005 0.16 0.025 0.004 0.87 0.002 0.001 0.0006 Pro. H: 100.4 0 1.25E-06 0.00005 0.003 0.008 0.002 0.24 0.15 0.0009 Pi. Hc: 100.6 0 3.51E-08 1.56E-09 0.79 0.0001 0.0004 0.02 0.80 Co. p(same): 1.34E-17 0 0.08 0.0003 0.38 0.00005 5.67E-06 3.64E-07 Pre. 0 0.0004 0.06 0.0005 0.00002 3.50E-07 Tr. 0 0.004 0.03 0.16 0.61 Se. 0 0.002 0.001 0.0004 Si. 0 0.24 0.005 Py. 0 0.14 R. 0 N. 2. Bas-pros 0 0.0005 0.0005 0.50 0.001 0.003 0.03 0.0006 0.001 0.0005 Pro. H: 122.9 0 8.83E-08 7.03E-08 1.00E-04 0.20 0.006 0.44 0.99 0.0002 Pi. Hc: 123 0 2.49E-08 9.11E-11 0.31 0.00007 0.00002 0.00002 0.97 Co. p(same): 3.43E-22 0 2.24E-06 0.00005 0.03 7.49E-08 2.56E-06 1.72E-08 Pre. 0 0.0006 0.22 0.00008 0.002 8.40E-09 Tr. 0 0.004 0.07 0.06 0.22 Se. 0 0.003 0.02 0.0005 Si. 0 0.5 0.0005 Py. 0 0.0007 R. 0 N. 3. Biast 0 0.009 0.003 0.02 0.40 0.01 0.07 0.0007 0.001 0.0006 Pro. H: 102.2 0 0.02 0.64 0.01 0.05 0.29 4.00E-07 1.78E-07 5.65E-06 Pi. Hc: 102.3 0 0.02 0.0001 0.66 0.03 0.002 8.24E-06 0.02 Co. p(same): 5.56E-18 0 0.06 0.03 0.35 3.27E-06 1.33E-06 0.00004 Pre. 0 0.007 0.36 3.90E-08 4.05E-08 7.29E-08 Tr. 0 0.03 0.005 0.0005 0.02 Se. 0 0.0002 0.0001 0.0004 Si. 0 0.012 0.35 Py. 0 0.002 R. 0 N. 212

Pro. Pi. Co. Pre. Tr. Se. Si. Py. R. N. n=5 n=28 n=26 n=18 n=33 n=9 n=7 n=25 n=17 n=28 4. Biecm 0 0.0006 0.0005 0.03 0.0004 0.003 0.006 0.0006 0.001 0.0005 Pro. H: 108.5 0 1.23E-07 0.002 0.005 0.00001 0.38 1.26E-06 6.15E-07 8.39E-08 Pi. Hc: 108.5 0 2.08E-07 0.0001 0.03 0.02 0.99 0.11 0.70 Co. p(same): 2.96E-19 0 2.99E-06 0.00003 0.008 1.92E-07 1.13E-06 6.20E-08 Pre. 0 0.00006 0.44 0.001 0.00004 0.0002 Tr. 0 0.004 0.009 0.13 0.022 Se. 0 0.02 0.006 0.01 Si. 0 0.20 0.92 Py. 0 0.08 R. 0 N. 5. Bienm 0 0.03 0.002 0.02 0.002 0.003 0.04 0.0006 0.008 0.0005 Pro. H: 89.36 0 0.00004 0.55 0.04 0.00005 0.98 5.54E-07 0.18 2.89E-06 Pi. Hc: 89.36 0 4.33E-06 0.001 0.02 0.0009 0.13 0.005 0.75 Co. p(same): 2.19E-15 0 0.007 0.00004 0.61 4.21E-07 0.07 6.18E-07 Pre. 0 0.0002 0.12 0.00002 0.85 0.0001 Tr. 0 0.001 0.24 0.001 0.03 Se. 0 0.0003 0.27 0.0004 Si. 0 0.0005 0.25 Py. 0 0.0002 R. 0 N. 6. Bifmt 0 0.009 0.03 0.07 0.17 0.005 0.57 0.0007 0.001 0.009 Pro. H: 102.7 0 0.67 0.02 0.0005 0.05 0.006 1.00E-06 5.45E-07 0.76 Pi. Hc: 102.7 0 0.11 0.02 0.03 0.02 1.70E-06 9.30E-07 0.53 Co. p(same): 4.44E-18 0 0.43 0.0005 0.11 1.68E-07 9.52E-07 0.01 Pre. 0 0.0002 0.35 1.11E-09 2.69E-08 0.0002 Tr. 0 0.002 0.09 0.01 0.06 Se. 0 0.00009 0.0002 0.005 Si. 0 0.04 1.00E-06 Py. 0 4.27E-07 R. 0 N. 213

Pro. Pi. Co. Pre. Tr. Se. Si. Py. R. N. n=5 n=28 n=26 n=18 n=33 n=9 n=7 n=25 n=17 n=28 7. Bizi 0 0.01 0.0009 0.02 0.002 0.003 0.05 0.0007 0.001 0.0006 Pro. H: 88.22 0 0.001 0.13 0.93 0.00005 0.17 0.01 2.68E-06 0.0002 Pi. Hc: 88.23 0 0.00009 0.009 0.001 0.0009 0.19 0.001 0.61 Co. p(same): 3.70E-15 0 0.18 0.00005 0.65 0.0002 6.63E-06 8.78E-06 Pre. 0 0.00005 0.13 0.0008 2.24E-06 0.0002 Tr. 0 0.001 0.0002 0.16 0.002 Se. 0 0.0009 0.0005 0.0003 Si. 0 0.00002 0.10 Py. 0 0.005 R. 0 N. 8. Bizs 0 0.63 0.0008 0.43 0.39 0.11 0.006 0.0007 0.001 0.33 Pro. H: 120.7 0 7.38E-09 0.25 0.35 0.03 0.00006 1.31E-09 1.48E-07 0.11 Pi. Hc: 120.7 0 0.00006 0.00006 0.001 0.00007 0.005 0.72 1.30E-09 Co. p(same): 9.70E-22 0 0.93 0.50 0.0002 1.03E-06 0.00007 0.03 Pre. 0 0.23 0.00004 1.71E-07 0.00006 0.01 Tr. 0 0.001 0.00005 0.0007 0.004 Se. 0 0.00007 0.0002 0.00006 Si. 0 0.02 6.67E-10 Py. 0 5.24E-08 R. 0 N. 9. Bizygo 0 0.02 0.006 0.63 0.18 0.003 0.42 0.03 0.004 0.004 Pro. H: 89.68 0 0.70 3.95E-06 0.005 0.0007 0.0009 0.41 0.01 0.02 Pi. Hc: 89.68 0 6.87E-07 0.0004 0.0006 0.0002 0.29 0.01 0.04 Co. p(same): 1.89E-15 0 0.0004 0.00004 0.67 0.00004 2.18E-06 2.40E-07 Pre. 0 0.00002 0.006 0.02 0.00001 0.00001 Tr. 0 0.001 0.0003 0.05 0.008 Se. 0 0.001 0.0005 0.0001 Si. 0 0.001 0.002 Py. 0 0.82 R. 0 N. 214

Pro. Pi. Co. Pre. Tr. Se. Si. Py. R. N. n=5 n=28 n=26 n=18 n=33 n=9 n=7 n=25 n=17 n=28 10. G-o 0 0.002 0.009 0.25 0.08 0.004 0.57 0.001 0.0004 0.001 Pro. H: 116.1 0 0.04 3.86E-06 0.0001 0.02 0.0002 0.11 0.00009 9.30E-07 Pi. Hc: 116.2 0 0.0007 0.03 0.002 0.005 0.001 0.60 4.01E-07 Co. p(same): 8.43E-21 0 0.09 0.0001 0.97 6.60E-07 2.38E-08 4.82E-07 Pre. 0 0.0001 0.15 6.13E-06 5.95E-09 3.03E-08 Tr. 0 0.002 0.05 0.21 0.79 Se. 0 0.0003 0.00006 0.0002 Si. 0 0.003 7.77E-06 Py. 0 0.23 R. 0 N. 11. Nas-pros 0 0.002 0.0006 0.002 0.93 0.008 0.05 0.006 0.18 0.0004 Pro. H: 130.8 0 0.007 1.51E-08 6.77E-09 0.06 0.36 0.32 0.004 3.53E-06 Pi. Hc: 130.8 0 2.49E-08 2.36E-10 0.38 0.08 0.01 0.0002 0.0002 Co. p(same): 7.96E-24 0 3.87E-07 0.00003 0.0002 8.58E-08 3.53E-06 1.51E-08 Pre. 0 0.00001 0.005 1.09E-06 0.05 3.07E-11 Tr. 0 0.11 0.04 0.002 0.07 Se. 0 0.78 0.25 0.001 Si. 0 0.06 1.20E-06 Py. 0 1.06E-06 R. 0 N. 12. Nas-rhi 0 0.04 0.03 0.03 0.20 0.11 0.006 0.22 0.23 0.0005 Pro. H: 125.8 0 0.44 3.24E-07 9.28E-07 0.94 0.02 0.18 0.001 7.99E-10 Pi. Hc: 125.9 0 2.69E-07 1.03E-06 0.58 0.001 0.39 0.001 3.48E-10 Co. p(same): 8.45E-23 0 0.06 0.00004 0.0002 1.49E-06 0.51 1.51E-08 Pre. 0 0.0002 0.00005 0.00007 0.83 2.40E-11 Tr. 0 0.11 0.32 0.01 0.00004 Se. 0 0.002 0.001 0.002 Si. 0 0.01 5.96E-10 Py. 0 7.67E-07 R. 0 N. 215

Pro. Pi. Co. Pre. Tr. Se. Si. Py. R. N. n=5 n=28 n=26 n=18 n=33 n=9 n=7 n=25 n=17 n=28 13. Ol-sta 0 0.0004 0.0005 0.28 0.003 0.003 0.12 0.001 0.001 0.0005 Pro. H: 134.3 0 4.17E-09 2.54E-08 4.18E-06 0.003 0.001 0.32 0.19 0.08 Pi. Hc: 134.3 0 2.49E-08 6.06E-11 0.33 0.00007 2.47E-09 3.61E-06 1.01E-06 Co. p(same): 1.52E-24 0 4.75E-07 0.00003 0.006 1.29E-07 8.05E-07 4.81E-08 Pre. 0 0.00001 0.23 0.006 9.85E-07 9.17E-06 Tr. 0 0.001 0.0003 0.03 0.03 Se. 0 0.03 0.0005 0.002 Si. 0 0.002 0.006 Py. 0 0.6 R. 0 N. 14. Zs-zgyi 0 0.01 0.01 0.28 0.004 0.14 0.87 0.08 0.006 0.008 Pro. H: 111 0 0.77 0.00002 4.65E-10 0.33 0.01 1.54E-07 0.79 0.13 Pi. Hc: 111 0 0.00003 7.99E-10 0.25 0.02 2.56E-07 0.86 0.07 Co. p(same): 9.33E-20 0 0.04 0.004 0.29 0.93 0.00007 5.17E-06 Pre. 0 0.0003 0.004 0.003 3.21E-08 1.51E-10 Tr. 0 0.07 0.003 0.36 0.58 Se. 0 0.18 0.009 0.005 Si. 0 7.59E-07 4.28E-08 Py. 0 0.24 R. 0 N. 15. Zs-zi 0 0.01 0.11 0.39 0.17 0.03 0.006 0.02 0.002 0.0005 Pro. H: 98.06 0 0.02 0.21 0.004 0.06 0.005 0.04 0.004 3.04E-07 Pi. Hc: 98.06 0 0.72 0.65 0.009 0.00007 0.72 8.19E-07 3.48E-10 Co. p(same): 3.89E-17 0 0.59 0.03 0.007 0.9 0.003 2.16E-06 Pre. 0 0.005 0.00008 0.23 9.85E-07 7.03E-11 Tr. 0 0.75 0.02 0.36 8.30E-01 Se. 0 0.00009 0.25 0.12 Si. 0 1.88E-06 5.96E-10 Py. 0 0.001 R. 0 N. 216

Proboscis monkey’s closest living relative (Groves, 1970; Delson, 1975; Whittaker et al, 2006). Bizygomaxillare superior breadth (bizs) revealed interesting similarities and dissimilarities between colobine genera. Regardless of its small size, this measurement for Procolobus was not significantly different from Piliocolobus, Presbytis, Trachypithecus, Semnopithecus and Nasalis.

5.2.3 One-way ANOVA results for cranial variables: Procolobus (n=5), Semnopithecus (n=9) and Simias (n=7) were excluded from all one-way ANOVA analyses due to their small sample sizes (as well as for the cranial indices examined, see below). Table 5.14 lists the results of applying one-way ANOVA to the remaining Colobinae genera, excluding those identified as not having a normal distribution via Shapiro-Wilk. In contrast to the other primate groupings examined thus far, very few variables produced the majority of variation between groups. Only two variables generated greater than 70% of variation between groups, sagittal length of the nasal bones (nas-rhi) and palatal length A (ol-sta). The remaining variables had percentages of variation between groups ranging from ~25% (bizi) to ~66% (bizs). Similar to results for Kruskal-Wallis and Mann-Whitney, there are far more significant differences than non-significant ones. As such, it’s more meaningful perhaps to mention variables that were not significantly different. Some of these included, the superior facial length (bas-pros) between Piliocolobus, Pygathrix and Rhinopithecus which were very similar (Table 5.14, no. 2). Nasalis is known for its narrow superior facial breadth (bifmt) but this dimension is not significantly different between it and Piliocolobus and Colobus but is significantly different from the other snub-nosed colobines (Table 5.14, no. 6). The sagittal length of the nasal bones (nas-rhi) revealed interesting similarities (i.e. not significantly different) in size between Piliocolobus, Colobus and Pygathrix on the one hand and Rhinopithecus, Trachypithecus and Presbytis on the other (Table 5.14, no. 12), whereas Nasalis was significantly different from all other colobine genera. Aside from Pygathrix and Trachypithecus, all palatal length A dimensions were significantly different between colobine genera (Table 5.14, no. 13). Lastly, the maximum length of the zygomatic (zs-zgyi) and zygomatico-maxillary suture length (zs-zi) exposed interesting similarities and (Table 5.14, no. 14 & 15). With regard to the former (zs-zgyi),

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Table 5.14: One-Way ANOVA results for cranial variables of the Colobinae (variables listed in alphabetical order by abbreviation). Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): p(same): Pi. Co. Pre. Tr. Py. R. 1. Bas-nas 0 .00002 .0004 0.12 0.79 0.04 Between F: 26.87 0 .00002 .00002 .0001 0.05 groups: 1847.33 48.68 5 369.466 26.75 5.94E-19 0.25 df: 60.31 0 0.55 .00002 .00002 Within groups: 1947.27 51.32 141 13.8104 p: 3.51E-14 0 0.002 .00002 Total: 3794.6 146 0 0.56 ex. Pro., N., Se., & Si. 0 2. Bas-pros Pi. Co. Pre. Tr. Py. R. Between F: 58.07 0 .00002 .00002 0.009 0.94 1 groups: 7556.01 62.71 5 1511.2 47.42 1.45E-28 0.02 df: 60.52 0 .00002 .00002 .00002 .00002 Within groups: 4493.68 37.29 141 31.87 p: 8.14E-22 0 0.0003 .00002 .00002 Total: 12049.7 146 0 0.0002 0.008 0 0.95 ex. Pro., N., Se., & Si. 0 3. Biast Pi. Co. Pre. Tr. Py. R. N. Between F: 25.76 0 0.41 0.99 0.27 .00003 .00003 .0001 groups: 2689.14 51.40 6 448.19 29.62 4.67E-24 0.78 df: 69.15 0 0.15 0.0005 0.006 .00003 0.14 Within groups: 2542.24 48.60 168 15.13 p: 7.44E-16 0 0.59 .00003 .00003 .00003 Total: 5231.38 174 0 .00003 .00003 .00003 0 0.02 0.94 0 .0004 ex. Pro. Se. & Si. 0

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Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. F comparisons sqrs: Var.: df: square: F: p(same): of Var. test: p(same): 4. Biecm p(same): Pi. Co. Pre. Py. R. N. Between F: 38.29 0 .00002 0.04 .00002 .00002 .00002 Pi. groups: 909.144 55.68 5 181.829 34.17 1.71E-22 0.17 df: 59.12 0 .00002 1 0.52 1 Co. Within groups: 723.692 44.32 136 5.32126 p: 2.52E-17 0 .00002 .00002 .00002 Pre. Total: 1632.84 141 0 0.46 1 Py. 0 0.44 R. ex. Pro. Se., Tr. & Si. 0 N. 5. Bienm Pi. Co. Pre. Tr. Py. R. N. Between F: 18.9 0 .00003 0.99 0.51 .00003 0.62 .00003 Pi. groups: 289.796 38.27 6 48.2993 17.36 1.41E-15 0.29 df: 69.93 0 .00003 0.02 0.60 0.01 0.99 Co. Within groups: 467.498 61.73 168 2.78273 p: 6.08E-13 0 0.13 .00003 0.19 .00003 Pre. Total: 757.294 174 0 .00004 1 0.003 Tr. 0 0.00003 0.93 Py. 0 0.001 R. ex. Pro. Se. & Si. 0 N. 6. Bifmt Pi. Co. Pre. Tr. Py. R. N. Between F: 36.56 0 0.99 0.11 0.02 .00003 .00003 1 Pi. groups: 4348.46 54.07 6 724.743 32.96 4.49E-26 0.18 df: 69.84 0 0.31 0.08 .00003 .00003 0.99 Co. Within groups: 3693.6 45.93 168 21.9857 p: 1.10E-19 0 0.99 .00003 .00003 0.07 Pre. Total: 8042.06 174 0 .00003 .00003 0.01 Tr. 0 0.35 .00003 Py. 0 .00003 R. ex. Pro. Se. & Si. 0 N.

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Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 7. Bizi p(same): Pi. Co. Pre. Tr. Py. N. Between F: 10.76 0 0.002 0.83 0.99 0.09 0.0001 Pi. groups: 784.44 24.90 5 159.89 10.08 2.38E-08 0.20 df: 68.49 0 .00003 0.001 0.87 0.98 Co. Within groups: 2365.94 75.10 152 15.57 p: 1.17E-07 0 0.59 0.002 .00002 Pre. Total: 3150.38 157 0 0.22 .0006 Tr. 0 0.43 Py. ex. Pro. Se., R. & Si. 0 N. 8. Bizs Pi. Co. Pre. Py. R. N. Between F: 53.46 0 .00002 0.62 .00002 .00002 0.77 Pi. groups: 3513.59 65.69 5 702.72 52.08 5.95E-30 0.25 df: 59.53 0 .00002 0.04 1 .00002 Co. Within groups: 1834.91 34.31 136 13.49 p: 9.41E-21 0 .00002 .00002 0.04 Pre. Total: 5348.5 141 0 0.05 .00002 Py. 0 .00002 R. ex. Pro. Se., Si. & Tr. 0 N. 9. Bizygo Pi. Co. Pre. Tr. R. Between F: 27.06 0 0.99 0.0001 0.04 0.006 Pi. groups: 2670.1 42.02 4 667.53 21.2 3.62E-13 0.02 df: 52.64 0 0.0001 0.01 0.02 Co. Within groups: 3683.78 57.98 117 31.49 p: 3.18E-12 0 0.03 0.0001 Pre. Total: 6353.88 121 0 0.0001 Tr. ex. Pro. Se., Si., Py. & N. 0 R.

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Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 10. G-o p(same): Pi. Co. Pre. Tr. N. Between F: 34.35 0 0.26 0.0006 0.23 .00002 Pi. groups: 2628.08 50.70 4 657.02 32.9 7.39E-19 0.61 df: 61.48 0 .00002 0.0004 .00006 Co. Within groups: 2555.94 49.30 128 19.97 p: 4.71E-15 0 0.30 .00002 Pre. Total: 5184.02 132 ex. 0 .00002 Tr. Pro., Py., Se., Si. & R. 0 N. 11. Nas-pros Pi. Co. Pre. Tr. Py. R. Between F: 84.37 0 0.25 .00002 .00002 0.95 0.003 Pi. groups: 2836.62 60.67 5 567.32 43.49 5.91E-27 0.002 df: 60.63 0 .00002 .00002 0.03 .00002 Co. Within groups: 1839.15 39.33 141 13.04 p: 5.39E-26 0 .00008 .00002 .00002 Pre. Total: 4675.77 146 0 .00002 0.10 Tr. 0 0.06 Py. ex. Pro. Se., Si. & N. 0 R. 12. Nas-rhi Pi. Co. Pre. Tr. Py. R. N. 0 0.98 .00003 .00005 0.82 .00003 .00003 Pi. Between F: 63.18 0 .00004 0.001 0.99 0.0002 .00003 Co. groups: 2885.93 73.93 6 480.99 77.05 4.92E-45 2.19E-06 df: 62.65 0 0.95 0.0002 0.99 .00003 Pre. Within groups: 1017.6 26.07 163 6.24 p: 107E-24 0 0.01 0.99 .00003 Tr. Total: 3903.53 169 0 0.002 .00003 Py. 0 .00003 R. ex. Pro., Se. & Si. 0 N.

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Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 13. Ol-sta p(same): Co. Pre. Tr. Py. R. Between F: 88.56 0 0.0001 0.0001 0.0001 0.0001 Co. groups: 4147.25 78.60 4 1036.81 104.7 3.12E-37 0.02 df: 51.01 0 0.0001 0.0001 0.0001 Pre. Within groups: 1129.11 21.40 114 9.9 p: 2.58E-22 0 0.1 0.0001 Tr. Total: 5276.36 118 ex. 0 0.004 Py. Pi., Pro., Se., Si. & N. 0 R. 14. Zs-zgyi Pi. Co. Pre. Tr. Py. R. N. Between F: 39.87 0 1 .00003 .00003 .00003 1 0.58 Pi. groups: 2093.98 55.42 6 349.00 34.81 3.84E-27 0.00006 df: 67.6 0 .00003 .00003 .00003 0.99 0.39 Co. Within groups: 1684.26 44.58 168 10.03 p: 2.40E-20 0 0.46 1 .00003 .00003 Pre. Total: 3778.24 174 0 0.51 .00003 .00003 Tr. 0 .00003 .00003 Py. ex. Pro., Se. & Si. 0 0.73 R. 0 N. 15. Zs-zi Pi. Co. Pre. Tr. Py. R. N. Between F: 27.38 0 0.15 0.66 0.09 0.25 0.01 .00003 Pi. groups: 935.3 57.09 6 155.88 37.25 1.66E-28 1.65E-08 df: 67.82 0 0.97 1 1 .00003 .00003 Co. Within groups: 703.03 42.91 168 4.18 p: 2.44E-16 0 0.93 0.99 .00003 .00003 Pre. Total: 1638.32 174 0 0.99 .00003 .00003 Tr. 0 .00003 .00003 Py. 0 .00006 R. ex. Pro., Se. & Si. 0 N.

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Piliocolobus, Colobus, Rhinopithecus and Nasalis are not significantly different, whereas Trachypithecus, Pygathrix and Presbytis were also not significantly different. With regard to the latter (zs-zgyi), Rhinopithecus and Nasalis were significantly different for every generic pairwise comparison but the same cannot be said for Piliocolobus, Colobus, Presbytis, Trachypithecus and Pygathrix.

5.2.4 Summary for cranial variables: In summary, box-plots revealed the extent to which some generic cranial variables are related to body size, although the genera of Nasalina sometimes do not follow this generalization.

5.3 Cranial Indices: Table 5.15 provides the mean and standard deviation (SD) for several cranial indices of the Colobinae. Many scaling trends are apparent. Figures 5.17 to 5.26 are box- plots which demonstrate general increases and decreases of cranial indices between genera. Furthermore, like the previous results, by converting raw data into cranial indices, sample variation, both between species, and males and females, decreases substantially. Furthermore, there are dramatic scaling trends related to body size but the patterns are different per subtribe. For example, whereas in the Colobina and Presbytina genera, there is a general decrease in the relative percentage of cranial height (bas-br) to superior facial length (bas-pros), and maxillo-alveolar breadth (biecm) to biasterionic breadth (biast) (Figures 5.17 & 5.18), the Nasalina genera are increasing, although the position of Nasalis is peculiar (see also Figures 5.20, 5.21, 5.23). However, the contributing proportion of the maxillo-alveolar breadth (biecm) to the entire length of the palate (ol-sta) for each subtribe there is a decrease in the relative percentage with increasing body size (Figure 5.19). Still, another example illustrates a proportional similarity between Nasalina genera despite increasing body size but for the Colobina and Presbytina genera there are increasing percentages of the palate length to increases in bizygomatic breadth (Figure 5.25, ol- sta/bizygo). Indices which distinguish a genus from other subtribe members included some of the following. Nasalis and Simias both have relatively long face (nas-pros) and nasal

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Table 5.15: Generic means for cranial indices (%; indices listed in alphabetical order by abbreviation).

Pro. Pi. Co. Pre. Tr. Se. 1. bas-br/bas-nas 80.40 80.99 73.84 83.44 82.51 81.21 SD 1.87 4.30 3.88 2.73 3.33 3.78 2. bas-br/bas-pros 73.31 67.45 58.66 77.54 71.00 67.89 SD 1.11 4.20 4.09 3.64 3.88 4.25 3. bas-br/biast 112.53 114.30 109.61 110.81 118.65 119.84 SD 5.00 5.20 9.77 6.20 9.37 9.39 4. bas-br/biaur 79.28 81.14 78.71 80.45 79.40 76.19 SD 2.11 5.15 5.19 2.63 4.69 4.11 5. bas-br/g-o 62.0 63.9 62.1 65.2 64.4 62.1 SD 1.6 3.3 3.2 2.7 2.8 3.1 6. bas-br/bizygo 63.67 64.03 63.76 69.63 66.27 60.29 SD 2.23 4.16 4.44 3.41 3.19 2.55 7. bas-br/pros-o 50.58 49.31 45.46 53.39 51.43 47.91 SD 0.92 2.46 2.91 2.26 2.59 2.66 8. bas-nas/bas-pros 91.20 83.30 79.39 92.93 86.04 83.58 SD 0.94 3.31 2.04 3.13 2.52 2.94 9. bas-pros/pros-o 69.00 73.23 77.56 68.88 72.51 70.61 SD 1.04 3.45 2.19 1.50 2.90 1.84 10. biast/biaur 70.62 71.09 72.07 72.79 67.29 63.94 SD 4.87 4.92 4.51 4.38 6.47 6.24 11. biast/g-o 55.2 56.0 57.0 59.0 54.2 51.5 SD 2.6 3.1 4.5 4.4 3.6 2.7 12. biast/lam-opn 163.1 160.1 175.5 154.8 152.4 142.1 SD 7.3 10.0 17.6 16.2 11.9 8.1 13. biaur/bizygo 80.34 79.07 81.03 86.57 83.57 79.23 SD 2.87 4.98 2.79 3.63 3.14 3.28 14. biaur/pros-o 63.83 60.90 57.82 66.39 64.90 62.93 SD 1.99 3.27 2.63 2.62 3.52 2.75 15. bicanex/bas-pros 41.76 38.21 36.85 38.80 37.56 39.37 SD 2.83 3.99 2.74 1.84 3.31 2.94 16. bicanex/nas-pros 80.25 74.22 78.34 89.03 81.54 75.61 SD 11.23 7.38 8.24 6.35 7.65 6.22 17. bicanex/ol-sta 95.13 81.79 73.20 93.71 84.76 77.52 SD 4.75 7.63 6.26 4.47 8.45 5.60 18. biecm/bas-br 63.25 63.89 71.96 62.59 68.96 73.54 SD 2.27 4.30 4.00 4.29 3.35 3.99 19. biecm/bas-nas 50.85 51.63 53.03 52.18 56.85 59.67 SD 1.97 2.71 2.41 3.31 2.58 3.28 20. biecm/biast 71.17 73.00 78.67 69.39 81.76 88.22 SD 3.90 5.80 5.88 6.64 6.82 9.27 21. biecm/bicanex 127.46 111.66 113.59 131.24 124.92 131.05 SD 14.03 11.68 12.58 10.33 5.97 12.12 22. biecm/bifmt 52.49 52.60 60.32 52.29 59.69 61.97 SD 3.80 3.14 5.19 3.43 3.33 6.56 23. biecm/bizi 64.18 65.17 67.90 62.52 67.89 64.81 SD 3.06 4.64 4.31 3.19 3.11 2.05 224

Pro. Pi. Co. Pre. Tr. Se. 24. biecm/bizygo 40.31 40.84 45.76 43.47 45.64 44.30 SD 2.67 2.96 2.33 1.85 1.99 2.32 25. biecm/g-o 39.24 40.74 44.63 40.71 44.36 45.64 SD 1.80 2.06 1.91 2.07 1.92 2.45 26. biecm/ol-sta 105.82 92.09 83.66 117.03 110.33 98.17 SD 6.50 4.92 5.95 7.31 7.13 5.30 27. biecm/pros-o 32.00 31.44 32.64 33.34 35.42 35.18 SD 1.38 1.61 1.54 1.55 1.69 1.81 28. biecm/zs-zgyi 84.22 87.25 99.16 96.60 112.74 103.55 SD 7.26 7.17 6.07 12.13 11.09 10.28 29. bien/bas-br 78.55 75.71 82.36 77.97 81.43 89.94 SD 5.05 7.61 6.00 4.79 7.02 4.72 30. bien/bas-nas 63.20 61.17 60.72 65.01 67.06 72.92 SD 5.19 5.42 4.34 3.89 4.82 2.68 31. bien/bas-pros 57.61 50.92 48.25 60.40 57.74 60.95 SD 4.38 4.52 4.12 3.88 5.00 2.90 32. bien/biast 107.69 88.22 86.47 81.43 86.36 96.46 SD 8.92 3.31 8.97 8.08 6.64 9.70 33. bifmt/biaur 95.74 98.46 94.07 96.32 91.78 90.97 SD 4.44 5.21 5.66 4.63 5.62 8.37 34. bifmt/bizi 122.67 122.50 113.13 120.06 113.91 105.51 SD 8.92 6.96 9.27 10.35 5.32 10.31 35. bifmt/bizygo 76.82 77.71 76.14 83.34 76.61 71.85 SD 0.79 4.30 3.84 4.57 4.02 4.05 36. bifmt/g-o 74.92 77.56 74.39 77.97 74.46 74.05 SD 3.68 3.13 5.50 3.31 3.79 4.61 37. bifmt/nas-pros 169.95 159.22 149.07 212.97 178.31 155.78 SD 19.77 10.95 15.84 16.95 13.97 16.76 38. bifmt/pros-o 61.11 59.84 54.36 63.90 59.47 57.10 SD 3.19 2.33 3.66 3.09 3.38 3.73 39. bipor/g-o 69.4 70.7 72.0 71.7 71.0 72.8 SD 1.6 3.0 3.3 2.6 3.0 3.6 40. bizi/bas-br 98.69 98.34 106.23 100.26 101.72 113.56 SD 5.01 7.61 6.86 7.48 5.57 6.79 41. bizi/biast 111.19 112.44 116.07 111.16 120.67 136.15 SD 9.77 10.49 8.53 11.15 11.36 13.78 42. bizi/bizygo 62.85 62.87 67.55 69.63 67.30 68.36 SD 3.86 5.13 3.99 3.32 3.13 3.11 43. bizi/nas-pros 133.8 128.8 130.7 176.7 156.7 152.3 SD 3.1 11.5 8.0 6.8 11.8 5.7 44. bizs/zs-zgyi 95.75 88.78 111.70 111.31 115.53 91.37 SD 3.37 11.46 13.57 24.14 23.23 16.36 45. bizs/zs-zi 246.85 214.25 293.13 248.40 249.53 194.18 SD 16.61 39.73 43.94 72.84 59.26 55.65 46. bizs/bizi 73.38 66.19 76.16 71.32 69.03 56.92 SD 7.12 6.79 5.66 8.29 8.63 6.61

225

Pro. Pi. Co. Pre. Tr. Se. 47. bizygo/pros-o 79.52 77.19 71.37 76.72 77.67 79.45 SD 3.47 4.47 2.33 2.00 3.30 2.05 48. br-lam/g-o 46.7 44.6 47.0 45.0 43.7 42.2 SD 1.9 4.6 3.5 4.2 3.5 2.9 49. g-o/biaur 127.86 126.98 126.68 123.55 123.31 122.70 SD 4.33 5.60 5.88 3.60 5.56 5.11 50. g-o/bizygo 102.70 100.22 102.57 106.87 102.96 97.09 SD 4.50 4.88 4.31 3.33 4.08 2.36 51. g-o/pros-o 81.57 77.17 73.14 81.95 79.86 77.13 SD 2.22 1.50 2.29 2.03 1.89 2.74 52. iv-pms/bicanex 49.19 57.70 64.73 52.81 60.33 60.22 SD 5.69 7.55 8.00 4.91 7.65 5.84 53. iv-pms/ol-sta 46.62 46.83 47.04 49.50 50.65 46.53 SD 3.73 4.56 3.99 5.42 3.75 3.82 54. lam-opn/bas-br 54.66 54.75 52.61 58.84 55.43 58.54 SD 3.53 4.27 4.26 4.32 3.55 4.14 55. lam-opn/bas-pros 40.08 36.95 30.79 45.55 39.37 39.79 SD 2.96 3.87 2.47 3.01 3.56 4.26 56. lam-opn/biaur 43.38 44.38 41.32 47.28 43.99 44.60 SD 3.77 3.97 3.34 3.02 3.66 3.86 57. lam-opn/g-o 33.9 35.1 32.7 38.3 35.7 36.3 SD 2.1 2.0 2.7 2.4 2.7 2.6 58. maxnawi/nas-rhi 70.7 71.3 87.0 96.2 79.1 69.8 SD 8.8 11.7 12.4 9.9 14.2 9.0 59. nas-br/g-o 63.8 63.4 61.1 69.0 66.4 64.1 SD 3.9 2.9 3.9 2.3 3.6 2.2 60. nas-pros/biaur 56.81 62.09 63.52 45.39 51.68 58.65 SD 5.52 5.09 5.19 2.82 4.14 4.91 61. nas-pros/bien 91.64 101.74 98.64 72.61 80.51 85.99 SD 12.00 7.79 10.37 5.56 8.19 9.64 62. nas-pros/g-o 44.51 48.87 50.22 36.73 41.94 47.87 SD 5.00 2.90 4.43 1.84 3.16 4.37 63. nas-rhi/nas-pros 36.81 36.92 33.57 36.58 33.43 34.18 SD 2.84 5.13 4.10 2.84 5.94 2.38 64. ol-sta/bas-br 59.86 69.51 86.44 53.66 62.71 75.10 SD 2.09 5.29 7.84 4.95 4.37 5.56 65. ol-sta/bas-nas 48.11 56.20 63.57 44.70 51.66 60.90 SD 1.38 3.99 3.37 3.32 2.89 4.05 66. ol-sta/bas-pros 43.87 46.74 50.42 41.46 44.45 50.81 SD 1.22 2.58 1.92 2.07 2.87 2.15 67. ol-sta/bizygo 38.10 44.44 54.86 37.24 41.48 45.19 SD 1.38 3.63 3.45 2.16 2.33 2.51 68. palhei/ol-sta 21.10 18.22 17.82 20.90 21.16 18.42 SD 3.00 3.37 1.88 2.04 3.62 3.25 69. zs-zgyi/bizi 76.71 75.17 68.67 65.66 60.76 63.02 SD 8.04 8.25 5.43 8.61 6.37 5.24

226

Pro. Pi. Co. Pre. Tr. Se. 70. zs-zgyi/bizygo 47.99 46.89 46.23 45.53 40.76 43.15 SD 2.73 2.11 2.33 4.69 3.31 4.80 71. zs-zgyi/ol-sta 126.14 106.08 84.48 122.94 98.56 95.28 SD 9.40 8.64 5.34 16.68 9.45 6.65 72. zs-zi/bizi 29.71 31.53 26.27 30.36 28.45 30.69 SD 1.82 4.17 2.33 6.10 3.75 5.60 73. zs-zi/bizygo 18.66 19.75 17.73 21.10 19.10 21.04 SD 1.51 2.67 1.65 4.18 2.29 4.30 74. zs-zi/zs-zgyi 38.95 42.08 38.38 46.36 46.96 48.40

SD 3.25 4.88 3.48 7.71 5.12 5.53

227

Table 5.15 concluded: Colobinae cranial indices (%; indices listed in alphabetical order by abbreviation). Si. Py. R. N. 1. bas-br/bas-nas 77.59 82.43 81.84 76.79 SD 3.54 3.96 4.10 4.86 2. bas-br/bas-pros 65.58 68.86 71.69 61.18 SD 3.91 4.48 4.91 5.72 3. bas-br/biast 103.32 104.19 99.36 105.13 SD 5.56 6.23 6.98 8.05 4. bas-br/biaur 76.55 79.02 74.25 75.15 SD 2.35 3.54 2.83 4.61 5. bas-br/g-o 59.8 62.9 59.4 59.2 SD 1.7 2.8 2.4 2.9 6. bas-br/bizygo 64.76 67.90 63.41 62.13 SD 2.39 3.30 4.51 4.44 7. bas-br/pros-o 46.43 49.27 49.21 43.95 SD 2.15 2.36 2.16 3.45 8. bas-nas/bas-pros 84.50 83.52 87.55 79.55 SD 2.32 3.20 2.81 3.00 9. bas-pros/pros-o 70.85 71.68 68.76 71.94 SD 1.56 2.91 2.32 1.67 10. biast/biaur 74.29 76.03 74.97 71.70 SD 4.96 4.61 4.42 4.64 11. biast/g-o 58.9 60.5 59.9 56.3 SD 2.6 3.2 1.7 3.5 12. biast/lam-opn 158.8 169.8 179.2 174.1 SD 1.8 12.9 18.2 11.3 13. biaur/bizygo 84.64 85.99 85.36 82.69 SD 3.59 3.45 4.23 3.47 14. biaur/pros-o 60.63 62.40 66.28 58.47 SD 1.14 2.48 1.69 2.53 15. bicanex/bas-pros 35.24 36.42 38.78 36.03 SD 2.64 2.97 2.65 2.12 16. bicanex/nas-pros 63.85 74.24 84.50 63.87 SD 7.31 7.21 9.38 4.80 17. bicanex/ol-sta 80.77 85.20 79.75 84.10 SD 7.08 7.11 5.46 4.80 18. biecm/bas-br 75.73 69.21 70.14 70.37 SD 4.86 3.51 3.68 4.07 19. biecm/bas-nas 58.67 57.00 57.28 53.92 SD 3.18 3.14 1.70 2.94 20. biecm/biast 78.31 71.95 69.64 73.90 SD 7.53 3.00 5.58 6.17 21. biecm/bicanex 141.43 131.24 129.95 119.49 SD 14.30 10.33 10.89 10.97 22. biecm/bifmt 60.48 52.43 52.07 59.11 SD 5.42 2.90 1.54 3.43 23. biecm/bizi 70.25 69.21 64.79 66.75 SD 4.94 3.69 2.35 4.34

228

Si. Py. R. N. 24. biecm/bizygo 49.03 46.93 44.37 43.65 SD 3.59 2.22 2.31 3.08 25. biecm/g-o 45.27 43.64 41.78 41.63 SD 3.27 2.44 1.82 1.63 26. biecm/ol-sta 113.55 110.05 103.15 100.25 SD 6.83 6.23 4.71 8.31 27. biecm/pros-o 35.11 34.05 34.46 30.83 SD 1.88 1.46 1.20 1.78 28. biecm/zs-zgyi 100.66 116.62 101.05 94.50 SD 7.36 7.85 5.16 7.45 29. bien/bas-br 74.95 78.11 86.98 85.93 SD 5.54 5.64 4.27 7.96 30. bien/bas-nas 58.04 64.33 71.18 65.77 SD 3.08 4.92 4.82 4.84 31. bien/bas-pros 49.04 53.77 62.33 52.32 SD 2.85 4.96 4.83 4.35 32. bien/biast 77.46 81.43 86.54 90.12 SD 7.26 8.08 8.58 8.66 33. bifmt/biaur 96.15 104.26 99.91 89.38 SD 6.50 2.97 3.07 3.92 34. bifmt/bizi 116.59 132.18 124.49 113.02 SD 9.19 7.08 5.06 5.88 35. bifmt/bizygo 81.29 89.60 85.28 73.86 SD 4.89 3.34 4.90 3.51 36. bifmt/g-o 75.05 83.27 80.23 70.57 SD 4.44 2.88 2.14 3.40 37. bifmt/nas-pros 149.59 185.46 210.27 129.13 SD 20.63 16.64 22.88 13.76 38. bifmt/pros-o 58.32 65.04 66.19 52.24 SD 4.52 2.81 1.97 2.88 39. bipor/g-o 66.5 70.0 69.4 70.5 SD 2.7 3.8 2.1 2.9 40. bizi/bas-br 107.92 100.19 108.38 105.74 SD 4.64 5.91 6.33 7.79 41. bizi/biast 111.43 104.29 107.58 110.91 SD 6.08 7.72 8.75 9.11 42. bizi/bizygo 69.86 67.94 68.54 65.52 SD 3.42 3.91 3.75 4.52 43. bizi/nas-pros 123.2 140.5 169.2 114.5 SD 6.5 12.9 19.9 12.7 44. bizs/zs-zgyi 70.89 141.58 110.43 81.33 SD 8.01 12.42 8.38 9.55 45. bizs/zs-zi 127.47 311.47 234.87 155.48 SD 11.60 35.85 22.96 24.77 46. bizs/bizi 49.28 84.00 70.71 57.39 SD 2.67 6.34 2.87 5.62

229

Si. Py. R. N. 47. bizygo/pros-o 71.73 72.62 77.77 70.76 SD 3.17 2.72 3.10 2.87 48. br-lam/g-o 33.1 42.8 40.6 38.5 SD 2.9 2.6 1.9 2.1 49. g-o/biaur 128.07 125.29 124.57 126.79 SD 2.81 3.83 4.02 5.33 50. g-o/bizygo 108.41 107.69 106.35 104.82 SD 5.36 4.56 6.58 5.66 51. g-o/pros-o 77.67 78.13 82.54 74.07 SD 2.85 2.86 2.59 3.14 52. iv-pms/bicanex 66.31 55.31 58.71 64.20 SD 8.62 13.35 11.73 5.23 53. iv-pms/ol-sta 53.14 47.07 46.43 53.84 SD 4.45 10.88 7.53 3.28 54. lam-opn/bas-br 61.50 56.87 56.82 54.20 SD 2.42 3.08 4.99 3.71 55. lam-opn/bas-pros 40.37 39.14 40.70 33.11 SD 3.38 3.10 4.31 3.30 56. lam-opn/biaur 47.10 44.90 42.18 40.67 SD 2.76 2.61 4.00 2.98 57. lam-opn/g-o 37.1 35.7 33.7 32.4 SD 1.9 1.8 3.0 2.0 58. maxnawi/nas-rhi 62.2 88.2 131.3 59.5 SD 8.6 13.4 24.7 7.5 59. nas-br/g-o 66.8 66.9 69.6 67.9 SD 2.6 3.7 2.5 2.4 60. nas-pros/biaur 64.87 56.63 48.02 69.84 SD 5.36 5.06 5.12 6.73 61. nas-pros/bien 113.15 92.40 74.74 108.96 SD 5.54 11.43 10.29 12.42 62. nas-pros/g-o 50.73 45.23 38.63 55.15 SD 5.14 4.07 4.83 5.54 63. nas-rhi/nas-pros 45.87 35.26 20.37 46.60 SD 3.30 4.68 14.86 3.62 64. ol-sta/bas-br 66.82 62.38 68.16 70.67 SD 4.74 4.73 4.95 7.22 65. ol-sta/bas-nas 51.75 51.37 55.61 53.98 SD 2.58 3.87 2.25 3.15 66. ol-sta/bas-pros 43.68 42.83 48.64 42.86 SD 1.45 2.62 1.39 1.45 67. ol-sta/bizygo 43.21 42.27 43.04 43.65 SD 2.32 2.69 1.90 2.63 68. palhei/ol-sta 22.40 18.92 14.06 17.58 SD 4.42 2.67 1.57 3.03 69. zs-zgyi/bizi 69.98 59.49 64.27 70.84 SD 5.37 3.65 3.96 4.58

230

Si. Py. R. N. 70. zs-zgyi/bizygo 48.77 40.34 43.96 46.29 SD 2.49 2.08 2.23 2.70 71. zs-zgyi/ol-sta 113.03 95.72 102.20 106.15 SD 6.47 6.70 4.62 4.71 72. zs-zi/bizi 38.83 27.14 30.31 37.40 SD 2.58 1.97 2.43 3.91 73. zs-zi/bizygo 27.11 18.43 20.76 24.49 SD 1.85 1.57 1.93 2.95 74. zs-zi/zs-zgyi 55.80 45.72 47.35 52.84

SD 6.01 3.70 5.04 4.91

(nas-rhi) and zygomatic (zs-zgyi) bones (Figures 5.20 & 5.23). Conversely, Rhinopithecus and Pygathrix are opposite to conditions described above for Nasalis and Simias. The small Olive-colobine (Procolobus) has a maxillo-alveolar breadth (biecm) that is relatively equal to or greater than palatal length (ol-sta), and a cranial height (bas-br) that is relatively larger in relation to superior facial length (bas-pros), compared to the larger species of Piliocolobus and Colobus (Figures 5.17 & 5.19). These comparisons are also similar for the equally small Surilis (Presbytis) in relation to the larger specie of Semnopithecus and Trachypithecus. Pygathrix has the widest bizygomaxillare superior breadth (bizs) in relation to bizygomaxillare inferior breadth (bizi) (Figure 5.22). Rhinopithecus has a relatively small facial height (nas-pros) in relation to superior facial breadth (bifmt) and bizygomaxillare inferior breadth (bizi) (Figures 5.20 & 5.21).

5.3.1 Shapiro-Wilk results for cranial indices: Table 5.16 lists the results of applying Shapiro-Wilk to several cranial indices. Only five indices for three genera were not normally distributed. The relative proportion of the maxillo-alveolar breadth (biecm) to biasterionic breadth (biast) and the maximum length of the zygomatic in relation to palatal length A (ol-sta) for Piliocolobus were not normally distributed. The relative proportion of cranial height (bas-br) to superior facial length (bas-pros) and the maximum length of the zygomatic in relation to palatal length A (ol-sta) for Presbytis were not normally distributed. Finally, the superior facial height (nas- pros) in relative proportion to cranial breadth (biaur) for Trachypithecus was also not normally distributed.

231

Box Plots for cranial indices of the Colobinae (Colobina: Pro. – Procolobus, n=5; Pi. – Piliocolobus, n=28; Co. – Colobus, n=26; Presbytina: Pre. – Presbytis, n=18; Tr. – Trachypithecus, n=33; Se. – Semnopithecus, n=9; & Nasalina: Si. – Simias, n=7; Py. - Pygathrix, n=25; R. – Rhinopithecus, n=17; & N. – Nasalis, n=28; indices in alphabetical order by abbreviation): Figure 5.17 Pre. 80 Tr. Se. Py. R. Pro. Pi. N.

70 Si.

Co. %

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: bas-br/bas-pros x 100

100 Figure 5.18 Se. Tr.

Co.

90 Si. N.

Pre. Pi. 80 Py. % Pro. R.

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: biecm/biast x 100

232

Figure 5.19

130 Pre. Py. Tr. 120 Si. Pro. N.

110 R. Se. Pi. % 100 Co.

70 0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: biecm/ol-sta x 100

Figure 5.20 270 260 250 Pre. R. 240 230 220 Py. 210 Tr 200 Pro. 190

% Pi. Si. 180 Co. Se. 170 N. 160 150 140 130 120 110

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: bifmt/nas-pros x 100

233

200 Figure 5.21 R. Pre 190 . 180 Tr. 170 Py. Se. 160 Pi. 150 Co. N. % 140 Pro. Si. 130 120 110 100

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: bizi/nas-pros x 100

Figure 5.22 100 Py. 90 Co. Pre. Tr. Pro. Pi. 80 R. N.

% 70 Se.

60 Si. 50

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: bizs/bizi x 100

234

90 Figure 5.23 N.

80 Pi. Co. Si. 70 Py.

% Tr. Pro. Se. 60 R.

Pre. 50

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: nas-pros/biaur x 100

60 Figure 5.24 59 58 57 56 Co. 55 Se. 54 53 52 Pi. R. 51 50 Tr. 49 48 % 47 Si. Py. 46 Pro. Pre. N. 45 44 43 42 41 40 39 38 37 36

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: ol-sta/bas-pros x 100

235

Figure 5.25 60 Co.

Pi. Se. 50 N. Si. R. Tr. Py. %

Pre. Pro 40 .

0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: ol-sta/bizygo x 100

Figure 5.26 150 Pre.

140

130 Pro.

120 Pi. Tr. Si. N. R.

% 110 Py. Se. 100 Co. 90

70 0 1 2 3 4 5 6 7 8 9 10 11 Colobinae: zs-zgyi/ol-sta x 100

236

Table 5.16: Shapiro-Wilk results for cranial indices of the Colobinae; those in bold represent significant departures from normality (indices listed in alphabetical order by abbreviation).

Pro. Pi. Co. Pre. Tr. Se. n=5 n=28 n=26 n=18 n=33 n=9 1. Bas-br/bas-pros W: 0.97 0.98 0.95 0.88 0.99 0.87 p(normal): 0.86 0.87 0.27 0.02 0.93 0.11 2. Biecm/biast W: 0.91 0.92 0.95 0.94 0.97 0.89 p(normal): 0.45 0.03 0.24 0.3 0.45 0.19 3. Biecm/ol-sta W: 0.87 0.98 0.99 0.97 0.96 0.89 p(normal): 0.26 0.81 0.99 0.81 0.2 0.18 4. Bifmt/nas-pros W: 0.84 0.98 0.98 0.9 0.98 0.92 p(normal): 0.17 0.96 0.94 0.07 0.89 0.39 5. Bizi/nas-pros W: 0.96 0.98 0.97 0.98 0.97 0.88 p(normal): 0.78 0.83 0.5 0.94 0.49 0.14 6. Bizs/bizi W: 0.96 0.98 0.95 0.95 0.99 0.94 p(normal): 0.77 0.85 0.25 0.45 0.92 0.62 7. Nas-pros/biaur W: 0.85 0.98 0.98 0.97 0.93 0.85 p(normal): 0.19 0.93 0.85 0.7 0.04 0.07 8. Ol-sta/bas-pros W: 0.98 0.97 0.94 0.95 0.96 0.99 p(normal): 0.92 0.5 0.12 0.38 0.26 0.99 9. Ol-sta/bizygo W: 0.95 0.97 0.94 0.97 0.99 0.95 p(normal): 0.72 0.69 0.13 0.77 0.93 0.64 10. Zs-zgyi/ol-sta W: 0.97 0.9 0.96 0.89 0.97 0.92 p(normal): 0.9 0.01 0.47 0.04 0.44 0.4

Si. Py. R. N. n=7 n=25 n=17 n=28 1. Bas-br/bas-pros W: 0.9 0.95 0.96 0.91 p(normal): 0.34 0.32 0.68 0.03 2. Biecm/biast W: 0.91 0.98 0.97 0.95 p(normal): 0.38 0.88 0.81 0.22 3. Biecm/ol-sta W: 0.82 0.97 0.95 0.95 p(normal): 0.06 0.69 0.39 0.23 4. Bifmt/nas-pros W: 0.94 0.98 0.96 0.94 p(normal): 0.69 0.84 0.57 0.12 5. Bizi/nas-pros W: 0.87 0.99 0.94 0.96 p(normal): 0.18 0.97 0.35 0.39 6. Bizs/bizi W: 0.86 0.95 0.86 0.95 p(normal): 0.15 0.23 0.04 0.21 7. Nas-pros/biaur W: 0.94 0.97 0.95 0.97 p(normal): 0.61 0.56 0.44 0.66 8. Ol-sta/bas-pros W: 0.97 0.96 0.93 0.98 p(normal): 0.93 0.54 0.21 0.83 9. Ol-sta/bizygo W: 0.93 0.95 0.97 0.98 p(normal): 0.59 0.22 0.79 0.92 10. Zs-zgyi/ol-sta W: 0.79 0.95 0.97 0.98 p(normal): 0.03 0.32 0.86 0.85

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5.3.2 Kruskal-Wallis and Mann-Whitney results for cranial indices: Table 5.17 provides the results of applying Kruskal-Wallis and Mann-Whitney to several cranial indices of the Colobinae to test for significant differences in generic medians. Akin to the previous results for the previous groupings, there are more significant differences than non-significant comparisons. For example, cranial height (bas-br) in relative proportion to the superior facial length (bas-pros) between the Piliocolobus and Semnopithecus samples are virtually identical (Table 5.17, no. 1). Likewise, the superior facial height (nas-pros) in relation to maximum cranial breadth (biaur) between the Procolobus and Semnopithecus samples are virtually identical (Table 5.17, no. 7). Similarly, there were near equal results between Procolobus and Presbytis, and Piliocolobus and Nasalis, with regard to the maximum length of the zygomatic (zs-zgyi) in relative proportion to palatal length A (ol-sta; (Table 5.17, no. 10). Surprisingly, despite their differences in absolute size, Nasalis compared to Procolobus, Piliocolobus, Simias and Pygathrix are not significantly different (based on medians) in the relative difference between maxillo-alveolar (biecm) and biasterionic (biast) breadths (Table 5.17, no. 2). As the box-plots indicated (Figure 5.20), Nasalis is significantly distinct from all other colobines in regard to the relative proportion of the superior facial breadth (bifmt) to superior facial height (nas-pros; Table 5.20, no. 4)). Simias and Pygathrix are extremes to one another in relationship between bizygomaxillare superior (bizs) and inferior (bizi) breadths. In the former, the superior breadth is nearly 50% of the inferior breadth, whereas in the latter it’s ~85%; all pairwise generic comparisons for these two genera are significantly different from the other colobine genera and are in between these extremes (Table 5.17, no. 6). Rhinopithecus is separated from the other odd-nosed colobines by the relative proportion of its palatal length A (ol-sta) to superior facial length (bas-pros) and is also significantly different from all colobine genera (Figure 5.25 and Table 5.17, no. 8).

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Table 5.17: Kruskal-Wallis results for cranial indices of the Colobinae with Mann-Whitney pairwise comparisons (p (same)) (indices listed in alphabetical order by abbreviation).

Pro. Pi. Co. Pre. Tr. Se. Si. Py. R. N. n=5 n=28 n=26 n=18 n=33 n=9 n=7 n=25 n=17 n=28 1. Bas-br/bas-pros 0 0.004 0.0005 0.005 0.19 0.05 0.006 0.04 0.12 0.001 Pro. H: 122.9 0 4.71E-08 3.28E-08 0.002 1 0.33 0.29 0.003 0.00009 Pi. Hc: 122.9 0 2.49E-08 9.11E-11 0.00003 0.002 1.27E-08 4.32E-09 0.30 Co. p(same): 3.44E-22 0 3.14E-07 0.0001 0.0002 2.19E-07 3.53E-06 1.96E-08 Pre. 0 0.02 0.005 0.09 0.68 7.90E-08 Tr. 0 0.60 0.49 0.04 0.007 Se. 0 0.14 0.001 0.05 Si 0 0.04 0.00002 Py. 0 2.39E-06 R. 0 N. 2. Biecm/biast 0 0.63 0.02 0.35 0.007 0.009 0.10 0.93 0.18 0.38 Pro. H: 83.18 0 0.0006 0.06 4.63E-06 0.0002 0.08 0.90 0.006 0.51 Pi. Hc: 83.18 0 0.00007 0.07 0.01 0.86 0.00005 3.54E-07 0.01 Co. p(same): 3.76E-14 0 2.35E-06 0.0002 0.01 0.04 0.99 0.02 Pre. 0 0.04 0.30 2.60E-07 2.54E-08 0.0001 Tr. 0 0.06 0.00007 0.00005 0.0006 Se. 0 0.02 0.002 0.17 Si 0 0.0009 0.27 Py. 0 0.003 R. 0 N.

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Pro. Pi. Co. Pre. Tr. Se. Si. Py. R. N. n=5 n=28 n=26 n=18 n=33 n=9 n=7 n=25 n=17 n=28 3. Biecm/ol-sta 0 0.0007 0.0005 0.01 0.23 0.02 0.10 0.16 0.31 0.13 Pro. H: 140.8 0 5.51E-06 1.51E-08 5.79E-11 0.005 0.00006 1.17E-09 1.90E-06 0.0003 Pi. Hc: 140.8 0 2.49E-08 6.06E-11 0.00004 0.00007 1.09E-09 8.65E-08 6.32E-09 Co. p(same): 7.18E-26 0 0.004 0.00004 0.29 0.003 1.33E-06 1.23E-06 Pre. 0 0.0002 0.13 0.99 0.0002 0.00003 Tr. 0 0.002 0.0001 0.09 0.82 Se. 0 0.24 0.002 0.002 Si. 0 0.00006 0.0001 Py. 0 0.33 R. 0 N. 4. Bifmt/nas-pros 0 0.38 0.02 0.002 0.36 0.29 0.19 0.12 0.008 0.0008 Pro. H: 152.1 0 0.0006 1.51E-08 1.85E-06 0.58 0.24 1.05E-07 8.83E-08 1.02E-08 Pi. Hc: 152.1 0 2.49E-08 1.71E-09 0.20 0.88 4.90E-09 4.97E-08 0.00001 Co. p(same): 3.27E-28 0 2.57E-08 0.00003 0.0002 0.00002 0.78 1.51E-08 Pre. 0 0.002 0.002 0.01 0.00001 3.23E-11 Tr. 0 0.6 0.002 0.00007 0.0006 Se. 0 0.0005 0.0004 0.02 Si. 0 0.001 9.60E-10 Py. 0 2.70E-08 R. 0 N.

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Pro. Pi. Co. Pre. Tr. Se. Si. Py. R. N. n=5 n=28 n=26 n=18 n=33 n=9 n=7 n=25 n=17 n=28 5. Bizi/nas-pros 0 0.41 0.30 0.0009 0.0007 0.003 0.009 0.12 0.003 0.003 Pro. H: 144.5 0 0.53 1.51E-08 1.45E-09 0.00002 0.29 0.003 4.54E-07 0.0002 Pi. Hc: 144.5 0 2.49E-08 1.07E-09 0.00003 0.03 0.003 6.34E-07 0.00001 Co. p(same): 1.22E-23 0 4.52E-07 0.00003 0.0002 3.75E-08 0.45 1.51E-08 Pre. 0 0.17 0.00004 0.00003 0.01 4.54E-11 Tr. 0 0.001 0.01 0.02 0.00001 Se. 0 0.002 0.0002 0.08 Si. 0 0.00004 1.15E-07 Py. 0 4.60E-08 R. 0 N. 6. Bizs/bizi 0 0.013 0.77 0.35 0.15 0.003 0.006 0.003 0.10 0.0006 Pro. H: 129.7 0 2.19E-06 0.05 0.11 0.002 0.00007 1.17E-09 0.01 0.00001 Pi. Hc: 129.7 0 0.07 0.001 0.00001 0.00007 4.90E-06 0.003 6.76E-10 Co. p(same): 1.36E-23 0 0.51 0.0006 0.0002 2.28E-06 0.78 1.30E-06 Pre. 0 0.0004 0.00004 3.91E-09 0.57 2.67E-07 Tr. 0 0.01 0.00001 0.00008 0.87 Se. 0 0.00007 0.0002 0.0008 Si. 0 5.56E-08 4.75E-10 Py. 0 1.38E-07 R. 0 N.

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Pro. Pi. Co. Pre. Tr. Se. Si. Py. R. N. n=5 n=28 n=26 n=18 n=33 n=9 n=7 n=25 n=17 n=28 7. Nas-pros/biaur 0 0.09 0.01 0.002 0.05 1 0.023 0.70 0.01 0.002 Pro. H: 140.8 0 0.30 1.51E-08 5.00E-09 0.04 0.22 0.0002 1.01E-07 0.00004 Pi. Hc: 140.8 0 2.86E-08 2.49E-09 0.013 0.64 0.00004 1.30E-07 0.0009 Co. p(same): 6.94E-26 0 1.84E-06 0.00003 0.0002 1.47E-07 0.15 1.51E-08 Pre. 0 0.0006 0.0001 0.0002 0.02 5.26E-11 Tr. 0 0.04 0.23 0.0004 0.0002 Se. 0 0.003 0.0003 0.08 Si. 0 0.00002 2.86E-08 Py. 0 2.70E-08 R. 0 N. 8. Ol-sta/bas-pros 0 0.02 0.0005 0.03 0.67 0.003 0.87 0.47 0.001 0.23 Pro. H: 129.8 0 3.64E-06 9.27E-07 0.003 0.0006 0.005 0.00001 0.003 3.32E-07 Pi. Hc: 129.8 0 2.49E-08 3.67E-10 0.49 0.00007 9.70E-10 0.002 3.12E-10 Co. p(same): 1.29E-23 0 0.0001 0.00003 0.02 0.003 5.73E-07 0.01 Pre. 0 0.00002 0.46 0.07 2.47E-06 0.01 Tr. 0 0.001 0.00001 0.009 8.89E-06 Se. 0 0.65 0.0002 0.21 Si. 0 1.30E-07 0.46 Py. 0 3.53E-08 R. 0 N.

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Pro. Pi. Co. Pre. Tr. Se. Si. Py. R. N. n=5 n=28 n=26 n=18 n=33 n=9 n=7 n=25 n=17 n=28 9. Ol-sta/bizygo 0 0.001 0.0005 0.31 0.003 0.003 0.006 0.002 0.002 0.001 Pro. H: 117.1 0 8.41E-10 1.57E-07 0.001 0.45 0.42 0.02 0.23 0.46 Pi. Hc: 117.1 0 2.49E-08 6.06E-11 0.00001 0.00007 9.70E-10 4.32E-08 3.12E-10 Co. p(same): 5.25E-21 0 1.24E-06 0.00003 0.0004 7.98E-07 1.57E-06 9.02E-08 Pre. 0 0.001 0.12 0.44 0.03 0.0004 Tr. 0 0.14 0.006 0.05 0.13 Se. 0 0.36 1 0.71 Si. 0 0.17 0.05 Py. 0 0.44 R. 0 N. 10. Zs-zgyi/ol-sta 0 0.0007 0.0005 0.97 0.0008 0.003 0.02 0.0005 0.001 0.0007 Pro. H: 117.9 0 1.30E-09 0.001 0.002 0.0009 0.006 4.11E-06 0.04 0.99 Pi. Hc: 117.9 0 2.86E-08 8.73E-08 0.0004 0.00007 1.48E-06 4.97E-08 3.12E-10 Co. p(same): 3.67E-21 0 8.03E-06 0.0007 0.12 6.26E-06 0.002 0.003 Pre. 0 0.48 0.002 0.45 0.04 0.0003 Tr. 0 0.001 0.91 0.01 0.0003 Se. 0 0.00007 0.0005 0.009 Si. 0 0.0009 1.27E-07 Py. 0 0.02 R. 0 N.

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5.3.3 One-way ANOVA results for cranial indices: Table 5.18 provides the results of applying one-way ANOVA to several cranial indices of the Colobinae. Again, Procolobus, Semnopithecus and Simias were excluded due to their small sample sizes. Half of the indices examined produced percentages of variation between groups > 70%, while the other half generated percentages ranging from >40% to <70%. Many indices highlight some similar cranial proportions between genera but many more significant differences based on the mean of the genus sample. For example, cranial height (bas-br) in relation to superior facial length (bas-pros) for Rhinopithecus and Trachypithecus was nearly identical (Table 5.18, no. 1). Likewise, the maxillo-alveolar breadth (biecm) in relative proportion to biasterionic breadth (biast) for Rhinopithecus and Presbytis was virtually the same (Table 5.18, no. 2). In another example, the bizygomaxillare inferior breadth (bizi) in relation to superior facial height (nas-pros) was nearly the same for Piliocolobus and Colobus (Table 5.18, no. 5). The relative proportion of maxillo-alveolar breadth (biecm) to palatal length A (ol-sta) for Pygathrix and Trachypithecus, and Nasalis and Rhinopithecus were also indistinguishable (Table 5.18, no. 3). Lastly, the relative size of the zygomatic bone’s length (zs-zgyi) in relation to the palate’s length (ol-sta) is not significantly different between Pygathrix, Rhinopithecus and Nasalis but significantly different to the Colobus and Trachypithecus (Table 5.18, no. 10). Facial dimensions of the colobines are highly distinctive. Significant differences between the genus sample means which further distinguish some colobine genera include the following. The relative proportions of bizygomaxillare inferior (bizi) to superior facial height (nas-pros); and bizygomaxillare superior (bizs) to bizygomaxillare inferior; and superior facial breadth (bifmt) to superior facial height (nas-pros) produced striking results which can easily separate colobine genera (Table 5.18, no. 4, 5 & 6). As noted above in the section of Kruskal-Wallis and Mann-Whitney results for cranial indices, it is interesting that the odd-nosed colobines of Nasalina have very similar proportions of their palate length (ol-sta) in relation to bizygomatic breadth (bizygo) but in the Colobina and Presbytina there are definite scaling trends. As such, Nasalina genera (although Trachypithecus and Piliocolobus also have similar percentages as the odd-nosed genera tested) are not significantly different to one another but significantly different to Colobus and Presbytis (Table 5.18, no. 9).

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Table 5.18: One-Way ANOVA results for cranial indices of the Colobinae (indices listed in alphabetical order by abbreviation). Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): p(same): Pi. Co. Tr. Py. R. 1. Bas-br/bas-pros 0 .00001 0.02 0.74 0.005 Between F: 42.83 0 .00001 .00001 .00001 groups: 2742.7 57.70 4 685.68 42.28 2.51E-22 0.55 df: 58.54 0 0.33 0.99 Within groups: 2010.84 42.30 124 16.22 p: 9.54E-17 0 0.16 Total: 4753.54 128 ex. Pro. Se., Si. & N. 0 Co. Pre. Tr. Py. R. N. 2. Biecm/biast 0 .00002 0.42 0.0007 .00002 0.05 Between F: 24.57 0 .00002 0.63 0.99 0.07 groups: 3450.5 43.44 5 690.1 21.66 4.81E-16 0.0003 df: 61.36 0 .00002 .00002 .00005 Within groups: 4493.14 56.56 141 31.87 p: 1.71E-13 0 0.24 0.85 Total: 7943.64 146 0 0.01 ex. Pro., Si., Se. & Pi. 0 3. Biecm/ol-sta Pi. Co. Pre. Tr. Py. R. N. Between F: 82.98 0 0.0006 .00002 .00002 .00002 .00003 .0003 groups: 18648.8 72.57 6 3108.13 74.06 1.27E-44 0.004 df: 69.9 0 .00002 .00002 .00002 .00002 .00002 Within groups: 7050.26 27.43 168 41.97 p: 8.26E-30 0 0.003 0.003 .00002 .00002 Total: 25699 174 0 1 0.0003 .00003 0 0.0003 .00003 0 0.99 ex. Pro., Si. & Se. 0

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Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. F comparisons sqrs: Var.: df: square: F: p(same): of Var. test: p(same): 4. Bifmt/nas-pros p(same): Pi. Co. Pre. Tr. Py. R. N. Between F: 87.14 0 0.08 .00003 0.005 .00003 .00003 .00003 Pi. groups: 132365 78.98 6 22060.8 105.2 2.95E-54 0.0008 df: 67.53 0 .00003 .00003 .00003 .00003 0.0003 Co. Within groups: 35237.1 21.02 168 209.75 p: 7.91E-30 0 .00003 .00003 0.99 .00003 Pre. Total: 167602 174 0 0.27 .00003 .00003 Tr. 0 .00003 .00003 Py. 0 .00003 R. ex. Pro., Si. & Se. 0 N. 5. Bizi/nas-pros Pi. Co. Pre. Tr. Py. R. N. Between F: 119.1 0 0.99 .00003 .00003 0.02 .00003 0.001 Pi. groups: 70810.2 73.86 6 11801.7 79.1 2.30E-46 .00004 df: 69.61 0 .00003 .00003 0.08 .00003 0.0001 Co. Within groups: 25064.5 26.14 168 149.19 p: 1.34E-34 0 .00003 .00003 0.35 .00003 Pre. Total: 95874.7 174 0 .0001 0.007 .00003 Tr. 0 .00003 .00003 Py. 0 .00003 R. ex. Pro., Si. & Se. 0 N. 6. Bizs/bizi Pi. Co. Pre. Tr. Py. N. Between F: 85 0 .00002 0.06 0.53 .00002 0.0004 Pi. groups: 11020.5 62.76 5 2204.1 51.23 6.43E-31 0.02 df: 66.78 0 0.09 0.03 0.0001 .00002 Co. Within groups: 6539.73 37.24 152 43.02 p: 1.38E-27 0 0.88 .00002 .00002 Pre. Total: 17560.2 157 0 .00002 .00002 Tr. 0 .00002 Py. ex. Pro., Si., R. & Se. 0 N.

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Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 7. Nas-pros/biaur p(same): Pi. Co. Pre. Py. R. N. Between F: 96.41 0 0.94 .00002 0.006 .00002 .00002 Pi. groups: 9607.75 71.84 5 1921.55 69.39 1.00E-35 0.11 df: 60.95 0 .00002 0.0002 .00002 0.0008 Co. Within groups: 3766.04 28.16 136 27.69 p: 1.28E-27 0 .00002 0.54 .00002 Pre. Total: 13373.8 141 0 .00002 .00002 Py. 0 .00002 R. ex. Pro., Si., Tr. & Se. 0 N. 8. Ol-sta/bas-pros Pi. Co. Pre. Tr. Py. R. N. Between F: 74.62 0 .00003 .00003 0.006 .00003 0.02 .00003 Pi. groups: 1470.95 67.60 6 245.16 58.42 1.30E-38 .001 df: 70.18 0 .00003 .00003 .00003 0.04 .00003 Co. Within groups: 705.056 32.40 168 4.2 p: 1.76E-28 0 .00003 0.02 .00003 0.21 Pre. Total: 2176 174 0 0.35 .00003 0.05 Tr. 0 .00003 0.98 Py. 0 .00003 R. ex. Pro., Si. & Se. 0 N. 9. Ol-sta/bizygo Pi. Co. Pre. Tr. Py. R. N. Between F: 108.1 0 .00002 .00003 0.003 0.04 0.50 0.94 Pi. groups: 3922.41 78.00 6 653.74 99.29 1.28E-52 0.03 df: 70.33 0 .00003 .00003 .00003 .00003 .00003 Co. Within groups: 1106.08 22.00 168 6.58 p: 1.76E-33 0 .00003 .00003 .00003 .00003 Pre. Total: 5028.49 174 0 0.99 0.49 0.10 Tr. 0 0.92 0.45 Py. 0 0.98 R. ex. Pro., Si. & Se. 0 N. 10. Zs-zgyi/ol-sta Co. Tr. Py. R. N. Between F: 65 0 .00002 .00002 .00002 .00002 Co. groups: 7003.53 54.45 4 1750.88 37.06 2.33E-20 .005 df: 58.81 0 0.56 0.25 0.0005 Tr. Within groups: 5858.57 45.55 124 47.25 p: 6.49E-21 0 0.004 .00002 Py. Total: 12862.1 128 0 0.26 R. ex. Pro., Pi., Si., Pre. & Se. 0 N.

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5.3.4 Summary for the cranial indices: In summary, like the cranial variables, some indices also displayed scaling trends related to body size. However, again, some indices for the Nasalina do not conform to this statement. Additionally, levels of variation decrease for indices when compared to cranial variables.

5.4 Bivariate Results: Table 5.19 presents the results of plotting the mean generic body weight against the generic mean for particular cranial variables. Compared to the other primate groupings examined thus far, the Colobinae produce the least number of cranial variables which are significantly related to increasing in body size. Figure 5.27 is a bivariate plot of mean generic body weight plotted against cranial vault length (g-o). Only one figure is provided because very few variables were significantly correlated to Ln generic mean body weight. The cause for so few cranial variables significantly correlated to body size is caused by the inclusion of the Nasalina genera. For example, Rhinopithecus includes one of the largest cercopithecines, in terms of body weight, (e.g. R. roxellanae) and yet this species has a small face (nas-pros; Figure 5.12) and very short nasal bones (nas-rhi; Figure 5.13). Another example would be N. larvatus. Again, this is one of the largest cercopithecines and yet this species upper facial breadth (bifmt) is similar in size to Colobus and Piliocolobus. These results require further investigation with larger samples sizes and further intra-group partitioning.

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Table 5.19: Results of body weight (Ln (g)) plotted against the generic mean for particular cranial measurements (Ln (mm)) (cranial variables listed in alphabetical order by abbreviation; those in bold represent cranial variables with an adjusted R squared value >0.70 and a p-value <0.05).

Adjusted P-value/ Mean Generic Body Weight (g) Ln (mm) R squared Significance F (Napier, 1985; Rowe, 1996; bas-br 0.59 0.006 Fleagle, 1998; Delson et al, 2000): bas-nas 0.65 0.003 bas-pros 0.58 0.006 Procolobus verus - 4450 biast 0.62 0.004 Piliocolobus spp. - 8285 biaur 0.82 0.0002 Colobus spp. - 9496 bicanex 0.43 0.02 Presbytis spp. - 6277 biecm 0.81 0.0002 Trachypithecus spp. - 8159 bien 0.64 0.003 Semnopithecus spp. - 11445 bienm 0.53 0.01 bifmo 0.55 0.008 Simias concolor - 7975 bifmt 0.52 0.01 Pygathrix spp. - 9720 bipor 0.68 0.002 Rhinopithecus spp. - 12847 bizi 0.78 0.0004 Nasalis larvatus - 15110 bizs 0.01 0.32 bizygo 0.65 0.003 br-lam -0.02 0.40 g-o 0.84 0.0001 lam-opn 0.51 0.01 nas-br 0.77 0.0005 nas-pros 0.36 0.04 nas-rhi 0.03 0.3 ol-sta 0.51 0.01 ol-pms 0.47 0.02 palhei -0.05 0.49 pros-o 0.76 0.0006 zs-zgyi 0.28 0.07 zs-zi 0.49 0.01

4.55 Figure 5.27: Adjusted R2: 0.82 Se. R. 4.50 Significance F & P-value 0.0001 N. 4.45 Py. y = 0.1859x + 2.694 Co. 4.40 Pi. 4.35 Pre. Tr. 4.30 Si Pro .

Ln Length Cranial (g-o) . 4.25

4.20 8.20 8.40 8.60 8.80 9.00 9.20 9.40 9.60 9.80 Ln Body Weight (g)

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5.5 Multivariate Statistics and Morphological Distances: To consider and evaluate the entire multivariate datasets simultaneously PCA and CVA were utilized. 5.5.1 PCA results for Ln transformed data: To identify the underlying structure (i.e. the cranial variables which account for most of the variation between genera) of multivariate datasets derived from Ln transformed generic means, PCA was utilized. By doing so, PCA based on the variance-covariance matrix results in the first (Eigenvalue – 0.37; 60.98%), second (Eigenvalue - 0.12; 20.12%) and third (Eigenvalue - 0.06; 9.14%) PCs accounting for 90.24% of variation within the subfamily (Figure 5.28). One hundred percent of the variation is explained by nine PCs. For PC 1, Semnopithecus receives the largest PCA object score, 20.12, closely followed by Nasalis at 20.08 (the object scores for PC 2 are negative and PC 3 are positive). In contrast, the smallest PCA object score is attained by Procolobus, 18.50, succeeded by Presbytis at 18.51. The variable loadings for the first PC are all positive and may be interpreted as differences due to size; the remaining PCs have mixed values and variable loadings are low to moderate (Table 5.20). The first PC was dominated by palatal and naso-facial lengths. The largest positively loaded dimension was palatal length A (ol-sta) with 0.27, closely followed by the other two palatal length measurements (iv-pms and ol-pms) each receiving 0.26. These measurements are closely followed by naso-facial dimensions including superior facial height (nas-pros), nasal height (nas-ns), sagittal height of the nasal aperture (rhi-ns), sagittal length of the nasal bones (nas-rhi), maxillo-alveolar length (pros-dm3) and superior facial length (bas-pros) with positive variable loadings of 0.24, 0.23, 0.23, 0.23, 0.22 and 0.20, respectively. The second PC was again governed by nasal dimensions including sagittal length of the nasal bones (nas-rhi), nasal height (nas-ns) and superior facial height (nas-pros) which procure positive variable loadings of 0.59, 0.26 and 0.21, correspondingly. Contrasting with these lengths were negatively loaded measurements such as bizygomaxillare superior (bizs) and the parietal sagittal chord (br-lam) with variable loadings of -0.44 and -0.24, respectively.

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Figure 5.28: Colobinae scatterplot of PC 1 & 2 and PC 2 & 3 for Ln transformed generic means. Co. - Colobus; Pi. - Piliocolobus; Pro. - Procolobus; Se. - Semnopithecus; Tr. - Trachypithecus; Pre. - Presbytis; Py. - Pygathrix; R. - Rhinopithecus; N. - Nasalis; and Si. - Simias.

-4 Nasal & -4.1 Facial -4.2 lengths -4.3 -4.4 % of -4.5 Si. -4.6 PC Eigenvalue Var. Cum. %. -4.7 N. 1 0.37 60.98 60.98 -4.8 Increase in body size 2 0.12 20.12 81.10 PC 2: -4.9 3 0.06 9.14 90.24 -5 Pi. 20.12% Pro. -5.1 4 0.02 3.63 93.87 -5.2 Co. Se. 5 0.02 2.58 96.45 Tr. -5.3 Pre. -5.4 Py. -5.5 -5.6 PCA -5.7 R. scores Axis 1 Axis 2 Axis 3 -5.8 Palatal & Nasal Co. 19.90 -5.15 1.20 Bizygomaxillare- 5.9 lengths Pi. 19.38 -4.98 1.47 Superior & Parietal Sagittal 18 . 4 18 . 5 18 . 6 18 . 7 18 . 8 18 . 9 19 19 . 1 19 . 2 19 . 3 19 . 4 19 . 5 19 . 6 19 . 7 19 . 8 19 . 9 20 20.120.2 Pro. 18.50 -5.02 1.44 Chord PC 1: 60.98% Se. 20.12 -5.15 1.54 Tr. 18.92 -5.23 1.48 Pre. 18.51 -5.33 1.66 2 Py. 19.47 -5.39 1.75 Length of R. R. 19.61 -5.69 1.96 Zygomatico- 1. 9 maxillary suture Si. N. 20.08 -4.67 1.79 1. 8 Si. 18.95 -4.49 1.89 N. Mean 19.35 -5.11 1.62 Py. 1. 7 Max. 20.12 -5.69 1.96 Pre. Min. 18.50 -4.49 1.20 1. 6 Range 1.62 1.20 0.76 PC 3: Se. SD 0.61 0.35 0.23 9.14% 1. 5 Tr. Pi. CV 3.13 6.81 14.51 Pro. 1. 4

1. 3

Bizygomaxillare 1. 2 Nasal & Superior & Co. Facial 1. 1 Parietal Sagittal lengths Chord Inferior breadth of the nasal bones & -5.9 -5.8 -5.7 -5.6 -5.5 -5.4 -5.3 -5.2 -5.1 -5 -4.9 -4.8 -4.7 -4.6 -4.5 -4.4 -4.3 -4.2 -4.1 -4 Palatal height PC 2: 20.12%

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Table 5.20: Variable loadings for Colobinae Ln transformed generic means. PC loadings PC 1 PC 2 PC 3 lam-opn 0.07 bizs -0.44 inbrnabo -0.53 inbrnabo 0.08 br-lam -0.24 palhei -0.31 bas-br 0.10 bifmo -0.18 br-lam -0.23 br-lam 0.10 bien -0.17 ol-sta -0.18 bizs 0.10 bifmt -0.17 bizs -0.18 bifmt 0.10 biast -0.12 ol-pms -0.16 bifmo 0.10 bipor -0.11 bicanex -0.13 biast 0.11 bas-br -0.11 iv-pms -0.07 nas-br 0.11 nas-br -0.11 bas-pros -0.07 g-o 0.12 biaur -0.11 bicanin -0.05 zs-zygi 0.12 bizi -0.10 maxalvlen -0.05 biaur 0.12 lam-opn -0.10 rhi-ns -0.03 bas-nas 0.13 g-o -0.08 bas-nas -0.03 palhei 0.13 bizygo -0.08 i1i2 -0.03 bipor 0.13 i1i2 -0.08 zs-zygi -0.02 bizygo 0.14 maxnawi -0.07 bizygo 0.00 bienm 0.15 biecm -0.06 bienm 0.00 bizi 0.15 bicanex -0.05 nas-pros 0.00 bicanin 0.16 bas-nas -0.05 bas-br 0.01 zs-zi 0.16 ol-sta -0.05 bipor 0.02 biecm 0.16 rhi-ns -0.04 biseptal 0.03 i1i2 0.16 bienm -0.04 bien 0.03 bien 0.17 biseptal -0.04 pros-o 0.03 biseptal 0.17 maxalvlen -0.03 nas-ns 0.03 pros-o 0.17 bicanin -0.02 bizi 0.06 bicanex 0.17 pros-o -0.02 biecm 0.08 maxnawi 0.19 bas-pros 0.01 g-o 0.09 bas-pros 0.20 zs-zygi 0.04 biaur 0.09 maxalvlen 0.22 ol-pms 0.05 nas-rhi 0.13 nas-rhi 0.23 iv-pms 0.05 lam-opn 0.14 rhi-ns 0.23 inbrnabo 0.12 bifmt 0.15 nas-ns 0.23 palhei 0.16 bifmo 0.18 nas-pros 0.24 zs-zi 0.17 biast 0.20 ol-pms 0.26 nas-pros 0.21 nas-br 0.24 iv-pms 0.26 nas-ns 0.26 maxnawi 0.29 ol-sta 0.27 nas-rhi 0.59 zs-zi 0.38

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The largest positively loaded measurements for the third PC include the zygomatico-maxillary suture length (zs-zi), maximum width of the nasal aperture (maxnawi), frontal sagittal chord (nas-br) and biasterionic breadth (biast) which attained variable loadings of 0.38, 0.29, 0.24 and 0.20, respectively. Juxtaposed against these dimensions were negatively loaded measurements such as the inferior breadth of the nasal bones (inbrnabo), palatal height (palhei) and the parietal sagittal chord (br-lam) with variable loadings of -0.53, -0.31 and -0.23, respectively. The scatterplots for PCs 1 & 2 and PCs 2 & 3 were both very informative. The scatterplot for PCs 1 & 2 was the result of size differences (Figure 5.28). From left to right along the first PC there is an increase in body size; on the left side there are Procolobus and Presbytis, while on the right side there are Semnopithecus and Nasalis. In contrast, second PC involves differences to facial form. Those below -5.0 have a relatively narrow upper face (e.g. Simias and Nasalis), whereas those greater than -5.0 had a relatively wide upper face (e.g. Pygathrix and Rhinopithecus). The scatterplot for PCs 2 & 3 was very different from the scatterplot for PCs 1 & 2 and also appears to convey some phylogenetic content, highlighting shape divergences. In the center were colobines which typify the cranial shape of many colobine monkeys (Semnopithecus and Piliocolobus); i.e. wide upper face but relatively short face and nasal bones with moderate facial projection (Delson, 1975). Then moving outward and away from the central genera were the more cranially derived genera such as Nasalis, Rhinopithecus and Colobus (Groves, 1970; Hull, 1979; Pan & Oxnard, 2001). Table 5.21 provides the inter-generic Euclidean distance matrix produced from PCA object scores per genus from the first to fifth principal components based on Ln transformed generic means. The average distance between colobine genera was 1.0 (Range 1.3). The largest distance produced was between Nasalis and Presbytis at 1.7 while the smallest distance was between Presbytis and Procolobus with 0.40. For comparison, Table 5.22 provides intra-generic Euclidean distances based on Ln transformed data. The largest mean intra-generic Euclidean distance was produced by Semnopithecus with 1.0 (followed by Nasalis with 0.91). This is due to the extreme size differences between Northern langur species, at the foothills of the Himalayas, compared to much smaller species in the South of

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India. The smallest mean intra-generic Euclidean distance was produced by Procolobus with 0.63 (succeeded by Presbytis with 0.67). In summary, the Nasalina genera are highly distinct compared to the other two subtribes, although Colobus is just as distinct from Procolobus and Piliocolobus (Figure 5.28). Table 5.21: Inter-generic Euclidean distances based on PCA scores from PC 1-5 per genus from Ln transformed generic means. Co. 0 Mean 1.02 Pi. 0.64 0 Range 1.32 Pro. 1.45 0.88 0 Se. 0.63 0.84 1.68 0 Tr. 1.06 0.59 0.59 1.23 0

Pre. 1.50 0.96 0.40 1.65 0.53 0 Py. 0.74 0.58 1.13 0.90 0.71 1.02 0 R. 0.99 0.89 1.40 0.92 0.99 1.20 0.47 0 N. 0.82 0.84 1.66 0.69 1.38 1.71 1.00 1.14 0 Si. 1.36 0.81 0.89 1.43 0.85 1.01 1.07 1.39 1.2 0 Co. Pi. Pro. Se. Tr. Pre. Py. R. N. Si. Table 5.22: Intra-generic Euclidean distances based on Ln transformed data.

Mean Max Min Range Co. 0.77 1.48 0.35 1.13 Pi. 0.83 1.67 0.36 1.30

Pro. 0.63 0.96 0.49 0.46 Se. 1.01 1.91 0.33 1.58 Tr. 0.85 1.68 0.39 1.29 Pre. 0.67 1.17 0.29 0.88 Py. 0.83 1.86 0.35 1.51 R. 0.88 1.77 0.22 1.55 N. 0.91 1.90 0.35 1.55 Si. 0.78 1.12 0.45 0.68 5.5.2 PCA results for MSV: PCA based on the variance-covariance matrix resulted in the first (Eigenvalue - 0.15; 56.34%) and second (Eigenvalue - 0.05; 17.93%) PCs accounting for 83.80% of variation within the subfamily (6.83% less variation explained than PCs 1 & 2 for Ln transformed generic means but 45.01% more variation explained than PCs 2 & 3 for Ln transformed generic means; Figure 5.29). One hundred percent of the variation is explained by nine PCs. The variable loadings for the first PC were not all positive (Tables 5.23), which again varied from low to moderate, and cannot be interpreted as size differences but

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Figure 5.29: Colobinae scatterplot of PC 1 & 2 of generic mean MSV. Co. - Colobus; Pi. - Piliocolobus; Pro. - Procolobus; Se. - Semnopithecus; Tr. - Trachypithecus; Pre. - Presbytis; Py. - Pygathrix; R. - Rhinopithecus; N. - Nasalis; and Si. - Simias.

Frontal Sagittal 1. 1 Chord & Length Table 5.23: Variable loadings for Si. Colobinae generic mean MSV. of nasal bones 1 PC loadings 0.9 N. PC 1 PC 2 0.8 nas-pros -0.28 bizs -0.46 nas-ns -0.28 ol-sta -0.32 Pre. 0.7 R. PC 2: nas-rhi -0.22 br-lam -0.31 17.93% Pro. bas-pros -0.17 bas-pros -0.23 0.6 Se. Py. Pi. Tr. ol-pms -0.14 ol-pms -0.15 0.5 ol-sta -0.13 maxalvlen -0.13 iv-pms -0.09 bicanex -0.13 0.4 maxalvlen -0.08 iv-pms -0.07

0.3 zs-zi -0.05 rhi-ns -0.06 pros-o -0.04 bas-nas -0.06 Nasal lengths Upper facial 0.2 Co. rhi-ns -0.03 inbrnabo -0.06 breadths Bizygomaxillare palhei -0.03 bien -0.06 Superior & Palatal 2.1 2.2 2.3 2.4 2.5 2.6 2.7 2.8 2.9 3 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 3.9 4 4.1 zs-zygi -0.03 palhei -0.03 length PC 1: 56.34% inbrnabo -0.02 bicanin -0.02 bicanex -0.01 i1i2 -0.02 % of bicanin 0.00 bizygo -0.01 PC Eigenvalue Var. Cum. %. biseptal 0.00 bipor -0.01 1 0.15 56.34 56.34 i1i2 0.01 bas-br 0.00 2 0.05 17.93 74.27 maxnawi 0.01 bienm 0.01 3 0.02 9.53 83.80 bienm 0.01 biseptal 0.01 4 0.02 6.64 90.45 biecm 0.04 maxnawi 0.04 5 0.01 3.34 93.78 bas-nas 0.07 biecm 0.04

PCA bizi 0.13 bizi 0.06 scores Axis 1 Axis 2 bizygo 0.13 pros-o 0.06 Co. 3.07 0.21 lam-opn 0.13 zs-zygi 0.09 Pi. 3.28 0.58 bien 0.14 bifmt 0.11 Pro. 3.60 0.64 bipor 0.17 biaur 0.11 Se. 3.32 0.60 br-lam 0.19 bifmo 0.12 Tr. 3.65 0.56 bas-br 0.19 lam-opn 0.13 Pre. 4.01 0.74 biast 0.20 nas-pros 0.13 Py. 3.77 0.60 biaur 0.21 biast 0.17 R. 4.00 0.72 g-o 0.22 g-o 0.18 N. 2.95 0.86 nas-br 0.24 nas-ns 0.21 Si. 3.12 1.04 bifmt 0.33 zs-zi 0.27 Mean 3.48 0.66 bifmo 0.34 nas-rhi 0.28 Max. 4.01 1.04 bizs 0.34 nas-br 0.33 Min. 2.95 0.21 Range 1.06 0.83 SD 0.38 0.22 CV 11.00 32.93

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Table 5.24: Inter-generic Euclidean distances based on PCA scores from PC 1-5 per genus from generic mean MSV.

Co. 0 Mean 0.67 Pi. 0.49 0 Range 0.82 Pro. 0.79 0.37 0 Se. 0.61 0.37 0.54 0 Tr. 0.71 0.45 0.37 0.40 0 Pre. 1.10 0.77 0.50 0.79 0.43 0 Py. 0.83 0.63 0.56 0.75 0.44 0.48 0 R. 1.09 0.80 0.63 0.78 0.51 0.42 0.46 0 N. 0.67 0.45 0.75 0.60 0.80 1.09 0.90 1.09 0 Si. 0.83 0.55 0.74 0.61 0.74 0.96 0.82 0.98 0.29 0 Co. Pi. Pro. Se. Tr. Pre. Py. R. N. Si.

Table 5.25: Intra-generic Euclidean distances based on MSV.

Mean Max Min Range Co. 0.58 0.95 0.25 0.70 Pi. 0.63 1.03 0.33 0.70 Pro. 0.50 0.56 0.45 0.10 Se. 0.58 0.90 0.24 0.67 Tr. 0.63 1.05 0.33 0.72 Pre. 0.57 0.84 0.28 0.56 Py. 0.55 0.98 0.29 0.68 R. 0.56 1.05 0.20 0.85 N. 0.58 1.00 0.29 0.72 Si. 0.54 0.72 0.33 0.39

instead shape dissimilarities or divergences; the second PC also had variable loadings of mixed polarity. The MSV that receive the largest variable loadings were different in comparison to those from Ln transformed data. Facial MSV such as bizygomaxillare superior (bizs), bifrontomalarorbitale (bifmo) and superior facial breadths (bifmt) have positive variable loadings of 0.34, 0.34 and 0.33, respectively. These MSV are then followed by neurocranial MSV including the frontal sagittal chord (nas-br), cranial vault length (g-o), biauriculare (biaur) and biasterionic breadths (biaur and biast) with variable loadings of 0.24, 0.22, 0.21, and 0.20, correspondingly. The largest positively loaded MSV for the second PC is the frontal sagittal chord (nas-br), sagittal length of the nasal bones (nas-rhi), zygomatico-maxillary suture length

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(zs-zi) and nasal height (nas-ns) with variable loadings of 0.33, 0.28, 0.27 and 0.21, respectively. Working against these MSV are negatively loaded dimensions such as the bizygomaxillare superior breadth (bizs), palatal length A (ol-sta), parietal sagittal chord (br- lam) and superior facial length (bas-pros) with variable loadings of -0.46, -0.32, -0.31 and - 0.23, correspondingly. The scatterplot for PCs 1 & 2 based on the Colobinae generic mean is quite informative in revealing shape similarities and dissimilarities between colobine genera, as well as agreeing with previous work (e.g. Groves, 1970; Hull, 1979; Figure 5.29); and similar to Ln transformed data PCs 2 & 3. Three of the most cranially distinct genera (Nasalis, Simias and Colobus) are separated from a cluster of the more typical colobine morphotype. Table 5.24 provides the inter-generic Euclidean distance matrix produced from PCA object scores from the first to fifth principal components based on generic mean MSV. The average distance between colobine genera was 0.67 (Range 0.82). The largest distance was between Presbytis and Colobus at 1.10 while the smallest distance was between Nasalis and Simias with 0.37. For comparison, Table 5.25 provides intra-generic Euclidean distances based on MSV. The largest mean intra-generic Euclidean distance was produced by Piliocolobus and Trachypithecus each at 0.63 (followed by Colobus, Semnopithecus and Nasalis, all three producing 0.58). The smallest mean intra-generic Euclidean distance was again produced by Procolobus at 0.50 (succeeded by Simias, 0.54). In summary, the multivariate results for MSV of the Colobinae reinforce the results for Ln transformed data (Figure 5.28, PCs 2 & 3). Nasalis, Simias and Colobus were located well away from the other colobines, highlighting their peculiar cranial shapes (Figure 5.29).

5.5.4 CVA results for Ln transformed data: To consider and summarize the entire Colobinae (n=186) multivariate dataset CVA was employed. After subjecting the Colobinae Ln transformed pooled sex multivariate dataset to MANOVA and CVA the first (Eigenvalue - 10.64; 30.24%) and second (Eigenvalue - 8.57; 24.34%) canonical variate axes were found to account for 54.58% of the variation (Figure 5.30). Table 5.28 provides the CVA object scores per genus for the

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Figure 5.30: Colobinae scatterplot of canonical variate axes 1 & 2 for Ln transformed pooled sex data with 95% confidence ellipses. Colobus - dot ; Piliocolobus - cross +; Procolobus - open square ; Semnopithecus – x; Trachypithecus - circle ; Presbytis - diamond ; Pygathrix - triangle ; Rhinopithecus - dash -; Nasalis - vertical line |; Simias - filled square .

Upper facial breadths 2 CVA Eigen- Eigen- vectors Axis 1 vectors Axis 2 ol-sta -0.37 nas-ns -0.36 1 inbrnabo -0.28 nas-rhi -0.32 CV ol-pms -0.19 ol-pms -0.27 Axis 2: palhei -0.18 nas-pros -0.22 24.34% zs-zygi -0.16 iv-pms -0.22 0 bizs -0.15 bas-pros -0.17 Palatal length A bicanex -0.14 inbrnabo -0.17 & Inferior br-lam -0.14 maxalvlen -0.16 breadth of the Frontal sagittal nasal bones chord & bas-nas -0.12 pros-o -0.16 Biorbital breadth bipor -0.07 bicanin -0.12 Nasal lengths 4.1 4.2 4.3 4.4 4.5 4.6 4.7 4.8 4.9 5 5.1 5.2 5.3 5.4 5.5 5.6 5.7 5.8 5.9 6 bicanin -0.05 g-o -0.08 CV Axis 1: 30.24% bas-pros -0.02 palhei -0.06 MANOVA bas-br 0.01 zs-zi -0.05 Wilks's Pillai bienm 0.02 bas-br -0.03 lambda: 0.00 trace: 6.07 i1i2 0.02 bicanex 0.01 df1: 324.00 df1: 324.00 rhi-ns 0.03 zs-zygi 0.01 df2: 1343.00 df2: 1431.00 biaur 0.04 bizygo 0.01 F: 12.54 F: 9.17 bizi 0.06 biseptal 0.02 p(same): 0.00 p(same): 0.00 bizygo 0.07 bizi 0.03 CVA iv-pms 0.07 nas-br 0.03 Eigenvalue 1: 10.64 Percent: 30.24 maxalvlen 0.07 bienm 0.04 Eigenvalue 2: 8.57 Percent: 24.34 bien 0.07 maxnawi 0.05 Total %: 54.58 nas-ns 0.08 rhi-ns 0.05 lam-opn 0.08 i1i2 0.05 Table 5.27: Inter-generic Euclidean distances based on mean CVA scores from axes 1-5 per nas-pros 0.10 bipor 0.05 genus from Ln transformed data. biecm 0.12 bas-nas 0.06 biseptal 0.14 biaur 0.08 Co. 0 Mean 0.77 bifmt 0.15 biecm 0.09 Pi. 0.51 0 Range 0.93 biast 0.16 biast 0.11 Pro. 1.08 0.64 0 g-o 0.17 lam-opn 0.12 Se. 0.48 0.56 1.11 0 zs-zi 0.20 bien 0.13 Tr. 0.79 0.48 0.46 0.74 0 pros-o 0.20 ol-sta 0.16 Pre. 1.15 0.76 0.35 1.12 0.45 0 nas-rhi 0.21 br-lam 0.20 Py. 0.68 0.57 0.89 0.62 0.58 0.76 0 maxnawi 0.27 bifmt 0.23 R. 0.86 0.78 1.11 0.68 0.83 0.95 0.43 0 bifmo 0.30 bizs 0.35 N. 0.66 0.63 1.21 0.59 1.0 1.27 0.78 0.92 0 nas-br 0.40 bifmo 0.38 Si. 0.94 0.52 0.66 0.89 0.66 0.77 0.79 1.05 0.72 0 Co. Pi. Pro. Se. Tr. Pre. Py. R. N. Si.

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Table 5.28: Colobus spp. (n=26; 15, 11). Semnopithecus spp. (n=9; 4, 5). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean 4.71 0.33 Mean 5.01 0.39 Max. 4.84 0.56 Max. 5.24 0.57 Min. 4.53 0.22 Min. 4.77 0.03 Range 0.30 0.34 Range 0.47 0.54 SD 0.08 0.09 SD 0.16 0.16 CV 1.62 27.66 CV 3.22 41.41 Piliocolobus spp. (n=28; 15, 13). Trachypithecus spp. (n=33; 15, 18). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean 4.84 0.34 Mean 4.79 0.62 Max. 5.07 0.57 Max. 4.95 0.98 Min. 4.65 0.16 Min. 4.68 0.44 Range 0.41 0.42 Range 0.26 0.54 SD 0.09 0.13 SD 0.07 0.12 CV 1.82 36.75 CV 1.45 19.74 Procolobus verus (n=5; 3, 2). Presbytis spp. (n=18; 8, 10). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean 4.69 0.56 Mean 4.88 0.83 Max. 4.81 0.68 Max. 5.02 0.96 Min. 4.60 0.43 Min. 4.72 0.70 Range 0.21 0.25 Range 0.30 0.26 SD 0.08 0.09 SD 0.09 0.07 CV 1.75 15.69 CV 1.80 8.25 Rhinopithecus spp. (n=17; 5, 12). Nasalis larvatus (n=28; 18, 10). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean 5.25 0.93 Mean 5.27 0.02 Max. 5.39 1.29 Max. 5.43 0.32 Min. 5.09 0.69 Min. 5.14 -0.23 Range 0.30 0.60 Range 0.29 0.55 SD 0.08 0.16 SD 0.07 0.16 CV 1.48 16.96 CV 1.28 977.55 Pygathrix spp. (n=25; 14, 11). Simias concolor (n=7; 2, 5). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean 5.14 0.73 Mean 5.13 0.13 Max. 5.29 0.99 Max. 5.24 0.26 Min. 4.99 0.51 Min. 5.05 -0.04 Range 0.30 0.48 Range 0.19 0.29 SD 0.07 0.12 SD 0.06 0.11 CV 1.27 16.67 CV 1.23 87.77

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first and second CV axes. On average, Nasalis scored the largest CVA score for CV axis 1 with 5.27 (Min. - Max., 5.14 - 5.43; closely followed by Rhinopithecus at 5.25), while Procolobus received the least at 4.69 (Min.-Max., 4.60 - 4.81; succeeded by Colobus at 4.71). Both CV axes 1 and 2 had mixed values with low to moderate variable loadings. Cranial measurements that were positively correlated with the first CV axis included the frontal sagittal chord (nas-br), bifrontomalarorbitale breadth (bifmo), maximum width of the nasal aperture (maxnawi), sagittal length of the nasal bones (nas- rhi), maximum cranial length (pros-o) and zygomatico-maxillary suture length (zs-zi) with variable loadings of 0.40, 0.30, 0.27, 0.21, 0.20 and 0.20, respectively (Table 5.26). However, juxtaposed against these cranial dimensions that were negatively correlated to first CV axis included, palatal length A (ol-sta) and the inferior breadth of the nasal bones with variable loadings of -0.37 and -0.28, respectively. The largest positively correlated MSV with the second CV axis was bifrontomalarorbitale breadth (bifmo) at 0.38; followed by bizygomaxillare superior breadth (bizs), bifrontomalartemporale breadth (bifmt) and the parietal sagittal chord (br- lam) scoring 0.35, 0.23 and 0.20, respectively. Contrasting with these MSV were negatively loaded nasal MSV such as nasal height (nas-ns), sagittal length of the nasal bones (nas-rhi), palatal length B (ol-pms), superior facial height (nas-pros) and the distance between incisivion and the central junction of the palatomaxillary suture (iv-pms) with variable loadings of -0.36, -0.32, -0.27, -0.22 and -0.22, correspondingly. The scatterplot for CV axes 1 & 2 revealed considerable overlap of colobine genera (Figure 5.30). However, similar to the scatterplots for PCA, genera such as Rhinopithecus (and Pygathrix) and Nasalis (and Simias) were found to lie on the outskirts of the more overlapping genera. The large 95% confidence ellipses for Procolobus and Semnopithecus were the result of the small sample sizes (n=5 and n=9). Please note, the overlap between colobine genera (for both Ln transformed data and MSV) was significantly reduced when only the subtribes are subjected to multivariate statistics. However, for the sake of brevity, those results will be presented elsewhere. Table 5.27 provides the inter-generic Euclidean distances distance matrix produced from generic mean CVA object scores from the first to fifth canonical variate axes based on Ln transformed pooled sex data. The average Euclidean distance between colobine genera

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is 0.77 (Range 0.93). The largest distance was between Nasalis and Presbytis at 1.27 (followed by Nasalis and Procolobus with 1.21) and the smallest distance was between Presbytis and Procolobus with 0.35 (succeeded by Rhinopithecus and Pygathrix with 0.43). In summary, CVA of the entire Colobinae multivariate datasets revealed the extent to which colobine crania are similar in size, as seen in the scatterplot of CV axes 1 & 2 (Figure 5.31).

5.5.5 CVA results for MSV: After converting the raw data of the Colobinae (n=186) into MSV and subjecting the entire multivariate dataset to MANOVA and CVA, the first (Eigenvalue - 10.01; 31.15%) and second (Eigenvalue - 8.40; 26.15%) canonical variate axes were found to account for 57.30% of the variation (Figure 5.31; 2.72% more variation explained than Ln transformed pooled sex data). Table 5.31 provides the CVA object scores per genus for the first (negative) and second (positive) CV axes. On average, Simias scored the largest CVA object scores for CV axis 1 at -1.10 (Min. - Max., -1.17 - -1.03; closely followed by Nasalis with -1.08), while Colobus received the least at -0.50 (Min. - Max., -0.64 - -0.38). The three largest positively loaded MSV for the first CV axis were palatal length A (ol-sta), bizygomaxillare superior breadth (bizs) and the parietal sagittal chord (br-lam; Table 5.29) with 0.45, 0.29 and 0.24, respectively. Contrasting with these MSV were negatively correlated MSV such as the sagittal length of the nasal bones (nas-rhi), frontal sagittal chord (nas-br), maximum width of the nasal aperture (maxnawi), nasal height (nas- ns), zygomatico-maxillary suture length (zs-zi), maximum cranial length (pros-o) and superior facial height (nas-pros) with variable loadings of -0.43, -0.27, -0.25, -0.23, -0.22, - 0.22 and -0.20, respectively. The MSV that are positively correlated with the second CV axis included bifrontomalarorbitale, bifrontomalartemporale and bizygomaxillare superior breadths (bifmo, bifmt and bizs), which along with the frontal sagittal chord (nas-br) procured variable loadings of 0.49, 0.35, 0.27 and 0.20, correspondingly. Juxtaposed against these MSV were dimensions which were negatively correlated with the second axis such as the inferior breadth of the nasal bones (inbrnabo), palatal length B (ol-pms) and nasal height (nas-ns) with variable loadings of -0.35, -0.27 and -0.24, respectively.

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Figure 5.31: Colobinae scatterplot of canonical variate axes 1 & 2 for pooled sex MSV with 95% confidence ellipses. Colobus - dot ; Piliocolobus - cross +; Procolobus - open square ; Semnopithecus - x; Trachypithecus - circle ; Presbytis - diamond ; Pygathrix - triangle ; Rhinopithecus - dash -; Nasalis - vertical line |; Simias - filled square .

3 Upper 2.9 Facial 2.8 Table 5.29: Variable loadings for breadths 2.7 Colobinae pooled sex MSV. 2.6 2.5 CVA 2.4 2.3 Eigen- Eigen- 2.2 vectors Axis 1 vectors Axis 2 CV 2.1 nas-rhi -0.43 inbrnabo -0.35 Axis 2: 2 26.15% 1. 9 nas-br -0.27 ol-pms -0.27 1. 8 maxnawi -0.25 nas-ns -0.24 1. 7 1. 6 nas-ns -0.23 bas-pros -0.17 1. 5 zs-zi -0.22 iv-pms -0.17 1. 4 pros-o -0.22 maxalvlen -0.12 1. 3 Palatal length & 1. 2 Nasal bone Bizygomaxillare nas-pros -0.20 nas-rhi -0.12 1. 1 length Superior biseptal -0.13 bicanin -0.09 Inferior breadth g-o -0.12 nas-pros -0.08 of the nasal -1.2 -1.1 -1 -0.9 -0.8 -0.7 -0.6 -0.5 -0.4 -0.3 -0.2 - iv-pms -0.09 palhei -0.06 bones & Palatal CV Axis 1: 31.15% length B biast -0.08 bas-br -0.04 bicanin -0.07 ol-sta -0.04 MANOVA bifmt -0.05 pros-o -0.04 Wilks's Pillai lambda: 0.00 trace: 5.80 maxalvlen -0.05 bas-nas -0.02 df1: 324.00 df1: 324.00 i1i2 -0.03 zs-zygi 0.01 df2: 1343.00 df2: 1431.00 bizygo -0.02 g-o 0.03 F: 11.14 F: 8.02 bizi -0.01 bizygo 0.04 p(same): 0.00 p(same): 0.00 bifmo -0.01 bizi 0.04 CVA bas-pros 0.01 rhi-ns 0.06 Eigenvalue 1: 10.01 Percent: 31.15 biecm 0.02 bienm 0.06 Eigenvalue 2: 8.40 Percent: 26.15 biaur 0.03 bipor 0.06 Total % 57.30 rhi-ns 0.03 bien 0.07 bienm 0.03 bicanex 0.07 bas-br 0.05 br-lam 0.09 Table 5.30: Inter-generic Euclidean distances based on mean CVA scores from axes 1-5 per genus from MSV. palhei 0.05 biecm 0.09 bipor 0.06 i1i2 0.09 Co. 0 Mean 0.57 lam-opn 0.06 zs-zi 0.09 Pi. 0.45 0 Range 0.74 ol-pms 0.07 biseptal 0.09 Pro. 0.73 0.33 0 zs-zygi 0.08 lam-opn 0.11 Se. 0.57 0.32 0.45 0 bien 0.08 biaur 0.12 Tr. 0.64 0.38 0.33 0.29 0 bicanex 0.08 maxnawi 0.19 Pre. 0.96 0.64 0.37 0.65 0.40 0 inbrnabo 0.16 biast 0.19 Py. 0.74 0.55 0.49 0.62 0.39 0.40 0 bas-nas 0.17 nas-br 0.20 R. 0.94 0.66 0.51 0.62 0.46 0.34 0.42 0 br-lam 0.24 bizs 0.27 N. 0.62 0.44 0.69 0.55 0.68 0.94 0.79 0.94 0 bizs 0.29 bifmt 0.35 Si. 0.71 0.42 0.58 0.47 0.56 0.78 0.67 0.78 0.22 0 ol-sta 0.45 bifmo 0.49

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Table 5.31: Colobus spp. (n=26; 15, 11). Semnopithecus spp. (n=9; 4, 5). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean -0.50 1.82 Mean -0.68 2.03 Max. -0.64 2.11 Max. -0.85 2.24 Min. -0.38 1.68 Min. -0.56 1.78 Range 0.25 0.43 Range 0.30 0.46 SD 0.06 0.10 SD 0.09 0.15 CV 13.07 5.55 CV 13.16 7.25 Piliocolobus spp. (n=28; 15, 13). Trachypithecus spp. (n=33; 15, 18). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean -0.71 2.00 Mean -0.62 2.24 Max. -0.85 2.23 Max. -0.77 2.56 Min. -0.55 1.83 Min. -0.45 2.01 Range 0.29 0.40 Range 0.33 0.55 SD 0.07 0.12 SD 0.06 0.12 CV 10.30 6.05 CV 9.51 5.26 Procolobus verus (n=5; 3, 2). Presbytis spp. (n=18; 8, 10). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean -0.67 2.23 Mean -0.65 2.56 Max. -0.79 2.30 Max. -0.73 2.73 Min. -0.59 2.10 Min. -0.48 2.42 Range 0.19 0.20 Range 0.25 0.31 SD 0.08 0.09 SD 0.07 0.08 CV 11.68 3.99 CV 10.54 3.28 Rhinopithecus spp. (n=17; 5, 12). Nasalis larvatus (n=28; 18, 10). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean -0.69 2.63 Mean -1.08 1.81 Max. -0.80 2.85 Max. -1.21 2.14 Min. -0.58 2.47 Min. -0.95 1.56 Range 0.22 0.38 Range 0.26 0.58 SD 0.06 0.11 SD 0.07 0.17 CV 9.04 4.05 CV 6.31 9.14 Pygathrix spp. (n=25; 14, 11). Simias concolor (n=7; 2, 5). CVA CVA scores Axis 1 Axis 2 scores Axis 1 Axis 2 Mean -0.71 2.48 Mean -1.10 1.99 Max. -0.85 2.64 Max. -1.17 2.12 Min. -0.61 2.18 Min. -1.03 1.76 Range 0.23 0.46 Range 0.14 0.36 SD 0.06 0.12 SD 0.04 0.15 CV 8.30 4.74 CV 3.75 7.60

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The scatterplot for CV axes 1 & 2 is very similar to the scatterplot for Ln transformed data (Figure 5.31). Again, there was considerable overlap and the more cranially distinct genera (Nasalis, Simias, Pygathrix, Rhinopithecus and Colobus) were found to lie to the periphery. Table 5.30 provides the inter-generic Euclidean distances distance matrix produced from generic mean CVA object scores from the first to fifth canonical variate axes based on pooled sex MSV. The average Euclidean distance between colobine genera was 0.57 (Range 0.74). The largest distance was between Colobus and Presbytis at 0.96 while the smallest distance was between Nasalis and Simias with 0.22. For comparison, Table 5.32 provides Cherry et al’s (1978) M distance between colobine genera. The average M distance between genera was 1.27 (Range 1.33). The largest morphological divergence was achieved by Presbytis and Colobus with 2.03 (followed by Nasalis and Presbytis at 1.93); the smallest were between, Semnopithecus and Piliocolobus with 0.70 (succeeded by Piliocolobus and Procolobus at 0.74). In summary, by exploring shape instead of size, CVA produced similar results as Ln transformed data, just orientated differently. CVA of colobine MSV appears to have slightly better separated Nasalina and Colobus, but there is still considerable overlap of the other colobine genera (Figure 5.31).

Table 5.32: Inter-generic Cherry et al’s M distance between colobine genera. Co. 0 Mean 1.27 Pi. 0.93 0 Range 1.33 Pro. 1.41 0.74 0 Se. 1.21 0.70 1.11 0 Tr. 1.42 0.90 0.76 0.78 0 Pre. 2.03 1.54 1.04 1.61 0.94 0 Py. 1.5 1.17 1.13 1.27 0.88 1.14 0 R. 1.95 1.42 1.3 1.36 1.03 0.93 1.0 0 N. 1.24 0.95 1.51 1.16 1.35 1.93 1.56 1.72 0 Si. 1.54 1.05 1.58 1.25 1.11 1.85 1.41 1.7 0.86 0 Co. Pi. Pro. Se. Tr. Pre. Py. R. N. Si.

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5.6 Limb Proportions: Published data on postcranial elements and limb proportions for the colobines (Tables 5.1-5.10) was gathered from Napier & Napier (1967), Groves (1989 & 2001a), Schultz (1986), Ciochon, (1993); and Whitehead et al (2005). Limb proportions of the Colobinae were found to be as diverse as the subfamily itself. First, the odd-nosed colobines, Nasalis, Simias, Rhinopithecus and Pygathrix all have very high intermembral indices, >90% (Groves, 2001a). Second, the ranges of variation for indices of Trachypithecus were on par with those of Cercopithecus and Macaca (Tables 5.1-5.10). For example, T. vetulus has an intermembral index of 76% while T. obscura and T. phayrei have 83% (Washburn, 1944; Napier, 1985; Fleagle, 1998). However, on average, intermembral indices for most Lutungs >80% with brachial index <101% (Weitzel et al, 1988). Third, in comparison to the olive (80%) and Black-and-white African colobines (76- 83%), Piliocolobus species were on average rather derived with a mean intermembral index of 86% (79-89%). Interestingly, the range of variation for the intermembral index for Presbytis species was low <80% (75-78%) and a brachial index of >106% (Weitzel et a1, 1988; Fleagle, 1998).

5.7 Genetics of the Colobinae: Until recently, the genetic relationships of the Colobinae were largely unknown. Most colobine species have 44 chromosomes but Nasalis has 48 (Bigoni et al, 2003). The genetic analyses of 424 bp mtDNA tRNAThr gene and cytochrome b gene fragment by Zhang & Ryder (1998; unfortunately this study did not include Procolobus verus, Piliocolobus spp. or Simias concolor) confirm the divergence of two subfamilies of the Cercopithecidae (Cercopithecinae and Colobinae) and another division between the African and Asian colobines. Furthermore, there is a complex evolutionary history within the Asian colobines – one that does not exactly agree with Groves’ generic taxonomy (2001a). For example, species placed in Trachypithecus (e.g. T. johnii) by Groves display greater genetic affinity to Semnopithecus species (e.g. S. entellus) based on the recent genetic analyses by Whittaker et al (2006; Table 5.33), also based on 424 bp from mtDNA tRNAThr gene and cytochrome b genes. Similar results were reported by Geissman et al, (2004), also based on mtDNA. In addition, Whittaker et al’s (2006) work also highlights the close genetic

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relationship between Nasalis larvatus and Simias concolor, which may strengthen arguments for the inclusion of the Simakobu within Nasalis. As mentioned previously, Brandon-Jones (2004) reports hybridizations between species of Semnopithecus and Trachypithecus. Knowledge of these hybridization zones compels Brandon-Jones (e.g. 1996 & 2004) to group together Trachypithecus species within Semnopithecus. Furthermore, like some macaque species, there are some colobine species which display sexual skin swellings whereas others do not (van Schaik et al, 1999; Shelmidine et al, 2007), once again, implying phylogenetic divergences and sexual (i.e. reproductive) behavioral differences within and between genera.

Table 5.33: Inter-generic uncorrected ‘p’ genetic distances between colobine genera from Whittaker et al, 2006. Si. 0 Mean 0.16 N. 0.061 0 Range 0.14 R. 0.134 0.144 0 Py. 0.136 0.171 0.127 0

Se. 0.155 0.167 0.162 0.195 0 Tr. 0.138 0.194 0.141 0.165 0.127 0 Co. 0.177 0.196 0.2 0.17 0.197 0.178 0 Pi. 0.179 0.194 0.178 0.199 0.168 0.18 0.161 0

Si. N. R. Py. Se. Tr. Co. Pi.

5.8 Discussion and Conclusion: From the forgoing analysis many sound inferences and conclusions may be drawn.

5.8.1 Question 1: Are the extant Catarrhini genera adequately defined and do they compose an ‘adaptively coherent’ grouping of one or more species which occupy an ‘adaptive zone’, as defined by Wood & Collard (1999a)? Aside from Trachypithecus and Semnopithecus, the remaining eight genera (Colobus, Piliocolobus, Procolobus, Presbytis, Pygathrix, Rhinopithecus, Nasalis and Simias) are adequately defined, each adaptively coherent and occupying different adaptive zones. Two species grouped within Trachypithecus by Groves (2001a; i.e. T. vetulus & T. johnii) are genetically more closely related to species of Semnopithecus (see Whittaker et al, 2006). Furthermore, T. vetulus and T. johnii have very different intermembral indices compared to the other species of Semnopithecus and some Trachypithecus spp. (Washburn, 1944;

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Napier, 1985; Fleagle, 1998). In addition, Brandon-Jones (2004) reported hybridizations in southern India between species of Trachypithecus and Semnopithecus.

Question 2: In terms of cranial morphology, how distinct are genera? In other words, how readily may a genus be determined from cranial dimensions, considered either singular or multiple? When compared to the other primate groupings thus far presented, cranial distinctions between colobine genera are less extreme. In contrast to the Cercopithecinae, very few cranial dimensions of the Colobinae are significantly correlated to body size (Table 5.19 and Figure 5.27). Cranially, Nasalis and Simias, and Pygathrix and Rhinopithecus are cranially quite peculiar and easily distinguishable from Colobina and Presbytina (Groves, 1970 & 1989). Procolobus verus: The Olive colobine is the smallest living colobine, <5-6 kg. On average, the superior facial height (nas-pros) was less than 40 mm and palatal length A (ol- sta) less than 30 mm (Figures 5.12 and 5.14). Sagittal crests are common in all African colobine genera but fairly absent from Asian colobines; for Procolobus, the sagittal crest begins much more anteriorly on the frontal bone compared to Colobus or Piliocolobus. The maxillo-breadth (biecm) for Procolobus is on average relatively greater than 100% of the palatal length A (ol-sta), whereas Piliocolobus and Colobus are less than 100% (Figure 5.19; Table 5.15, no. 26). The cranial height (bas-br) for Procolobus verus is greater than 70% of superior facial length (bas-pros; Figure 5.17). In addition, the length of the zygomatic bone (zs-zgyi) is 120% longer than palate length (ol-sta; Figure 5.25). Piliocolobus spp.: The Red colobines are generally in between the extremes exhibited by Procolobus and Colobus; although Piliocolobus is larger and much more speciose than Procolobus. The Red and Black-and-White colobines overlap in dimensions such as, superior facial and bizygomatic breadths (bifmt and bizygo; Figures 5.7 and 5.10). Interestingly, despite differences in size, Piliocolobus and Procolobus overlap in their bizygomaxillare superior breadths (bizs; Figure 5.9). Colobus spp.: Black-and-white colobines are the largest of the African colobines and exhibit more terrestrial behaviors than either the Olive or Red colobines. Black-and- white colobines have a longer face (nas-pros) and palate (ol-sta), and wider maxillo- alveolar breadth (biecm) compared to either Procolobus or Piliocolobus (Figure 5.14). The

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cranial height (bas-br) for Colobus spp. were less than 70% of superior facial length (bas- pros; Figure 5.17). Furthermore, the length of the zygomatic bone (zs-zgyi) is on average less than 100% of the palate length (ol-sta; Figure 5.25). Presbytis spp.: The Surilis are very small colobines in Southeast Asia, similar in size to Procolobus. In fact, Surilis on average have a smaller superior facial height than the olive colobine, Procolobus (Figure 5.12). Metrically and proportionally, Presbytis is very similar to Procolobus; which is most likely the result of similar body size between the two genera. However, the relative percentage of cranial height (bas-br) to superior facial length (bas-pros) for Presbytis is on average greater than the other members of the Presbytina (and Procolobus) (Figure 5.17; Table 5.15, no. 2). Trachypithecus spp.: The Lutungs are geographically and cranially in between Semnopithecus and Presbytis (similar in most regards to Piliocolobus with Colobus and Procolobus; see above); Trachypithecus spp. are sympatric with Semnopithecus spp. in the most Western portion of their geographic range, whereas sympatric with Presbytis spp. in the most Eastern portion of its geographic range (Bennett & Davies, 1994). Although sagittal crests are not common in Asian colobines, they may have deep or strong temporal lines. The length of the face (nas-pros) in Lutungs is on average larger than Presbytis but smaller than Semnopithecus (Figure 5.12). Semnopithecus spp.: The Langurs are the largest distributed colobines and occupy forests at sea level or up to +3000m above sea level. Compared to the samples for Presbytis and Trachypithecus, Semnopithecus species from northern India are very large but species in the south are much smaller. Within the Presbytina, the sample for Semnopithecus generated some of the largest maxillo-alveolar and bizygomatic breadths (biecm) and superior facial heights (nas-pros; Figures 5.5, 5.10 & 5.12). Pygathrix spp.: The Douc Langurs are highly arboreal and, like Rhinopithecus, have an absolutely and relatively wide upper face (Figure 5.7). Metrical and proportional overlap between Pygathrix and Rhinopithecus is common (Figures 5.2, 5.3 and 5.5). On average, Pygathrix and Rhinopithecus have large biasterionic breadths (Figure 5.4); the latter produced some of the largest figures. Pygathrix spp. have some the largest bizygomaxillare superior breadths (bizs) of all colobines (Figure 5.9); this dimension is significantly different to Rhinopithecus.

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Rhinopithecus spp.: The snub-nosed colobines have long fascinated primatologists and all are at present endangered. There is a large difference in size between R. roxellanae and R. avunculus (see Figure 5.11), as well as each occupying radically different habitats. Rhinopithecus spp. have greatly reduced nasal bones and sometimes maybe completely absent. Increase in size for larger species could be a consequence of climate and latitude (Davison, 1982). Nasalis larvatus: The Proboscis monkey is one of the largest cercopithecines and can be readily identified by its absolutely and relatively long nasal bones (Figure 5.13) and narrow interorbital pillar (Groves, 1970 & 1989). Compared to Pygathrix and Rhinopithecus, Nasalis and Simias both have narrow upper faces with long nasal bones, but there is overlap with some African colobines (Figure 5.7). Additionally, considering that all Nasalina genera have intermembral indices above 90% could suggest these genera, or their ancestors, were much more terrestrial than extant representatives. Simias concolor: The Simakobu is most likely the closest living relative to Nasalis yet they appear dramatically different. The bizygomaxillare superior breadth (bizs) for the Simias sample was different from all other colobines (Figure 5.9); however, this finding needs better substantiation with larger sample sizes. Within the Nasalina genera, Simias has the smallest maxillo-alveolar breadth (biecm; Figure 5.5). For its body size, Simias has a very long zygomatico-maxillary suture (zs-zi; Figure 5.16), compared to Pygathrix and Rhinopithecus but similar to Nasalis. Question 3: How much morphological variation is encompassed within a Catarrhini genus? Morphological variation in colobine genera can produce coefficients of variation < 5% to ~ 20% (Table 5.11). Highly variable measurements are associated with the nasal and facial region, especially nasal height (nas-ns), sagittal length of the nasal bones (nas-rhi), inferior breadth of the nasal bones (inbrnabo) palatal length and height (ol-sta and palhei). In contrast, some neurocranial variables exhibit much less variation. Intra-generic Euclidean distances based on Ln transformed data were on average 0.82 (Max. - Min., Semnopithecus, 1.01 – Procolobus, 0.63) while those based on MSV were 0.57 (Max. - Min., Piliocolobus and Trachypithecus, 0.63 - Procolobus, 0.50) (Tables 5.22 & 5.25).

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Question 4: How much has one genus morphologically and genetically diverged from another? With regard to morphological divergence, measuring size or shape differences produces different results. When Ln transformed data was analyzed, the greatest Euclidean distance produced was between Nasalis and Presbytis (Tables 5.21 and 5.27); the smallest distance was between Procolobus and Presbytis. However, when shape differences were measured, the greatest Euclidean distance produced was between Colobus and Presbytis (Tables 5.24 and 5.30); the smallest distance was between Nasalis and Simias. By calculating Cherry et al’s M distance, the largest distance was again between Presbytis and Colobus but the smallest distance was between Piliocolobus and Semnopithecus (Table 5.32). For comparison, genetic analyses by Whittaker et al (2006; Table 5.33) support close relationship between Nasalis and Simias but the largest distance was between Piliocolobus and Pygathrix.

Question 5: Do cranial morphometric similarities and/or differences reflect adaptive zones? Cranial morphometrics and/or dispositions which may indicate the adaptive zones of these genera include the following. Procolobus verus: The Olive colobine, the smallest living colobine, is highly arboreal and folivorous and prefers well-watered, closed, dense foliage of the lower forest strata. Associated with this is a small cranium with very little facial projection and both males and females can develop sagittal crests which begins placed anteriorly on the frontal bone. The palate is relatively and absolutely very small (see also generic description for question 2 above and generic summaries below). Piliocolobus spp.: Red colobines prefer the middle to upper forest strata and has a diverse diet but exhibits some terrestrial behaviors to forage for food. Males and females can develop strong temporal lines and/or sagittal crest. Size of some cranial dimensions (i.e. size) overlaps more with those of Colobus than Procolobus, although when relative percentages are examined Piliocolobus generally falls in between these genera (see also generic description for question 2 above and generic summaries below). Colobus spp.: Black-and-White colobines exploit a variety of niches not limited to particular forest strata; can and do consume mature foliage and do exhibits some

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terrestrial behaviors to forage for food (Oates, 1977; Rowe, 1996). In general Black-and- White colobines, males and females, have a cranium that has relatively robust and dramatic, with strong nuchal and temporal lines compared to Piliocolobus and Procolobus. The palate is relatively and absolutely longer than Procolobus (see also generic description for question 2 above and generic summaries below). Presbytis spp.: Surilis, the smallest of the Asian colobines, are highly arboreal, and during particular seasons can and do consume seeds and fruits. Associated with this is a globular cranium similar to Procolobus in size but lacking a sagittal crest. Surilis have a relatively small face with very little facial projection but relatively large neurocranium (see also generic description for question 2 above and generic summaries below). Trachypithecus spp.: Lutungs are highly arboreal, can range in body size from 6 to 20 kg (Weitzel et al, 1988) and can consume mature foliage. Associated with this is a cranium similar in size to Piliocolobus although sagittal crests are lacking but robust temporal lines can develop. Size of some cranial dimensions (i.e. size) overlaps more with those of Presbytis and Semnopithecus, although when relative percentages are examined Trachypithecus generally falls in between these two genera (see also generic description for question 2 above and generic summaries below). Semnopithecus spp.: Langurs are arboreal and terrestrial and have varied diets. The body sizes of langurs are clinally distributed with large species in the North (e.g. S. schistaceus) and smaller species in the South (S. priam). Associated with this is a cranium that is very large in northern species and greatly reduced in southern species. On average, the palatal length was greater than Trachypithecus spp (see also generic description for question 2 above and generic summaries below). Pygathrix spp.: Doucs langurs are highly arboreal, have peculiar soft-tissue nasal anatomy similar to Rhinopithecus spp. Associated with this is a cranium with a very wide upper face and the nasal bones may or may not be present (this is true for Rhinopithecus as well). Douc langurs and Rhinopithecus spp. do not develop sagittal crests (see also generic description for question 2 above and generic summaries below). Rhinopithecus spp.: Snub-nosed monkeys are arboreal and terrestrial, have varied diets, and have peculiar soft-tissue nasal anatomy similar to Pygathrix. Associated with this is a cranium with a wide upper face (bifmt), small facial height (nas-pros) and the nasal

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bones may or may not be present. The neurocranium is in general globular (see also generic description for question 2 above and generic summaries below). Nasalis larvatus: The Proboscis monkey is arboreal and terrestrial, adult males develop a large bulbous nose but females and juveniles have peculiar soft-tissue nasal anatomy similar to Simias. Associated with this is a cranium with a long but narrow face. Sagittal crests are common, as are strongly developed nuchal crest. These features, along with limb proportions, could suggest Nasalis, or the ancestor of Nasalis was once much more terrestrial (see also generic description for question 2 above and generic summaries below). Simias concolor: The Simakobu is arboreal and terrestrial and peculiar soft-tissue nasal anatomy similar to Nasalis. Associated with this is a cranium very similar to Nasalis but much smaller in size. In addition, the cranium has relatively long zygomatic bones and long zygomatico-maxillary suture but a narrow bizygomaxillare superior breadth. The palatal length is similar in size to Trachypithecus spp (see also generic description for question 2 above and generic summaries below).

Question 6: What analogies may be drawn from extant Catarrhini genera for the interpretation of fossil hominin genera? The colobines and their generic arrangement can provide analogies and considerations for the interpretation of fossil hominin genera. First, when treated with multivariate statistics colobine genera overlap considerable, suggesting the crania are in general very similar. This may suggest that colobine crania are rather conservative, not having dramatically altered between genera, despite niche differences, although the crania of Nasalina genera are particularly distinct. Second, sister species (or genera) can have radically different phenotypes (e.g. Nasalis and Simias). Third, increased terrestrialism may result in increases to body size and more facial projection. Lastly, limb proportions reveal stark differences between genera, which are related to locomotion and substrate use (Schultz, 1986; Napier, 1985).

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5.8.2 Generic Summaries: Subfamily: Colobinae (or Colobidae, see Groves, 1989 and Jablonski, 2002; leaf-eating, complex stomach monkeys) Tribe Colobini, Subtribe: Colobina (African colobines - Genetic analyses by Goodman et al (2001) suggests African and East Asian colobines diverged ~ 10 Ma.) Colobus Illiger, 1811. Black-and-White colobines, a polytypic genus comprising five species; C. satanas (Black Colobus), C. angolensis (Angola Colobus), C. polykomos [type species of genus; King Colobus; 2n = 44], C. vellerosus (Ursine Colobus) & C. guereza (Mantled Colobus). Three-chambered stomach as opposed to four. In addition, fossils have been allocated to this genus (Delson, 1975; Leakey, 1988; Pickford & Senut, 1988; Delson, 1994; Benefit, 2000; Gundling & Hill, 2000; Delson et al, 2000; Jablonski, 2002). Many in the past have grouped all the African colobine species under Colobus (e.g. James, 1960 and Napier & Napier, 1967). Actually, Colobus species are the most derived and distinctive of the African colobines (Hull, 1979), while Piliocolobus and Procolobus share several features in common (which is why some group them under one genus, Procolobus e.g. Oates et al, 1994; Delson et al, 2000). Piliocolobus Rochebrune, 1887. Red colobines, a polytypic species comprising nine species; P. badius [type species of genus; Western Red; 2n – 44], P. pennantii (Pennant’s Colobus), P. preussi (Preuss’s Colobus), P. tholloni (Thollon’s Red Colobus), P. foai (Central African Red Colobus), P. tephrosceles (Ugandan Red Colobus), P. gordonorum (Uzungwa Red Colobus), P. kirki (Zanzibar Red Colobus; 2n – 44), & P. rufomitratus (Tana River Red Colobus). Surprisingly, unlike Colobus, female species of Piliocolobus and Procolobus have been observed to temporarily allomother their infants (Davies & Oates, 1994). Where sympatric, Piliocolobus species are vulnerable prey to chimpanzees (Boesch & Boesch, 1989; Mitani & Watts, 1999). Procolobus Rochebrune, 1887. A monotypic genus, P. verus. The Olive Colobus is the smallest living colobine (~ 4.5kg), they have greatly reduced thumbs, a four- chambered stomach (another characteristic shared by Piliocolobus) and females display cyclical sexual swelling, while young males have perineal organ (Oates et al, 1994). Additionally, female olive colobines are the only Old World monkeys which carry their infants in their mouth while in transit (Rowe, 1996). Some consider Piliocolobus a

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subgenus of Procolobus (Oates et al, 1994; Rowe, 1996; Delson et al, 2000; Grubb et al, 2003).

Tribe Presbytini, Subtribe Presbytina (Langur group) Semnopithecus Desmarest, 1822; Sacred, Gray or Indian Langurs. A polytypic genus comprising seven species; S. schistaceus (Nepal Gray Langur), S. ajax (Kashmir Gray Langur), S. hector (Tarai Gray Langur), S. entellus [type species of genus; Northern Plains Gray Langur; 2n = 44], S. hypoleucos (Black-footed Gray Langur), S. dussmieri (Southern Plains Gray Langur) & S. priam (Tufted Gray Langur). Semnopithecus species occupy the widest range of altitudes next only to humans; from sea-level to >4000m (Rowe, 1996). Thus, the flexible adaptability of Semnopithecus is not surprising, as well as its sympatry with Trachypithecus and Macaca (Bennet & Davies, 1994). In addition, there is an clinal trend in size within Semnopithecus (akin to clinal distributions seen in Homo, Macaca and Papio) because species in the north (S. schistaceus & S. ajax) of India, at the foothills of the Himalayas, are very large compared to species in the south (S. priam) (Bennett & Davies, 1994).In contrast to the other Asian colobines, Semnopithecus species are known for infanticide (Yeager & Kool, 2000). Barry (1987) reported the presence of “Presbytis sivalensis” fossils in the Siwalik formations of Pakistan and India but this is most likely not the Presbytis as it is known today (Groves, 2001a) and probably represents a species of langur. Trachypithecus Reichenbach, 1862; Lutungs. A polytypic genus comprising 17 species in five species groups; (1-Vetulus-group T. vetulus; Purple-faced Langur, sometimes given generic or subgeneric status, Kasi, e.g. Fleagle, 1998 and Hill, 1934 & 1936; T. johnii (Nilgiri Langur, also, given generic or subgeneric status, Kasi)), (2- Cristatus-group T. auratus [type species of genus; Javan Lutung], T. cristatus (Silvery Leaf Monkey or Lutung; 2n = 44), T. germaini (Indochinese Lutung), T. barbei (Tenasserim Lutung)), (3-Obscurus-group T. obscurus (Dusky or Spectacled Leaf Monkey; 2n = 44), T. phayrei (Phayre’s Leaf Monkey)), (4-Pileatus-group T. pileatus (Capped Langur), T. shortridgei (Shortridge’s Langur), T. geei (Gee’s Golden Langur)), (5- Francoisi-group T. francoisi (Francois’s Langur), T. hatinhensis (Hatinh Langur), T. poliocephalus (White-headed Langur), T. laotum (Loatian Langur), T. delacouri

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(Delacour’s Langur) & T. ebenus (Indochinese Black Langur)). Bennett & Davies (1994) describe Trachypithecus as one of the most ‘diverse array’ colobine species. Furthermore, Presbytis and Trachypithecus species are sympatric in some areas. However, species are exploiting different areas of the same environment or prefer dissimilar plant-food species, both of which vary with seasonality. In addition, fossil species have also been allocated to this genus from Java, ~ 1.9 Ma (Jablonski & Tyler, 1999). Presbytis Eschscholtz, 1821; Surilis. A polytypic genus comprising 11 species; P. melalophos [type species of genus; Sumatran Surili], P. femoralis (Banded Surili), P. natunae (Natuna Islands Surili), P. chrysomelas (Sarawak Surili), P. siamensis (White- thighed Surili), P. frontata (White-fronted Langur), P. comata (Javan Surili), P. thomasi (Thomas’s Langur), P. hosei (Hose’s Langur), P. rubicunda (Maroon Leaf Monkey) & P. potenziani (Mentawai Langur or Joja). In the past the genus Presbytis incorporated most East Asian colobines now placed in Semnopithecus and Trachypithecus (e.g. James, 1960; Napier & Napier, 1967). Generally, the stomachs of Presbytis species are relatively smaller compared to Trachypithecus (Bennett & Davies, 1994), although the former does eat more fruit and seeds than the latter, which eats more mature leaves (Caton, 1999; Yeager & Kool, 2000). Presbytis species, similar to Trachypithecus, have considerable variation in fur color and pattern and their infants may be a different color (Rowe, 1996; MacDonald, 2001; Hutchins et al, 2003).

Tribe Presbytini, Subtribe Nasalina (Odd-nosed group) (Please note, the monophyly of the odd-nosed colobines and their relationship to fossil species, e.g. Mesopithecus, have been questioned (morphological cladistics - Jablonski, 1995 & 1998) and supported (molecular data - Ming et al, 2004; Sterner et al, 2006).) Pygathrix E. Geoffroy, 1812; Douc langurs. A polytypic genus comprising 3 species; P. nemaeus [type species of genus; Red-shanked Douc; 2n = 44], P. nigripes (Black-shanked Douc) & P. cinerea (Gray-shanked Douc). Genetic analysis by Bigoni et al (2004) of East Asian colobinae chromosomal karyotypes demonstrated Pygathrix nemaeus is the most basal, and karyotypically the most conservative. Furthermore, this genetic analysis confirms the monophyly and divergence of African and East Asian colobinae clades. Also, the results of Bigoni et al (2004) are congruent with the cladistic analysis by Jablonski (1995 & 1998), which also placed Pygathrix as one of the primitive members of

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the odd-nosed colobines, having changed less than the others from the hypothesized ancestral mophotype. In addition, fossil species have also been allocated to this genus (Delson, 1994). In some areas, Pygathrix is sympatric with Hylobates, Tachypithecus and Macaca (Lippold, 1998). Rhinopithecus Milne-Edwards, 1872; Snub-nosed monkeys. A polytypic genus comprising 4 species; R. roxellana [type species of genus; Golden Snub-nosed Monkey], R. bieti (Black Snub-nosed Monkey), R. brelichi (Gray Snub-nosed Monkey) & R. avunculus (Tonkin Snub-nosed Langur, sometimes given subgeneric status, Presbyticus). Like Pygathrix, Rhinopithecus are known for their colorful pelage and unique soft-tissue nasofacial anatomy, which unites them phylogenetically (Groves, 2001a). In addition, fossil species have also been allocated to this genus (Jablonski & Yumin, 1991). Like northern species of Semnopithecus, species of Rhinopithecus must also endure below freezing temperatures and snow during the winter months (Bennet & Davies, 1994). Sexual dimorphism between males and females can be quite large (Jablonski & Pan, 1995). Species of Rhinopithecus, particularly Rhinopithecus roxellanae and R. bieti, can exploit a range of terrestrial and arboreal niches, some of which are subject to snowstorms during the winter months (Davison, 1982; Wu, 1993; Rowe, 1996; Long, 1998; Su et al, 1998; Li, 2007). Fossil specimens have been allocated to this genus (Jablonski & Yuerong, 1988; Jablonski, 2002). Nasalis E. Geoffroy, 1812; Proboscis monkey. A monotypic genus, N. larvatus. 2n = 48. The Proboscis monkey, confined to Borneo, is famous for the male’s large bulbous nose and ease with moving between habitats by means of swimming (Oates & Davies, 1994). Cranially, Nasalis and Simias are peculiar. Features which distinguish the two include the longer sagittal length of the nasal bones in comparison to other colobines and a narrow interorbital pillar (Groves, 1970). Nasalis is further separated from the other colobines by thinner tooth enamel, the mandibular ascending ramus slopes posteriorly and supraorbital ridges are absent (Groves, 1989). In addition, Havarti (2000) has reported that the dental eruption sequence of Nasalis is unlike other colobines. Furthermore, karyological analyses by Bigoni et al (2003) places Nasalis as the most derived member of the Asian colobines, yet still a member of this clade. Just as Pygathrix and Rhinopithecus are united by soft nasofacial anatomy, Nasalis and Simias also share unique soft-tissue features as

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well (Groves, 2001a). If Simias is subsumed under Nasalis, N. larvatus is the type species of the genus. No fossil specimens have been allocated to this genus. Simias Miller, 1903; Simakobu or Pig-tailed langur. A monotypic genus, S. concolor. 2n = 48? The simakobu monkey is a very little studied primate and is critically endangered, deserving immediate conservation attention. A recent genetic analysis by Whittaker et al (2006) conclusively shows a more immediate relationship with Nasalis than with any other Asian colobine. This perhaps strengthens morphological arguments for its subgeneric placement within Nasalis (e.g. Groves, 1970 and Szalay & Delson, 1979). However, conservation efforts could perhaps be improved if distinctiveness were stressed. Oddly though, Simias is the only colobine with a very short tail (Rowe, 1996). No fossil specimens have been allocated to this genus.

277 Chapter 6: Results for the genera of the Hominoidea

Hylobates Pongo Gorilla Pan Homo

Panina Hominina

Gorillini Hominini

Ponginae Homininae

Hylobatidae Hominidae Hominoidea Figure 6.1: A taxonomic and phylogenetic Catarrhini diagram representing the likely relationships Simiiformes within the Hominoidea based on Haplorrhini biomolecular and morphological data. Primates

6.1 Introduction: The purpose of this chapter is to report and discuss the results for the extant genera of the Hominoidea. The Superfamily Hominoidea (Simpson, 1931) includes five genera; Hylobates, Pongo, Gorilla, Pan and Homo, which together comprise 21 described species (Figure 6.1; Groves, 2001a; see also Andrews & Harrison, 2005). However, over half of these species (14) belong to Hylobates, the gibbons and siamang, which may be further subdivided into four subgenera. The average number of species per genus in the extant Hominoidea is 4.2, ranging from one to fourteen with a standard deviation of 5.5. In addition, if Hylobates is excluded and only the great apes are considered, the average number of species per genus drops to 1.8, ranging from one to two with a standard deviation of 0.5; lastly, if only hylobatid subgenera are considered, the average number of species per subgenus is 3.5, ranging from one to seven with a standard deviation of 3.0. In contrast, the Superfamily Cercopithecoidea, sister-superfamily to the Hominoidea, have on average 6.2 species per genus (n=21) ranging from one to twenty- five with a standard deviation of 6.8. However, if genera which contain ten or more species are omitted (Macaca, Cercopithecus, Trachypithecus, Presbytis), the average number of species per genus is reduced to 3.4, ranging from one to nine with a standard deviation of 2.5. Based on paleontological and genetic data, the origins of the Hominoidea lie in the Oligocene-Miocene (Simons, 1985; Gebo et al, 1997; Ward et al, 1999; Begun, 2002b; Kelley, 2002; Pilbeam, 2002; Ward & Duren, 2002; Steiper et al, 2004; Finarelli & Clyde, 2004; Raaum et al, 2005). The species of hominoid genera range in size from less than five kilograms (Hylobates) to greater than 200 kg (Gorilla). Gorilla and Pan are largely terrestrial quadrupeds (Sarmiento, 1998); although both can and do exploit arboreal resources and the latter sleeps in trees and both are restricted to Africa’s tropical forested environments. Hylobates and Pongo are strictly arboreal (rarely coming to or travelling upon the ground) and occupy tropical forests of East and Southeast Asia. Homo is an obligate biped and globally distributed but originated in Africa during the Plio-Pleistocene (Leakey et al, 1964; Leakey, 1973a, 1973b, 1974 & 1976; Leakey & Wood, 1973; Day et al, 1973, 1975 & 1976; Leakey & Walker, 1976 & 1985; Brown et al, 1985; Wood, 1991 & 1992; Tobias, 1991; Hill et al, 1992; Schrenk et al, 1993; Prat et al, 2005). Hominoids are morphologically united by shared features found in the dentition and postcranial anatomy, particularly the pectoral girdle and forelimb (Groves, 1972; Andrews & Groves, 1976; Andrews, 1985). The living apes may be characterized mainly as frugivores (Andrews & Martin, 1991; Singleton, 2004) but animal prey (including vertebrates and/or invertebrates) is also consumed and species of Gorilla and H. Symphalangus can and do consume vegetation and leaves (Chivers, 1974; Remis, 2003). Currently, the fossil record is virtually silent on the evolution of Gorilla and Pan. However, 10-10.5 Myr old (Late Miocene) gorilla-like dental specimens assigned to Chororapithecus abyssinicus have been recovered from Ethiopia (Suwa et al, 2007). In addition, dental specimens of Pan were recently recovered from mid-Pleistocene age deposits in Kenya (McBrearty & Jablonski, 2005). In contrast, fossil material exists which can be demonstrated as belonging to the evolutionary lineages related to Pongo and Homo (e.g. Sivapithecus for the former and Australopithecus for the latter; see Andrews & Cronin, 1982; McHenry & Skelton, 1985; McHenry & Coffing, 2000). The Miocene pliopithecoids and their relationship to modern gibbons, if any, remains to be conclusively demonstrated and perhaps may be the result of primitive or convergent facial morphology (Simons & Fleagle, 1973;

279 Fleagle, 1984; Begun, 2002a). Tables 6.1 - 6.5 provide important features describing the extant hominoids and their respective adaptive zones.

Table 6.1: Hylobates (including subgenera Hylobates, Nomascus, Hoolock & Symphalangus) spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) Average Habitat (,; (range) Min.-Max.) 7135, 6500*; Intermembral Brachiation, Frugivore with Arboreal/ Monogamous/ 4000-15000 129 (121-155) below branch, some folivory Tropical rain H. lar 2.1 & H. Brachial bipedal on forest, middle hoolock 1.89- 113 (105-124) large branches and upper 1.94 (~110 g Crural & ground strata average adult 83 brain weight)

Table 6.2: Pongo spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) Average Habitat (,; (range) Min.-Max.) 77500, Intermembral Suspensory- Frugivore + Arboreal and 1 solitary male 37000*; 144 (135-150) below branch, some animal limited territory which 35000- Brachial Quadrumanous prey terrestrial/ encompasses 85000 100 (92-109) climbing Tropical rain the territories Crural forest of different 90 females/ 1.08-1.69 (413.3g adult brain weight - P. pygmaeus)

Table 6.3: Gorilla spp. Body Weight Limb Indices Locomotion Diet Substrate/ SS/EQ (g) Average Habitat (,; (range) Min.-Max.) 170000, Intermembral Quadrupedal- Herbivore + Terrestrial with 1 male- 83100*; 117 (110-125) knuckle- frugivore arboreal multifemale/ 70000- Brachial walking, with capabilities/ .85-1.34 180000 80 (73-86) climbing, Tropical rain (505.9g adult Crural below branch forest brain weight - 84 (82-86) G. gorilla)

280 Table 6.4: Pan spp. Body Weight Intermembral Locomotion Diet Substrate/ SS/EQ (g) Index Habitat (,; Min.-Max.) 45000, Intermembral Quadrupedal- Frugivore + Terrestrial & Multimale- 39500*; 107 (102-117) knuckle- animal prey arboreal/ multifemale 30000-55000 Brachial walking, with Tropical forest/ fission-fusion/ 93 (87-100) climbing above woodland 1.42-1.89 Crural & below (410.3g adult 86 branch brain weight - Pan troglodytes)

Table 6.5: Homo sapiens Body Weight Intermembral Locomotion Diet Substrate/ SS/EQ (g) Index Habitat (&; Average Min.-Max.) (range) 85000, Intermembral Habitual Omnivore Terrestrial/ Varied, 40000; 72 (65-79) Bipedalism Varied Monogamy & 40000-125000 Brachial polygamy/ 76 (65-85) 4.3-5.8 (1250g Crural adult brain 84 weight)

With regard to competing classificatory schemes for the Hominoidea there is consensus for the hylobatids but there is considerable disagreement as to the arrangement and hierarchical partitioning of the great ape clade (Cela-Conde, 1998; Andrews & Harrison, 2005; Tuttle, 2006). In general, most previous classifications grouped all gibbons within Hylobates (Groves, 1972, 1989 & 2001a) although sometimes the siamangs are given generic status, Symphalangus (e.g. Schwartz et al, 1978; Goodman et al, 1998). However, in recent years further intra-generic divisions have been employed because of differing diploid numbers between species, genetic divergence estimates and biogeography (Hall et al, 1998; Roos & Geissmann, 2001; Geissmann, 2002; Chatterjee, 2006). As such, Hylobates contains four subgenera, including Hylobates (7 species; 2n=44), Hoolock (2n=38; see Mootnick & Groves, 2005), Nomascus (5 species; 2n=52) and Symphalangus (2n=50). In contrast, all the Great ape species have 48 chromosomes but modern humans have 46. Largely as a result of Simpson’s mammalian taxonomy (1945 & 1963) the lesser and great apes (excluding Homo) were placed in the family Pongidae, while Hominidae was reserved for humans and their bipedal fossil ancestors. In fact, during the 1970’s some actually grouped gorillas and chimpanzees together in Pan (e.g. Tuttle & Basmajian, 1974). However, continued research has unequivocally demonstrated the

281 sister species status of chimpanzees and modern humans based on hard and soft tissue anatomy and genetics (Groves, 1986; Begun, 1992 & 1994; Ruvolo, 1997a; Page & Goodman, 2001; Gibbs et al, 2002; Lockwood et al, 2004; Uddin et al, 2004). However, this conclusion continues to be challenged by a small minority (Tuttle, 2006; Grehan, 2006 and references therein). Still others question the reliability of phylogenetic methods (using morphological features) in constructing the correct evolutionary history of a species and their closest relatives (Corruccini, 1994; Collard & Wood, 2000 and 2001b).

6.2 Descriptive Statistics and Univariate Analyses for cranial variables: Table 6.6 provides the descriptive statistics for the pooled sex samples of the thirty-six measurements collected for the Hominoidea. Due to sexual dimorphism, the samples for Gorilla and Pongo produced the largest standard deviations and coefficients of variation. Like the other primate groupings for this study, the nasal and palatal regions and their associated dimensions were the most highly variable. In particular, the inferior breadth of the nasal bones (inbrnabo), height of the nasal aperture (rhi-ns), palatal height (palhei) and the maximum width of the nasal aperture (maxnawi) varied considerably. In contrast, cranial dimensions that were not highly variable include maxillary and palatal breadths (biecm and bienm) and interentoglenoid breadth (bien). The box-plots for the Hominoidea generally reveal two patterns. First, due to the extreme differences in size, Hylobates and Gorilla are easily separated from Pan, Pongo and Homo. As such, like the cercopithecins, the extant apes form a small (<20kg), medium (>20-<100kg) and large (>100kg) ape scale (although the scale, range of variation and sexual dimorphism are greater in apes). Second, because of the exceptionally large neurocranium and greatly reduced non-projecting viscerocranium, Homo is quite distinct from the other apes. As such, fewer variables exhibit clear scaling trends related to body size as seen in the other primate groupings. Examples which demonstrate these patterns may be seen in Figures 6.2 to 6.16. For example, the cranial height (Figure 6.2, bas-br) of Homo was well beyond that of the other apes. However, Gorilla had the greatest inferior cranial length (Figure 6.3, bas-nas). The degree to which Homo has experienced a reduction of the viscerocranium may be observed in such variables as superior facial height and length, and palatal length A (nas-pros, bas-pros and ol-sta, Figures 6.4, 6.13 and 6.13).

282 Table 6.6: Hominoidea pooled sex. g-o pros-o biast bipor biaur bizygo bifmo bifmt bizs bizi zs-zi zs-zgyi Hylo. (n=31) Mean 86.9 112.7 56.1 63.2 68.5 75.5 59.4 61.9 40.4 54.5 20.0 38.8 Median 84.7 110.0 55.2 62.4 66.7 73.0 58.1 62.0 39.9 51.6 19.9 39.0 Maximum 99.0 130.0 67.0 72.0 81.6 92.0 73.4 74.7 49.9 66.3 25.8 45.3 Minimum 74.0 94.0 46.9 53.0 58.0 62.0 53.1 50.0 28.7 44.4 14.9 32.9 Range 25.0 36.0 20.1 19.0 23.6 30.0 20.3 24.7 21.2 21.8 10.9 12.4 SD 7.0 11.6 4.1 5.9 7.2 8.8 4.2 6.0 5.3 7.0 3.1 3.8 CV 8.0 10.3 7.3 9.4 10.6 11.7 7.1 9.8 13.2 12.8 15.3 9.9

Pongo (n=26) Mean 125.8 208.7 105.0 113.6 121.7 149.7 82.2 101.7 62.3 112.5 40.4 57.1 Median 124.8 212.4 103.0 112.2 120.5 145.5 80.8 99.9 61.6 112.8 39.7 57.5 Maximum 144.0 246.0 132.0 139.0 149.0 183.0 95.0 124.2 75.6 135.0 55.6 75.4 Minimum 115.0 181.0 84.0 97.0 102.0 119.0 72.5 87.0 54.1 87.1 28.1 40.5 Range 29.0 65.0 48.0 42.0 47.0 64.0 22.5 37.3 21.5 47.9 27.6 34.9 SD 7.3 19.1 13.3 11.5 12.4 19.3 5.7 11.2 5.8 12.1 7.2 8.2 CV 5.8 9.1 12.7 10.1 10.2 12.9 6.9 11.0 9.2 10.7 17.7 14.4

Gorilla (n=34) Mean 176.5 258.6 115.4 129.1 134.6 161.7 106.8 126.0 58.1 118.7 49.0 76.7 Median 181.6 252.9 112.6 128.4 130.5 162.1 106.0 126.7 59.5 116.7 48.5 76.6 Maximum 216.9 325.3 157.1 150.1 155.6 188.0 128.3 147.1 79.3 140.0 65.7 100.5 Minimum 143.0 217.0 93.0 112.0 118.0 137.0 89.4 107.0 39.7 97.9 37.1 61.8 Range 73.9 108.3 64.1 38.1 37.6 51.0 39.0 40.1 39.6 42.1 28.6 38.7 SD 20.8 33.2 16.8 10.5 11.2 16.1 8.8 13.2 8.7 11.6 7.1 9.8 CV 11.8 12.8 14.6 8.1 8.3 9.9 8.3 10.5 15.0 9.8 14.5 12.7

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g-o pros-o biast bipor biaur bizygo bifmo bifmt bizs bizi zs-zi zs-zgyi Pan (n=33) Mean 135.3 190.3 82.9 104.7 113.9 125.8 88.4 104.6 53.5 87.7 33.0 56.9 Median 136.0 189.3 82.0 104.0 113.0 125.0 88.0 105.0 54.3 87.0 33.6 55.9 Maximum 148.2 210.3 97.3 122.0 134.2 141.7 100.7 118.0 63.1 96.7 39.0 67.1 Minimum 124.0 167.5 72.0 92.8 102.9 113.7 75.9 93.0 41.2 78.8 27.3 50.0 Range 24.2 42.8 25.3 29.2 31.2 28.0 24.8 25.0 21.9 18.0 11.7 17.1 SD 6.0 10.4 5.9 6.4 6.7 7.8 5.2 5.3 5.5 4.3 3.4 4.7 CV 4.5 5.5 7.1 6.1 5.9 6.2 5.9 5.1 10.3 4.9 10.2 8.3

Homo (n=20) Mean 174.8 198.7 105.3 110.1 118.6 122.6 96.3 102.4 56.0 97.8 31.6 52.2 Median 175.7 199.3 105.1 109.9 118.6 123.9 96.8 102.8 55.9 99.4 32.2 53.1 Maximum 190.7 220.8 120.3 121.7 130.3 140.5 108.3 116.6 67.7 108.1 38.8 61.2 Minimum 160.5 178.2 96.8 98.1 107.1 99.7 83.4 88.9 45.5 85.3 24.3 42.8 Range 30.2 42.6 23.5 23.7 23.2 40.9 24.9 27.7 22.2 22.9 14.6 18.4 SD 8.5 10.5 5.7 7.1 6.1 9.7 5.7 6.8 6.2 6.5 3.8 4.4 CV 4.9 5.3 5.4 6.5 5.2 7.9 6.0 6.6 11.1 6.6 12.1 8.5

284 Table 6.6 continued: Hominoidea pooled sex. bas-pros bas-nas bas-br nas-pros nas-br br-lam lam-opn nas-rhi inbrnabo rhi-ns maxnawi nas-ns Hylo. (n=31) Mean 84.2 68.3 55.5 34.9 60.1 34.1 31.1 11.4 9.7 19.5 15.2 30.6 Median 80.0 67.0 55.8 33.6 61.0 33.6 30.4 11.1 10.0 19.0 15.4 29.3 Maximum 104.0 82.0 69.0 45.0 70.0 47.8 43.2 19.0 14.1 28.9 18.5 41.1 Minimum 69.0 58.0 46.0 24.1 49.4 22.0 21.0 7.1 5.3 13.2 13.0 21.7 Range 35.0 24.0 23.0 20.9 20.6 25.8 22.2 11.9 8.8 15.7 5.5 19.5 SD 11.6 7.5 4.9 5.8 5.0 6.6 4.7 2.6 2.2 3.9 1.5 5.4 CV 13.7 10.9 8.8 16.7 8.3 19.4 15.1 22.4 22.6 19.8 9.9 17.5

Pongo (n=26) Mean 154.4 99.1 98.3 93.9 70.2 60.9 67.1 33.5 11.7 33.9 28.7 67.0 Median 156.2 98.0 99.5 94.2 70.7 61.2 65.4 33.7 12.3 34.5 29.5 67.4 Maximum 191.0 115.0 111.1 125.4 83.5 76.0 94.4 42.2 15.7 41.0 34.3 86.1 Minimum 128.2 89.0 89.0 70.7 58.2 46.8 49.6 24.6 5.6 23.2 21.4 43.9 Range 62.8 26.0 22.1 54.7 25.4 29.2 44.8 17.5 10.1 17.8 12.9 42.2 SD 16.8 6.8 6.0 13.8 7.0 6.9 9.4 4.9 2.9 5.1 3.7 9.7 CV 10.9 6.9 6.1 14.7 10.0 11.3 14.0 14.7 25.1 15.0 12.8 14.5

Gorilla (n=34) Mean 178.9 129.8 108.6 109.0 92.5 76.8 82.3 51.2 18.9 33.3 35.6 85.2 Median 179.5 131.7 105.0 106.6 90.1 76.3 77.8 50.9 18.0 33.1 34.4 84.7 Maximum 222.6 161.3 128.0 135.6 114.0 112.9 123.2 68.0 29.0 42.4 49.2 107.9 Minimum 149.6 110.0 91.0 81.7 76.5 49.3 60.1 33.0 11.8 25.4 28.7 60.7 Range 73.0 51.3 37.0 53.9 37.5 63.6 63.0 35.0 17.3 17.0 20.6 47.2 SD 21.6 12.5 11.1 13.2 11.3 14.8 17.9 8.3 4.2 4.6 4.4 11.4 CV 12.1 9.6 10.2 12.2 12.2 19.3 21.8 16.2 22.2 13.8 12.3 13.4

285 bas-pros bas-nas bas-br nas-pros nas-br br-lam lam-opn nas-rhi inbrnabo rhi-ns maxnawi nas-ns Pan (n=33) Mean 134.3 101.8 90.1 83.3 75.9 62.2 55.0 32.3 11.9 26.2 26.8 58.0 Median 134.0 101.0 91.0 83.8 76.4 62.2 55.5 31.4 11.8 26.8 26.7 57.4 Maximum 148.9 111.9 97.3 98.2 86.9 72.4 64.0 39.4 18.0 34.3 31.9 68.2 Minimum 115.9 88.0 83.0 70.8 63.2 49.6 46.3 26.2 7.1 19.1 20.8 48.6 Range 33.1 23.9 14.3 27.4 23.7 22.8 17.6 13.2 10.9 15.1 11.2 19.6 SD 8.9 6.2 3.1 7.0 5.6 5.1 3.7 3.6 2.5 3.6 2.8 4.8 CV 6.6 6.1 3.5 8.4 7.3 8.2 6.7 11.3 21.2 13.9 10.5 8.3

Homo (n=20) Mean 96.2 97.9 131.9 64.3 109.4 109.5 97.4 22.1 16.3 28.8 26.8 48.9 Median 94.9 98.4 130.9 65.1 108.1 108.7 95.0 22.1 16.0 29.3 26.7 49.8 Maximum 109.7 107.2 145.2 72.8 119.5 126.7 117.3 27.3 19.6 32.1 31.1 60.4 Minimum 87.5 88.3 123.8 53.5 103.5 96.2 83.3 15.8 14.1 25.4 23.4 41.4 Range 22.2 18.9 21.4 19.3 16.0 30.5 34.1 11.5 5.5 6.7 7.6 19.0 SD 5.9 5.7 6.2 5.4 4.9 7.3 8.9 3.2 1.5 2.0 2.0 4.0 CV 6.1 5.8 4.7 8.4 4.4 6.7 9.1 14.5 9.2 6.9 7.4 8.1

286 Table 6.6 concluded: Hominoidea pooled sex. maxalvlen biecm iv-pms i1i2 bicanin bicanex palhei biseptal bienm ol-sta ol-pms bien Hylo. (n=31) Mean 44.7 37.5 18.4 12.1 19.3 32.6 7.4 21.3 21.2 42.7 27.9 37.5 Median 42.2 37.5 17.8 11.9 18.8 31.4 7.9 21.0 20.7 40.2 26.9 37.3 Maximum 59.0 44.5 28.0 15.3 25.8 39.2 9.5 25.7 25.5 58.0 39.4 44.0 Minimum 35.0 30.3 13.7 9.4 14.1 25.4 5.0 16.7 18.2 31.6 20.7 30.0 Range 24.0 14.1 14.3 5.9 11.8 13.9 4.5 9.0 7.3 26.4 18.6 14.0 SD 6.9 4.4 3.0 1.6 3.1 3.9 1.2 2.6 2.2 7.8 4.0 3.6 CV 15.5 11.6 16.5 12.9 16.1 11.8 15.8 12.3 10.2 18.2 14.3 9.7

Pongo (n=26) Mean 77.2 66.2 38.7 24.4 38.3 64.2 15.2 42.8 37.9 77.0 59.8 65.5 Median 77.5 67.0 39.4 24.2 37.4 65.7 14.6 42.3 38.2 77.4 60.1 66.7 Maximum 100.0 75.1 48.3 31.8 55.2 81.4 21.0 53.7 47.4 99.0 75.7 77.5 Minimum 65.9 57.5 27.7 20.5 29.4 51.7 11.0 36.4 28.8 62.1 46.1 53.0 Range 34.1 17.6 20.7 11.3 25.8 29.7 10.0 17.3 18.7 36.9 29.6 24.5 SD 7.7 5.6 6.0 2.5 5.7 8.9 3.0 4.4 4.6 8.8 7.0 5.6 CV 10.0 8.4 15.5 10.3 15.0 13.8 19.8 10.3 12.2 11.4 11.8 8.5

Gorilla (n=34) Mean 91.3 70.4 48.6 24.9 39.2 66.9 17.4 44.8 37.9 95.3 69.8 66.8 Median 89.7 70.5 45.1 24.1 39.1 67.1 17.5 44.3 37.5 95.0 67.0 66.7 Maximum 111.3 80.8 73.2 35.1 48.9 86.7 23.7 61.1 44.2 126.2 103.4 79.3 Minimum 78.0 62.5 35.3 21.8 32.8 53.0 11.0 37.3 32.5 75.7 52.1 52.5 Range 33.3 18.3 37.9 13.3 16.0 33.7 12.7 23.8 11.7 50.5 51.2 26.8 SD 9.3 4.2 8.8 2.8 3.7 8.0 3.2 4.4 3.1 13.8 11.3 6.8 CV 10.2 6.0 18.1 11.2 9.5 11.9 18.6 9.8 8.2 14.5 16.2 10.2

287

maxalvlen biecm iv-pms i1i2 bicanin bicanex palhei biseptal bienm ol-sta ol-pms bien Pan (n=33) Mean 66.4 58.4 35.3 22.7 35.2 56.2 11.8 39.9 34.1 68.4 51.5 56.6 Median 65.2 58.0 35.4 22.4 35.3 56.6 12.0 39.7 33.6 68.0 52.0 56.1 Maximum 79.4 66.3 46.1 28.6 43.7 63.9 15.5 49.2 40.2 92.7 67.1 63.1 Minimum 55.6 52.7 27.5 19.8 30.4 46.6 7.7 32.9 29.1 56.1 43.6 49.8 Range 23.8 13.6 18.6 8.8 13.4 17.3 7.8 16.4 11.2 36.6 23.6 13.3 SD 5.8 3.1 3.9 1.9 3.2 4.2 1.8 3.3 2.9 7.2 5.2 4.0 CV 8.8 5.3 10.9 8.5 9.2 7.5 15.1 8.2 8.5 10.5 10.1 7.1

Homo (n=20) Mean 49.0 62.9 27.0 15.1 25.6 40.3 9.7 28.4 40.1 46.0 35.8 72.6 Median 48.5 64.5 26.5 15.0 25.8 40.3 10.0 28.5 40.8 45.3 35.3 72.1 Maximum 59.0 69.5 39.8 17.8 27.8 43.5 13.6 32.5 44.7 55.0 41.5 82.8 Minimum 41.7 55.7 18.1 12.3 22.1 36.4 5.6 23.1 35.1 39.4 28.0 63.5 Range 17.3 13.8 21.6 5.5 5.7 7.1 8.0 9.5 9.6 15.6 13.5 19.3 SD 4.1 3.9 4.6 1.3 1.4 1.9 2.2 2.5 2.8 4.2 3.6 5.2 CV 8.3 6.1 17.2 8.9 5.4 4.6 22.4 8.7 6.9 9.1 10.0 7.2

288 Box Plots for cranial variables of the Hominoidea (Hy. – Hylobates spp., n=31; Pan spp., n=33; Po. – Pongo pygmaeus, n=26; Ho. – Homo sapiens, n=20 & Go. – Gorilla gorila, n=34; variables in alphabetical order by abbreviation): 150 Figure 6.2 Ho. 180 Figure 6.3 140 170 Go. 130 Go. 160 150 120 Po. 140 110 130 100 Pan 120 Po. Pan Ho. mm mm 90 110 100 80 Hy. Hy. 70 90 80 60 70 50 60

1 2 3 4 5 6 1 2 3 4 5 6 Hominoidea: Cranial Height (bas-br) Hominoidea: Inferior Cranial Length (bas-nas)

Figure 6.4 180 Figure 6.5 200 Po. 170 190 160 Go. 180 Pan Go. 150 170 140 Po. 160 130 Ho. 150 120 140 110 130 mm Pan

mm 100 120 Ho. 90 110 Hy. 80 Hy 100 70 . 90 60 80 50 70 1 2 3 4 5 6 1 2 3 4 5 6 Hominoidea: Biasterionic Breadth (biast) Hominoidea: Superior Facial Length (bas-pros)

Figure 6.6 Figure 6.7 Go. 90 80 Ho. Po. 80 Po. Go. Ho. 70 Pan 70 Pan 60 60 mm mm 50 Hy. 50 Hy. 40 40

30 30

1 2 3 4 5 6 1 2 3 4 5 6 Hominoidea: Maxillo-alveolar Breadth (biecm) Hominoidea: Interentoglenoid Breadth (bien)

289 Figure 6.8 Figure 6.9 Go. Go. 150 150 Po. 140 140 130 Po. Pan Ho. 130 Pan Ho. 120 120 110 110 100 100 mm mm 90 Hy. 90 80 80 Hy. 70 70 60 60 50 50 1 2 3 4 5 6 1 2 3 4 5 6 Hominoidea: Bifrontomalartemporale Breadth (bifmt) Hominoidea: Biporionic Breadth (bipor)

Figure 6.10 Figure 6.11 Go. 200 140 Po. 190 Po. Go. 130 180 170 120 Ho. 160 110 150 Pan Ho. 100 Pan 140 130

mm 90 mm 80 120 Hy. 110 70 100 Hy. 60 90 50 80 40 70 1 2 3 4 5 6 1 2 3 4 5 6 Hominoidea: Bizygomaxillare Inferior Breadth (bizi) Hominoidea: Bizygomatic Breadth (bizygo)

Figure 6.13 170 160 130 Figure 6.12 Go. 150 120 Ho. 140 Go. 110 130 Po. 100 Po. 120 90 110 Pan 80 100

mm 90 70 Pan mm 80 Ho. 60 70 50 Hy. 60 Hy. 40 50 30 40 20 30

1 2 3 4 5 6 1 2 3 4 5 6 Hominoidea: Occipital Sagittal Chord (lam-opn) Hominoidea: Superior Facial Height (nas-pros)

290 Figure 6.14 Figure 6.15 Go. 110 130 Go. 120 100 110 Po. 90 100 Pan 80 Po. 90 70 Pan 80 Ho. mm 70 Hy mm 60 . Ho. 60 50 Hy. 50 40 40 30 30

1 2 3 4 5 6 1 2 3 4 5 6 Hominoidea: Palatal Length A (ol-sta) Hominoidea: Maximum Length of the Zygomatic (zs-zgyi)

Figure 6.16 70 Go.

60 Po. 50 Pan Ho. 40 mm

30 Hy.

1 2 3 4 5 6 Hominoidea: Zygomatico-maxillary Suture Length (zs-zi)

One cranial variable which did exhibit a relationship to an increase in body size was biasterionic breadth (Figure 6.5, biast). The lack of variables which demonstrate clear scaling relationships with increasing body size is actually due to Homo, which does not correspond to the other apes because of its absolutely and relatively larger neurocranium and smaller viscerocranium dimensions.

6.2.1 Shapiro-Wilk results for cranial variables: The majority of cranial variables were found to have a normal distribution. However, there are some exceptions for Gorilla and Hylobates. Table 6.7 lists the results of the Shapiro-Wilk analysis of particular cranial measurements. Four cranial dimensions of Gorilla were not normally distributed. This is due to the extreme sexual dimorphism of Gorilla. These included cranial height (bas-br), biasterionic breath (biast), superior facial breadth (bifmt) and the occipital sagittal chord (lam-opn).

291 Table 6.7: Shapiro-Wilk results for cranial variables of the Hominoidea; those in bold represent significant departures from normality (variables listed in alphabetical order by abbreviation).

Hy. Pan Po. Ho. Go. n=31 n=33 n=26 n=20 n=34 1. Bas-br W: 0.98 0.98 0.95 0.92 0.93 p(normal): 0.73 0.69 0.24 0.11 0.04 2. Bas-nas W: 0.92 0.97 0.96 0.97 0.95

p(normal): 0.02 0.6 0.31 0.67 0.18 3. Bas-pros W: 0.90 0.96 0.95 0.93 0.96 p(normal): 0.07 0.32 0.29 0.18 0.32 4. Biast W: 0.95 0.98 0.96 0.95 0.93 p(normal): 0.14 0.80 0.40 0.40 0.04 5. Biecm W: 0.94 0.98 0.93 0.91 0.95 p(normal): 0.08 0.84 0.88 0.08 0.11 6. Bien W: 0.98 0.95 0.98 0.98 0.98 p(normal): 0.85 0.13 0.87 0.90 0.75 7. Bifmt W: 0.98 0.98 0.94 0.96 0.92 p(normal): 0.86 0.85 0.11 0.46 0.01

8. Bipor W: 0.93 0.98 0.95 0.96 0.95 p(normal): 0.04 0.82 0.22 0.49 0.17 9. Bizi W: 0.9 0.98 0.98 0.97 0.96 p(normal): 0.009 0.8 0.94 0.66 0.22 10. Bizygo W: 0.94 0.96 0.94 0.98 0.92 p(normal): 0.1 0.25 0.12 0.94 0.02 11. Lam-opn W: 0.93 0.99 0.95 0.92 0.91 p(normal): 0.06 0.99 0.21 0.12 0.009 12. Nas-pros W: 0.95 0.98 0.98 0.97 0.97 p(normal): 0.17 0.8 0.86 0.74 0.45 13. Ol-sta W: 0.89 0.91 0.96 0.96 0.95 p(normal): 0.005 0.01 0.42 0.48 0.15 14. Zs-zgyi W: 0.93 0.94 0.98 0.97 0.96 p(normal): 0.04 0.08 0.93 0.76 0.28

15. Zs-zi W: 0.96 0.95 0.97 0.99 0.97 p(normal): 0.29 0.15 0.6 0.99 0.6

Surprisingly, five cranial variables for Hylobates were also not normally distributed. These included inferior cranial length (bas-nas), biporionic breadth (bipor), bizygomaxillare inferior breadth (bizi), palatal length A (ol-sta) and maximum length of the zygomatic (zs-zgyi). However, unlike Gorilla, the cause of these non-normally distributed variables is not sexual dimorphism. Hylobates species are not sexually dimorphic like genera such as Gorilla, Pongo, Papio or Mandrillus. Instead, these variables are not normally distributed because of the size differences between the smaller gibbons (H. Hylobates, H. Hoolock and H. Nomascus) and larger siamangs (H. Symphalangus). Thus, when applying one-way ANOVA to these particular cranial dimensions, Gorilla and Hylobates were excluded. Lastly, apart from palatal length A 292 for Pan, which was not normally distributed, all other variables for the chimpanzees, Pongo and Homo are normally distributed.

6.2.2 Kruskal-Wallis and Mann-Whitney results for cranial variables: As the box-plots indicated, many cranial variables were significantly different between genera while some are remarkably similar. Table 6.8 lists the results of applying Kruskal-Wallis to several hominoid cranial variables. Box-plots illustrated the polarity in size between Hylobates and Gorilla. All cranial variables for Hylobates subjected to Kruskal-Wallis were significantly different from the other hominoids. Likewise, most cranial variables for Gorilla were also significantly different except for two dimensions. The interentoglenoid breadth (bien, Figure 6.7) and bizygomaxillare inferior breadth (bizi, Figure 6.10) of Gorilla overlapped with that of Pongo and were thus not significantly different (although the latter approaches significance, p-value 0.08). In addition, the superior facial length (bas-pros, Figure 6.4) of Pan and Gorilla is very similar and is not significantly different. Furthermore, most cranial variables between Pan and Pongo were significantly different but three measurements were not. The inferior cranial length (bas-nas), superior facial breadth (bifmt) and maximum length of the zygomatic (zs-zgyi) for Pan, Pongo and Homo overlapped and were thus not significantly different. Lastly, most cranial dimensions of Homo were also significantly different from the other hominoids but seven variables are very similar to Pan and Pongo. For example, the inferior cranial length (bas-nas), biasterionic breadth (biast), superior facial breadth (bifmt), and biporionic breadth (bipor) were not significantly different between the Pongo and Homo samples (bifmt was also not significantly different between Pan and Homo). In addition, the bizygomatic breadth (bizygo) and zygomatico-maxillary suture length (zs-zi) were not significantly different for the Pan and Homo samples.

6.2.3 One-way ANOVA results for cranial variables: Similar to the results of Kruskal-Wallis, one-way ANOVA also revealed many significant differences but with greater statistical power more similarities were apparent. Table 6.9 lists the results of one-way ANOVA. For most cranial variables, the greater majority (>70%) of the variation was between groups, not within them. However, this statement was not true for biporionic (bipor), bizygomaxillare inferior (bizi) breadths and the maximum length of the zygomatic (zs-zgyi). The inferior cranial length

293 Table 6.8: Kruskal-Wallis results for cranial variables of the Hominoidea with Mann-Whitney pairwise comparisons (p (same)) (variables listed in alphabetical order by abbreviation).

Hy. Pan Po. Ho. Go. n=31 n=26 n=33 n=20 n=34 1. Bas-br 0 6.67E-12 1.13E-10 2.36E-09 4.68E-12 Hy. H: 125.3 0 1.16E-06 1.48E-09 1.64E-11 Pan Hc: 125.3 0 8.92E-09 0.0003 Po. p(same): 4.00E-26 0 6.85E-09 Ho. 0 Go. 2. Bas-nas 0 6.67E-12 1.13E-10 2.36E-09 4.68E-12 Hy. H: 116.6 0 0.11 0.03 4.63E-12 Pan Hc: 116.6 0 0.76 9.07E-11 Po. p(same): 2.81E-24 0 1.19E-09 Ho. 0 Go. 3. Bas-pros 0 6.67E-12 1.13E-10 0.002 6.67E-12 Hy. H: 109.1 0 0.00002 1.48E-09 0.99 Pan Hc: 109.1 0 8.92E-09 0.00002 Po. p(same): 1.13E-22 0 1.48E-09 Ho. 0 Go. 4. Biast 0 6.67E-12 1.13E-10 2.36E-09 4.68E-12 Hy. H: 114.5 0 1.93E-08 1.66E-09 3.10E-12 Pan Hc: 114.5 0 0.36 0.008 Po. p(same): 8.08E-24 0 0.04 Ho. 0 Go. 5. Biecm 0 6.67E-12 1.13E-10 2.36E-09 4.68E-12 Hy. H: 113.2 0 4.17E-07 0.0001 3.55E-12 Pan Hc: 113.2 0 0.06 0.007 Po. p(same): 1.49E-23 0 1.78E-07 Ho. 0 Go. 6. Bien 0 6.67E-12 1.13E-10 2.36E-09 4.68E-12 Hy. H: 109 0 1.38E-07 1.48E-09 3.55E-08 Pan Hc: 109 0 0.0002 0.47 Po. p(same): 1.19E-22 0 0.003 Ho. 0 Go. 7. Bifmt 0 6.67E-12 1.13E-10 2.36E-09 4.68E-12 Hy. H: 107.6 0 0.10 0.27 1.80E-10 Pan Hc: 107.6 0 0.51 6.13E-08 Po. p(same): 2.35E-22 0 1.05E-08 Ho. 0 Go.

294

Hy. Pan Po. Ho. Go. n=31 n=33 n=26 n=20 n=34 8. Bipor 0 6.67E-12 1.13E-10 2.36E-09 4.68E-12 Hy. H: 110 0 0.001 0.008 8.57E-12 Pan Hc: 110 0 0.47 6.63E-06 Po. p(same): 7.23E-23 0 8.95E-08 Ho. 0 Go. 9. Bizi 0 6.67E-12 1.13E-10 2.36E-09 4.68E-12 Hy. H: 121.8 0 5.43E-10 1.60E-06 2.08E-12 Pan Hc: 121.8 0 0.00006 0.08 Po. p(same): 2.19E-25 0 2.81E-08 Ho. 0 Go. 10. Bizygo 0 6.67E-12 1.13E-10 2.36E-09 4.68E-12 Hy. H: 116.4 0 1.03E-06 0.29 3.88E-12 Pan Hc: 116.4 0 4.27E-06 0.01 Po. p(same): 3.19E-24 0 1.49E-09 Ho. 0 Go. 11. Lam-opn 0 6.67E-12 1.13E-10 2.36E-09 4.68E-12 Hy. H: 121.9 0 4.41E-08 1.48E-09 3.88E-12 Pan Hc: 121.9 0 2.85E-08 0.001 Po. p(same): 2.15E-25 0 0.003 Ho. 0 Go. 12. Nas-pros 0 6.67E-12 1.13E-10 2.36E-09 4.68E-12 Hy. H: 121.5 0 0.002 2.92E-09 5.65E-11 Pan Hc: 121.5 0 1.71E-08 0.0002 Po. p(same): 2.60E-25 0 1.19E-09 Ho. 0 Go. 13. Ol-sta 0 8.83E-12 1.13E-10 0.03 4.68E-12 Hy. H: 121.3 0 0.0002 1.48E-09 2.42E-11 Pan Hc: 121.3 0 8.92E-09 2.92E-06 Po. p(same): 2.87E-25 0 1.19E-09 Ho. 0 Go. 14. Zs-zgyi 0 6.67E-12 4.36E-10 4.76E-09 4.68E-12 Hy. H: 113.8 0 0.91 0.002 1.32E-11 Pan Hc: 113.8 0 0.03 3.18E-09 Po. p(same): 1.13E-23 0 1.19E-09 Ho. 0 Go. 15. Zs-zi 0 6.67E-12 1.13E-10 3.77E-09 4.68E-12 Hy. H: 116.3 0 0.00004 0.20 3.55E-12 Pan Hc: 116.3 0 0.00002 0.00006 Po. p(same): 3.24E-24 0 1.49E-09 Ho. 0 Go.

295 Table 6.9: One-Way ANOVA results for cranial variables of the Hominoidea (variables listed in alphabetical order by abbreviation). Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 1. Bas-br p(same): Hy. Pan Po. Ho. Between F: 806.7 0 0.0001 0.0001 0.0001 groups: 73898.9 96.53 3 24633 981.9 3.71E-77 0.007 df: 49.66 0 0.0001 0.0001 Within groups: 2659.13 3.47 106 25.09 p: 4.20E-42 0 0.0001 Total: 76558 109 ex. Go. 0 2. Bas-nas Pan Po. Ho. Go. Between F: 59.88 0 0.66 0.36 0.0001 groups: 21505 72.59 3 7168.33 96.22 1.67E-30 7.58E-06 df: 57.47 0 0.96 0.0001 Within groups: 8120.18 27.41 109 74.5 p: 1.10E-17 0 0.0001 Total: 29625.2 112 ex. Hy. 0 3. Bas-pros Pan Po. Ho. Go. Between F: 187.8 0 0.0001 0.0001 1 groups: 39313.7 72.85 3 13104.6 96.6 1.89E-30 0.00002 df: 57.78 0 0.0001 0.0001 Within groups: 14651.1 27.15 108 135.66 p: 9.34E-30 0 0.0001 Total: 53964.8 111 ex. Hy 0 4. Biast Hy. Pan Po. Ho. Between F: 448.9 0 0.0001 0.0001 0.0001 groups: 43711.11 85.74 3 14570.4 212.4 1.15E-44 3.79E-09 df: 51.32 0 0.0001 0.0001 Within groups: 7272.75 14.26 106 68.61 p: 8.44E-37 0 0.94 Total: 50983.8 109 ex. Go. 0

296 Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. F comparisons sqrs: Var.: df: square: F: p(same): of Var. test: p(same): 5. Biecm p(same): Hy. Pan Po. Ho. Go. Between F: 259.7 0 0.00002 0.00002 0.00002 0.00002 Hy. groups: 20492 89.04 4 5123 282.3 1.17E-65 0.0005 df: 63.96 0 0.00002 0.00002 0.00002 Pan Within groups: 2522.52 10.96 139 18.15 p: 8.85E-39 0 0.04 0.002 Po. Total: 23014.5 143 0 0.00001 Ho. 0 Go. 6. Bien Hy. Pan Po. Ho. Go. Between F: 274.4 0 0.00002 0.00002 0.00002 0.00002 Hy. groups: 21296.5 85.06 4 5324.13 197.8 2.52E-56 0.01 df: 63.76 0 0.00002 0.00002 0.00002 Pan Within groups: 3741.22 14.94 139 26.92 p: 2.04E-39 0 0.00002 0.88 Po. Total: 25037.7 143 0 0.0003 Ho. 0 Go. 7. Bifmt Hy. Pan Po. Ho. Between F: 328.4 0 0.0001 0.0001 0.0001 Hy. groups: 37825.8 86.30 3 12608.6 222.7 1.33E-45 0.00008 df: 52.04 0 0.48 0.07 Pan Within groups: 6002.58 13.70 106 56.63 p: 8.72E-34 0 0.98 Po. Total: 43828.4 109 ex. Go. 0 Ho. 8. Bipor Pan Po. Ho. Go. Between F: 43.83 0 0.003 0.10 0.0001 Pan groups: 10820.9 54.03 3 3606.96 42.71 2.50E-18 0.001 df: 54.69 0 0.50 0.0001 Po. Within groups: 9206.25 45.97 109 84.46 p: 1.43E-14 0 0.0001 Ho. Total: 20027.1 112 ex. Hy. 0 Go.

297

Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 9. Bizi p(same): Pan Po. Ho. Go. Between F: 95.66 0 0.0001 0.0001 0.0001 Pan groups: 18799 66.55 3 6266.34 72.29 8.25E-26 2.13E-07 df: 50.52 0 0.0001 0.08 Po. Within groups: 9448.77 33.45 109 86.69 p: 7.79E-21 0 0.0001 Ho. Total: 28247.8 112 ex. Hy. 0 Go. 10. Bizygo Hy. Pan Po. Ho. Between F: 244.4 0 0.0001 0.0001 0.0001 Hy. groups: 84327.5 84.59 3 28109.2 193.9 6.95E-43 3.37E-10 df: 51.85 0 0.0001 0.77 Pan Within groups: 15367.4 15.41 106 144.98 p: 1.43E-30 0 0.0001 Po. Total: 99694.9 109 ex. Go. 0 Ho. 11. Lam-opn Hy. Pan Po. Ho. Between F: 395 0 0.0001 0.0001 0.0001 Hy. groups: 55846.9 92.06 3 18615.6 409.7 3.89E-58 0.0002 df: 48.51 0 0.0001 0.0001 Pan Within groups: 4816.61 7.94 106 45.44 p: 4.57E-34 0 0.0001 Po. Total: 60663.5 109 ex. Go. 0 Ho. 12. Nas-pros Hy. Pan Po. Ho. Go. Between F: 375.1 0 0.00002 0.00002 0.00002 0.00002 Hy. groups: 101597 88.13 4 25399.3 258 2.95E-63 1.29E-06 df: 65.75 0 0.0007 0.00002 0.00002 Pan Within groups: 13684.4 11.87 139 98.45 p: 1.76E-44 0 0.00002 0.00002 Po. Total: 115282 143 0 0.00002 Ho. 0 Go.

298

Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 13. Ol-sta p(same): Po. Ho. Go. Between F: 263.6 0 0.0001 0.0001 Po. groups: 30622.7 78.11 2 15311.3 137.4 3.99E-26 0.00004 df: 48.79 0 0.0001 Ho. Within groups: 8583.01 21.89 77 111.47 p: 7.03E-27 ex. Hy. & Pan 0 Go. Total: 39205.7 79 14. Zs-zgyi Pan Po. Ho. Go. Between F: 52.66 0 0.99 0.09 0.0001 Pan groups: 10775.6 64.56 3 3591.88 66.18 1.90E-24 0.00002 df: 56.17 0 0.07 0.0001 Po. Within groups: 5915.15 35.44 109 54.27 p: 2.49E-16 0 0.0001 Ho. Total: 16691.1 112 ex. Hy. 0 Go. 15. Zs-zi Hy. Pan Po. Ho. Go. Between F: 159.3 0 0.00002 0.00002 0.00002 0.00002 Hy. groups: 14792.3 79.25 4 3698.08 132.7 1.94E-46 5.44E-07 df: 63.64 0 0.00002 0.85 0.00002 Pan Within groups: 3874.2 20.75 139 27.87 p: 2.12E-32 0 0.00002 0.00002 Po. Total: 18666.5 143 0 0.00002 Ho. 0 Go.

299 (bas-nas), biporionic breadth (bipor) and maximum length of the zygomatic bone (zs- zgyi) for Pan, Pongo and Homo were not significantly different between each other but all were significantly different from Gorilla (Figures 6.3 and 6.15). The biasterionic breadth (biast) of Pongo and Homo is remarkably similar (Figure 6.5) however their respective surface morphologies and dispositions are very different.

6.2.4 Summary for cranial variables: To summarize, cranial variables of the Hominoidea revealed, 1) Hylobates and Gorilla measurements easily separated these genera from the rest due to extreme differences in size; 2) there is size overlap for some cranial variables for all Great apes (including humans); and 3) modern Homo crania are remarkably distinct and has changed dramatically in comparison to the other apes.

6.3 Cranial Indices: To correct for absolute differences in size many cranial indices were examined. Table 6.10 lists the mean and standard deviation (SD) for several cranial indices. Figures 6.17 to 6.26 are box-plots which display interesting scaling trends and distinguishing proportions between the hominoid genera. Again, Hylobates and Homo were clearly distinct but three indices revealed proportional similarities. These two genera overlapped in the relative proportion of the breadth of the lower face (bizi) to the superior facial height (nas-pros); the superior facial height (nas-pros) to the interentoglenoid breadth (bien); and the superior facial height (nas-pros) to the cranial vault length (g-o) (Figures 6.22, 6.24 and 6.25). There were also proportions which the African apes, Gorilla and Pan, share to the exclusion of the Asian ape, Pongo. In fact, two of them included the proportions which showed overlap with Hylobates and Homo. In Figures 6.18, 6.22 and 6.25 Pan and Gorilla overlap, whereas Pongo does not. Unlike the other apes, the upper facial breadth of Pongo was relatively less than the lower facial breadth (Figure 6.20).

300 Table 6.10: Generic means for cranial indices (%; indices listed in alphabetical order by abbreviation).

Hy. Pan Po. Ho. Go. 1. bas-br/bas-nas 81.54 88.71 100.14 135.00 84.45 SD 6.19 3.98 4.98 7.11 4.84 2. bas-br/bas-pros 66.51 66.93 64.58 137.47 61.49 SD 6.66 4.13 5.11 8.45 4.06 3. bas-br/biast 99.11 109.11 96.56 125.56 95.71 SD 7.54 7.37 11.18 8.21 8.45 4. bas-br/biaur 81.36 79.28 81.78 111.39 81.50 SD 6.61 3.76 5.15 6.16 7.10 5. bas-br/g-o 63.9 66.7 78.9 75.6 61.8 SD 3.7 2.5 4.5 3.9 4.0 6. bas-br/bizygo 73.91 71.77 66.83 108.12 67.87 SD 6.22 3.67 6.06 8.42 4.48 7. bas-br/pros-o 49.42 47.41 47.69 66.46 42.63 SD 3.65 2.14 3.34 3.13 3.74 8. bas-nas/bas-pros 81.4 75.5 64.5 101.8 72.9 SD 3.0 4.3 3.7 3.5 3.6 9. bas-pros/pros-o 74.54 70.95 73.94 48.43 69.38 SD 3.26 2.54 3.08 2.18 4.78 10. biast/biaur 82.38 72.87 85.59 88.86 85.47 SD 7.36 4.35 9.28 4.18 7.18 11. biast/g-o 64.7 61.0 83.1 60.3 65.5 SD 4.9 3.9 8.4 2.8 6.4 12. biast/lam-opn 182.0 150.4 159.6 109.7 143.4 SD 16.7 12.0 13.9 9.8 19.6 13. biaur/bizygo 90.89 90.60 81.65 97.20 83.49 SD 3.57 3.89 3.86 7.38 3.83 14. biaur/pros-o 60.86 59.87 58.33 59.83 52.49 SD 3.38 2.69 2.49 4.18 4.49 15. bicanex/bas-pros 38.94 41.70 41.49 41.98 37.48 SD 3.01 2.07 2.28 2.77 2.35 16. bicanex/nas-pros 94.83 67.38 68.82 62.33 62.53 SD 10.72 5.08 7.44 5.62 8.92 17. bicanex/ol-sta 77.41 83.42 83.27 88.16 70.65 SD 7.65 6.02 5.22 7.79 5.54 18. biecm/bas-br 67.65 64.89 66.89 47.74 64.74 SD 6.50 3.53 4.89 2.72 5.37 19. biecm/bas-nas 54.89 57.52 66.86 64.39 54.54 SD 3.82 3.26 4.23 4.05 3.85 20. biecm/biast 67.02 70.70 63.44 59.85 61.93 SD 7.99 4.65 6.33 3.83 7.03 21. biecm/bicanex 115.00 104.14 103.98 156.36 106.18 SD 7.02 6.64 7.63 8.36 8.44 22. biecm/bifmt 60.75 55.91 65.39 61.58 56.28 SD 7.20 3.04 4.02 3.73 4.52 23. biecm/bizi 68.91 66.69 59.14 64.45 59.66 SD 4.46 3.40 4.90 2.81 4.36

301 Hy. Pan Po. Ho. Go. 24. biecm/bizygo 49.74 46.54 44.53 51.49 43.82 SD 3.74 2.81 3.20 3.14 3.17 25. biecm/g-o 43.05 43.23 52.62 36.03 40.28 SD 3.08 2.42 3.13 2.12 3.60 26. biecm/ol-sta 88.91 86.69 86.35 137.55 75.01 SD 9.19 5.96 5.36 10.82 8.29 27. biecm/pros-o 33.26 30.73 31.77 31.70 27.52 SD 2.13 1.46 1.38 1.81 2.49 28. biecm/zs-zgyi 95.95 103.10 115.97 121.05 92.83 SD 8.24 8.14 9.33 7.62 9.52 29. bien/bas-br 67.83 62.89 66.85 54.58 61.43 SD 5.54 4.08 6.01 4.46 7.53 30. bien/bas-nas 55.10 55.79 66.84 73.68 51.68 SD 3.80 4.45 5.55 7.09 5.32 31. bien/bas-pros 44.90 42.06 43.12 75.04 37.66 SD 3.77 3.44 4.56 7.64 4.40 32. bien/biast 67.06 68.60 64.34 68.42 58.61 SD 5.67 6.28 7.89 5.92 7.59 33. bifmt/biaur 90.79 92.05 83.56 86.44 93.57 SD 7.58 4.88 4.44 5.81 5.49 34. bifmt/bizi 114.42 119.40 90.59 104.99 106.22 SD 10.82 4.98 7.21 7.41 5.66 35. bifmt/bizygo 82.38 83.29 68.17 83.68 78.01 SD 5.93 3.89 3.85 3.45 3.85 36. bifmt/g-o 71.36 77.42 80.74 58.61 71.71 SD 5.56 4.14 6.65 3.31 5.44 37. bifmt/nas-pros 178.46 125.49 109.35 158.53 117.64 SD 20.54 10.07 11.65 16.81 13.27 38. bifmt/pros-o 55.18 55.05 48.71 51.59 48.99 SD 4.71 2.85 2.84 3.47 3.71 39. bipor/g-o 72.6 77.4 90.2 63.1 73.6 SD 2.9 4.1 5.9 4.9 5.7 40. bizi/bas-br 98.44 97.40 113.56 74.12 108.64 SD 10.12 4.59 9.16 3.66 6.68 41. bizi/biast 97.55 106.26 109.20 92.96 103.81 SD 12.59 8.47 11.91 6.16 9.48 42. bizi/bizygo 72.25 69.86 75.51 80.03 73.53 SD 4.21 4.07 4.98 5.92 3.54 43. bizi/nas-pros 156.6 105.2 121.1 152.4 108.7 SD 14.4 8.6 12.9 8.3 8.5 44. bizs/zs-zgyi 104.67 92.76 110.79 107.90 78.06 SD 14.81 14.29 15.53 13.96 16.20 45. bizs/zs-zi 205.18 161.20 158.16 180.78 122.54 SD 33.43 29.28 27.40 34.07 26.02 46. bizs/bizi 74.41 59.88 55.77 57.40 49.80 SD 8.06 7.53 6.16 6.43 7.79

302 Hy. Pan Po. Ho. Go. 47. bizygo/pros-o 67.01 66.12 71.58 61.71 62.85 SD 3.61 2.43 4.31 4.14 4.38 48. br-lam/g-o 39.2 46.0 49.5 62.2 44.0 SD 7.1 3.8 4.0 2.9 4.8 49. g-o/biaur 127.38 119.03 103.85 147.65 131.05 SD 7.40 5.76 6.15 8.96 10.88 50. g-o/bizygo 115.73 107.74 84.91 143.11 109.19 SD 7.38 5.28 7.89 8.49 7.55 51. g-o/pros-o 77.34 71.17 60.52 88.01 68.45 SD 2.96 2.81 3.45 2.45 4.44 52. iv-pms/bicanex 56.55 63.03 60.51 68.42 72.71 SD 7.10 7.29 6.78 9.75 10.14 53. iv-pms/ol-sta 43.85 52.36 50.24 60.06 51.12 SD 7.77 5.38 5.09 8.60 6.46 54. lam-opn/bas-br 55.38 61.13 67.50 73.83 74.34 SD 6.44 4.12 6.33 5.75 9.18 55. lam-opn/bas-pros 36.89 40.88 43.42 101.62 45.62 SD 6.22 3.42 3.52 11.51 5.70 56. lam-opn/biaur 45.13 48.45 55.10 82.20 60.72 SD 6.86 3.82 5.13 7.46 9.96 57. lam-opn/g-o 35.7 40.7 52.2 55.3 46.3 SD 3.2 3.1 4.2 4.7 6.2 58. maxnawi/nas-rhi 135.5 84.1 85.1 121.3 69.3 SD 24.8 13.2 14.9 14.8 10.1 59. nas-br/g-o 69.4 56.1 55.8 62.7 52.5 SD 6.3 3.0 4.4 2.2 3.2 60. nas-pros/biaur 50.70 73.70 77.21 54.91 80.38 SD 4.82 5.72 8.65 5.01 8.92 61. nas-pros/bien 92.58 148.37 142.90 90.85 162.21 SD 9.97 12.78 23.10 9.40 17.34 62. nas-pros/g-o 39.94 62.02 74.59 37.30 61.60 SD 4.57 5.30 9.42 3.80 7.38 63. nas-rhi/nas-pros 32.90 38.68 36.39 33.98 47.29 SD 5.21 4.43 4.52 3.50 4.34 64. ol-sta/bas-br 76.83 75.16 77.61 34.86 86.91 SD 10.73 6.08 5.56 2.79 8.14 65. ol-sta/bas-nas 62.18 66.64 77.61 46.94 73.20 SD 5.94 5.80 5.42 2.84 5.74 66. ol-sta/bas-pros 50.52 50.15 49.89 47.75 53.18 SD 3.60 3.07 2.14 2.35 2.73 67. ol-sta/bizygo 56.33 53.87 51.66 37.56 58.79 SD 5.35 4.25 3.73 2.61 4.46 68. palhei/ol-sta 17.73 17.46 19.87 21.13 18.40 SD 3.15 2.51 3.62 4.48 3.42 69. zs-zgyi/bizi 71.61 64.99 50.70 53.44 64.64 SD 5.63 5.09 4.15 4.10 5.24

303 Hy. Pan Po. Ho. Go. 70. zs-zgyi/bizygo 51.63 45.28 38.17 42.58 47.45 SD 3.68 2.84 2.63 1.90 3.40 71. zs-zgyi/ol-sta 92.52 84.48 74.15 113.77 81.04 SD 11.49 7.85 6.31 8.08 7.46 72. zs-zi/bizi 36.69 37.63 35.77 32.25 41.22 SD 3.33 3.51 3.97 2.97 3.52 73. zs-zi/bizygo 25.88 25.44 25.66 24.41 29.40 SD 5.73 5.27 6.45 6.79 6.11 74. zs-zi/zs-zgyi 51.40 58.06 70.61 60.68 63.97 SD 4.74 5.22 6.15 7.21 5.35

304 Box Plots for cranial indices of the Hominoidea (Hy. – Hylobates spp., n=31; Pan spp., n=33; Po. – Pongo pygmaeus, n=26; Ho. – Homo sapiens, n=20 & Go. – Gorilla gorilla, n=34; indices in alphabetical order by abbreviation):

Figure 6.18 Figure 6.17 110 Ho. 230 Hy. 220 100 210 200 Po. 90 Hy. 190 Pan Go. Pan 180

% Go. 170 80 160 Po. % 150 70 140 Ho. 130 60 120 110 100 1 2 3 4 5 6 90 Hominoidea: bas-nas/bas-pros x 100 80 1 2 3 4 5 6 Hominoidea: biast/lam-opn x 100 Figure 6.19 Figure 6.20 Ho. 140 Po. Hy. 140 Pan 130 Pan 130 120 Go. Hy. Go. 120 Ho. 110 % 110 Po. 100 % 100 90 80 90 70 80 1 2 3 4 5 6 Hominoidea: biecm/zs-zgyi x 100 1 2 3 4 5 6 Hominoidea: bifmt/bizi x 100 Figure 6.21 Po. Figure 6.22 200 100 190 Hy. 180 Ho. 90 Pan 170 Go. 160 80 Hy. 150 Po. % Ho. 140 % Go. 130 Pan 70 120 110 60 100 90 80 1 2 3 4 5 6 1 2 3 4 5 6 Hominoidea: bipor/g-o x 100 Hominoidea: bizi/nas-pros x 100

305 Figure 6.23 70 210 Figure 6.24 Ho. Go. 200 Po. 190 Pan Po. 180 60 Go. 170 160 150

% 50 140

Pan % Hy. 130 120 Hy. Ho. 110 40 100 90 80 70 1 2 3 4 5 6 Hominoidea: lam-opn/g-o x 100 1 2 3 4 5 6 Hominoidea: nas-pros/bien x 100

Figure 6.25 Figure 6.26 100 80 Po. Ho. Po. 90 Go. Pan 80 Pan Go. 70 70 Hy.

% 60 % 60 Hy. 50 Ho. 40 50 30

1 2 3 4 5 6 1 2 3 4 5 6 Hominoidea: nas-pros/g-o x 100 Hominoidea: zs-zi/zs-zgyi x 100

6.3.1 Shapiro-Wilk results for cranial indices: Table 6.11 provides the results of applying Shapiro-Wilk to several hominoid cranial indices. The relative contribution of the occipital sagittal chord (lam-opn) to the cranial vault length (g-o) for Hylobates was the only index not normally distributed. However, as mentioned previously, this is not due to sexual dimorphism but rather the difference in size (and proportions) between the smaller gibbons and larger siamangs.

6.3.2 Kruskal-Wallis and Mann-Whitney results for cranial indices: Table 6.12 provides the results of applying Kruskal-Wallis to several cranial indices and confirms the patterns seen from the box-plots. There were fewer generic pairwise comparisons which were not significantly different than those that were. In fact, there were two indices in which every comparison is statistically significant. These two indices included the inferior cranial length (bas-nas) in relation to the superior

306 Table 6.11: Shapiro-Wilk results for cranial indices of the Hominoidea; those in bold represent significant departures from normality (indices listed in alphabetical order by abbreviation).

Hy. Po. Pan Ho. Go. n=31 n=26 n=33 n=20 n=34 1.Bas-nas/bas-pros W: 0.96 0.99 0.95 0.93 0.95 p(normal): 0.38 0.97 0.2 0.18 0.16 2. Biast/lam-opn W: 0.98 0.99 0.94 0.96 0.98 p(normal): 0.74 0.99 0.17 0.55 0.68 3. Biecm/zs-zgyi W: 0.98 0.98 0.95 0.93 0.97 p(normal): 0.73 0.65 0.22 0.15 0.35 4. Bifmt/bizi W: 0.98 0.98 0.97 0.95 0.98 p(normal): 0.93 0.87 0.56 0.34 0.84 5. Bipor/g-o W: 0.98 0.98 0.98 0.97 0.97 p(normal): 0.83 0.81 0.88 0.82 0.47 6. Bizi/nas-pros W: 0.96 0.98 0.95 0.96 0.96 p(normal): 0.36 0.91 0.25 0.64 0.21 7. Lam-opn/g-o W: 0.86 0.96 0.96 0.93 0.95 p(normal): 0.001 0.21 0.39 0.16 0.1 8. Nas-pros/bien W: 0.96 0.94 0.96 0.96 0.98 p(normal): 0.21 0.06 0.47 0.5 0.63 9. Nas-pros/g-o W: 0.97 0.96 0.98 0.96 0.95 p(normal): 0.6 0.23 0.77 0.58 0.16 10. Zs-zi/zs-zgyi W: 0.98 0.99 0.98 0.94 0.95 p(normal): 0.7 0.99 0.88 0.1945 0.14

307 Table 6.12: Kruskal-Wallis results for cranial indices of the Hominoidea with Mann-Whitney pairwise comparisons (p (same)) (indices listed in alphabetical order by abbreviation).

Hy. Pan Po. Ho. Go. n=31 n=33 n=26 n=20 n=34 1. Bas-nas/bas-pros 0 5.24E-07 1.13E-10 2.36E-09 2.55E-10 Hy. H: 115.4 0 4.05E-10 1.48E-09 0.001 Pan Hc: 115.4 0 8.92E-09 3.32E-09 Po. p(same): 5.05E-24 0 1.19E-09 Ho. 0 Go. 2. Biast/lam-opn 0 4.74E-08 0.00003 2.65E-09 2.87E-08 Hy. H: 83.57 0 0.52 1.86E-09 0.10 Pan Hc: 83.58 0 8.92E-09 0.001 Po. p(same): 3.05E-17 0 1.33E-07 Ho. 0 Go. 3. Biecm/zs-zgyi 0 0.002 1.17E-08 2.36E-09 0.14 Hy. H: 86.48 0 4.01E-06 2.31E-08 0.00007 Pan Hc: 86.48 0 0.12 2.31E-09 Po. p(same): 7.35E-18 0 1.19E-09 Ho. 0 Go. 4. Bifmt/bizi 0 0.49 1.97E-09 0.003 0.0008 Hy. H: 91.09 0 6.06E-11 1.22E-08 6.68E-11 Pan Hc: 91.09 0 1.70E-06 2.53E-09 Po. p(same): 7.73E-19 0 0.71 Ho. 0 Go. 5. Bipor/g-o 0 6.58E-06 1.13E-10 8.62E-08 0.19 Hy. H: 98.25 0 5.70E-10 4.08E-09 0.005 Pan Hc: 98.25 0 8.92E-09 1.35E-10 Po. p(same): 2.32E-20 0 4.39E-07 Ho. 0 Go. 6. Bizi/nas-pros 0 6.67E-12 5.93E-10 0.35 4.68E-12 Hy. H: 106.3 0 8.88E-06 1.48E-09 0.16 Pan Hc: 106.3 0 1.60E-08 0.0004 Po. p(same): 4.48E-22 0 0.0004 Ho. 0 Go.

308 Hy. Pan Po. Ho. Go. n=31 n=33 n=26 n=20 n=34 7. Lam-opn/g-o 0 1.24E-06 1.91E-10 2.36E-09 1.15E-09 Hy. H: 100.8 0 3.02E-10 1.48E-09 0.0001 Pan Hc: 100.9 0 0.05 0.003 Po. p(same): 6.49E-21 0 0.00002 Ho. 0 Go. 8. Nas-pros/bien 0 6.67E-12 1.25E-10 0.64 4.68E-12 Hy. H: 106.3 0 0.27 1.48E-09 0.00006 Pan Hc: 106.3 0 1.02E-08 0.0006 Po. p(same): 4.44E-22 0 1.19E-09 Ho. 0 Go. 9. Nas-pros/g-o 0 1.40E-11 1.13E-10 0.02 6.67E-12 Hy. H: 110.1 0 2.71E-06 1.49E-09 0.9 Pan Hc: 110.1 0 8.92E-09 1.57E-06 Po. p(same): 6.97E-23 0 1.48E-09 Ho. 0 Go. 10. Zs-zi/zs-zgyi 0 9.29E-06 1.13E-10 0.00002 3.02E-10 Hy. H: 83.86 0 5.98E-10 0.35 0.00009 Pan Hc: 83.86 0 0.00003 0.00003 Po. p(same): 2.65E-17 0 0.06 Ho. 0 Go. facial length (bas-pros) along with the occipital sagittal chord (lam-opn) and its contribution to the cranial vault length (g-o) (see Figures 6.17 and 6.23).

6.3.3 One-way ANOVA results for cranial indices: Table 6.13 presents the results of applying one-way ANOVA to several cranial indices for the hominoids. There were four indices and their respective variation between groups was greater than 70%. The remaining indices produced variation percentages between groups ranging from >50-69%. Akin to the results for Kruskal- Wallis, there were far fewer generic pairwise comparisons which were not significantly different than those that were, thus confirming the results of the former. The relative proportion of the zygomatico-maxillary suture length (zs-zi) to the maximum length of the zygomatic (zs-zgyi) was not significantly different between Pan and Homo is not significantly different (Figure 6.26), while the superior facial height (nas-pros) in proportion to the cranial vault length (g-o) and the interentoglenoid breadth (bien) were also not significantly different between Hylobates and Homo (Figures 6.24, 6.25 and 6.26). In addition, the relative length of the zygomatico-maxillary suture (zs-zi) to the maximum length of the zygomatic (zs-zgyi), as well as, the proportional relationship between superior facial (bifmt) and bizygomaxillare inferior breadths (bizi) were not

309 Table 6.13: One-Way ANOVA results for cranial indices of the Hominoidea (indices listed in alphabetical order by abbreviation). Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 1. Bas-nas/bas-pros p(same): Hy. Pan Po. Ho. Go. Between F: 343.4 0 0.00002 0.00002 0.00002 0.00002 groups: 17586 95.45 4 4396.49 327.1 1.21E-69 0.45 df: 65.27 0 0.00002 0.00002 0.06 Within groups: 1868.24 10.14 139 13.44 p: 4.63E-43 0 0.00002 0.00002 Total: 18424.2 143 0 0.00002 0 2. Biast/lam-opn Hy. Pan Po. Ho. Go. Between F: 81.53 0 0.00002 0.00002 0.00002 0.00002 groups: 66108.1 63.73 4 16527 61.06 1.11E-29 0.005 df: 66.7 0 0.33 0.00002 0.52 Within groups: 37625.7 36.27 139 270.69 p: 5.97E-25 0 0.00002 0.005 Total: 103734 143 0 0.00002 0 3. Biecm/zs-zgyi Hy. Pan Po. Ho. Go. Between F: 53.15 0 0.02 0.00002 0.00002 0.67 groups: 15819.2 60.27 4 3954.79 52.71 5.93E-27 0.52 df: 65.74 0 0.00002 0.00002 0.0001 Within groups: 10429.3 39.73 139 75.03 p: 6.54E-20 0 0.19 0.00002 Total: 26248.5 143 0 0.00002 0

310 Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. of F comparisons sqrs: Var.: df: square: F: p(same): Var. test: p(same): 4. Bifmt/bizi p(same): Hy. Pan Po. Ho. Go. Between F: 81.07 0 0.09 0.00002 0.00004 0.0004 Hy. groups: 13733.6 64.06 4 3433.39 61.94 5.91E-30 0.0002 df: 62.92 0 0.00002 0.00002 0.00002 Pan Within groups: 7705.17 35.94 139 55.43 p: 4.08E-24 0 0.00002 0.00002 Po. Total: 21438.7 143 0 0.97 Ho. 0 Go. 5. Bipor/g-o Hy. Pan Po. Ho. Go. Between F: 77.78 0 0.002 0.00002 0.00002 0.94 Hy. groups: 9186.21 74.22 4 2296.55 100 6.46E-40 0.002 df: 62.81 0 0.00002 0.00002 0.03 Pan Within groups: 3191.58 25.78 139 22.96 p: 1.27E-23 0 0.00002 0.00002 Po. Total: 12377.8 143 0 0.00002 Ho. 0 Go. 6. Bizi/nas-pros Hy. Pan Po. Ho. Go. Between F: 159.3 0 0.00002 0.00002 0.61 0.00002 Hy. groups: 67971.9 80.53 4 16993 143.7 2.34E-48 0.001 df: 64.79 0 0.00002 0.00002 0.74 Pan Within groups: 16435.5 19.47 139 118.24 p: 9.15E-33 0 0.00002 0.0002 Po. Total: 84407.4 143 0 0.00002 Ho. 0 Go. 7. Lam-opn/g-o Pan Po. Ho. Go. Between F: 74.62 0 0.0001 0.0001 0.0001 Pan groups: 3339.87 57.82 3 1113.29 49.81 2.36E-20 0.0003 df: 53.37 0 0.08 0.0002 Po. Within groups: 2436.06 42.18 109 22.35 p: 4.29E-19 0 0.0001 Ho. Total: 5775.93 112 ex. Hy. 0 Go.

311 Levene’s test for Welch Tukey’s pairwise Sum of % of Mean Homo. F comparisons sqrs: Var.: df: square: F: p(same): of Var. test: p(same): 8. Nas-pros/bien p(same): Hy. Pan Po. Ho. Go. Between F: 211.5 0 0.00002 0.00002 0.99 0.00002 Hy. groups: 129090 80.36 4 32272.5 142.1 4.31E-48 0.00002 df: 65.73 0 0.66 0.00002 0.0009 Pan Within groups: 31557.4 19.64 139 227.03 p: 9.11E-37 0 0.00002 0.00002 Po. Total: 160647 143 0 0.00002 Ho. 0 Go. 9. Nas-pros/g-o Hy. Pan Po. Ho. Go. Between 0 0.00002 0.00002 0.55 0.00002 Hy. groups: 26241.4 81.94 4 6560.35 156.8 1.74E-50 0.00004 F: 194.6 0 0.00002 0.00002 0.99 Pan Within groups: 5815.6 18.16 139 41.84 df: 66.39 0 0.00002 0.00002 Po. Total: 32027 143 p: 6.75E-36 0 0.00002 Ho. 0 Go. 10. Zs-zi/zs-zgyi Hy. Pan Po. Ho. Go. Between F: 69.44 0 0.00004 0.00002 0.00002 0.00002 Hy. groups: 5521.96 58.87 4 1380.49 49.74 6.39E-26 0.01 df: 64.23 0 0.00002 0.35 0.0003 Pan Within groups: 3857.83 41.13 139 27.75 p: 1.28E-22 0 0.00002 0.0002 Po. Total: 9379.79 143 0 0.14 Ho. 0 Go.

312 significantly different between Gorilla and Homo are not statistically significant. Pan and Gorilla were very similar in three indices which describe the relative proportion of the occipital bone and facial dimensions (biast/lam-opn, bizi/nas-pros and nas-pros/g-o) and were not significantly different (Figures 6.18, 6.22 and 6.25).

6.3.4 Summary of cranial indices: In summary, akin to previous results for other catarrhines, levels of variation decrease for cranial indices, despite sexual dimorphism. Furthermore, Homo crania continued to be easily diagnosable for the other genera, although a few indices for Hylobates did overlap with those for humans. Additionally, some indices distinguish Pongo from the African apes .

6.4 Bivariate Results: Table 6.14 presents the results of plotting mean generic body against the generic mean for particular cranial variables. Several comparisons are statistically significant. Figure 6.27 is a bivariate plot of mean generic body weight plotted against biauriculare breadth (biaur). However, unlike the previous bivariate plots, interpretation of the hominoids was more difficult. Whereas earlier bivariate plots could be interpreted as the distribution of the genera across the landscape and their respective substrates, the hominoids do not but instead simply indicate size. Pan, Pongo and Homo were situated very close to one another around the linear regression line and yet all three occupy very different habitats with radically different forms of locomotion. Again, Hylobates and Gorilla were well segregated from the rest due to their extreme differences in size.

313 Table 6.14: Results of body weight (Ln (g)) plotted against the generic mean for particular cranial measurements (Ln (mm)) (cranial variables listed in alphabetical order by abbreviation; those in bold represent cranial variables with an adjusted R squared value >0.70 and a p-value <0.05).

P-value/ Mean Generic Body Weight (g) Adjusted Significance (Rowe, 1996; Fleagle, 1998): Ln (mm) R squared F bas-br 0.74 0.04 Hylobates spp. - 6729 bas-nas 0.92 0.006 Pan spp. - 43875 bas-pros 0.49 0.12 Pongo spp. - 56950 biast 0.93 0.005 Homo sapiens - 60000 biaur 0.97 0.001 Gorilla spp. - 124683 bicanex 0.65 0.10 biecm 0.96 0.002 bien 0.83 0.02 bienm 0.85 0.02 bifmo 0.91 0.007 bifmt 0.97 0.0001 bipor 0.99 0.0004 bizi 0.93 0.005 bizs 0.81 0.02 bizygo 0.91 0.007 br-lam 0.57 0.08 g-o 0.80 0.03 lam-opn 0.77 0.03 nas-br 0.40 0.15 nas-pros 0.84 0.02 nas-rhi 0.81 0.02 ol-sta 0.41 0.15 ol-pms 0.59 0.08 palhei 0.64 0.06 pros-o 0.99 0.0002 zs-zgyi 0.83 0.02 zs-zi 0.88 0.01

5.00 Figure 6.27: Adjusted R2: 0.97 4.90 Significance F & P-value 0.001 Po. 4.80 Go. y = 0.2455x + 2.2968 Pan 4.70 Ho. 4.60 4.50 4.40 4.30 Ln Biauriculare Breadth Ln Biauriculare Hy. 4.20 4.10 8.00 8.50 9.00 9.50 10.00 10.50 11.00 11.50 12.00 Ln Body Weight (g)

314 6.5 Multivariate Statistics and Morphological Distances: To consider and evaluate the entire multivariate datasets simultaneously PCA and CVA were utilized. 6.5.1 PCA results for Ln transformed data: To identify the underlying structure (i.e. the cranial variables which account for most of the variation between genera) of the Hominoidea (n=5) dataset based on Ln transformed generic means, PCA was employed. By doing so, PCA based on the variance-covariance matrix resulted in the first (Eigenvalue - 3.11; 84.24%), second (Eigenvalue - 0.49; 13.38%) and third (Eigenvalue - 0.06; 1.63%) PCs accounting for 99.25% of variation within the superfamily. One hundred percent of the variation is explained by the four PCs (Figure 6.28). Table 6.17 lists the variable loadings for the cranial measurements. The cranial dimensions receive low to moderate variable loadings. The variable loadings for the first PC were all positive and may be interpreted as size differences between genera; the remaining PCs had mixed polarities. Evidence for this can be seen in the mean PCA object scores per genus. For axis one, Gorilla, the largest living primate, received the highest mean PCA score of 24.96, while Hylobates, the smallest living apes, obtained the least - 20.26; Pan, Homo and Pongo had object scores in between these two extremes. The three highest variable loadings for PC 1 were nasal and facial length dimensions. These include the sagittal length of the nasal bones (nas-rhi), superior facial height (nas-pros) and nasal height (nas-ns) which procured variable loadings of 0.31, 0.25 and 0.22, respectively (Table 6.17). The variable loadings for PC 2 had mixed values. The positively loaded measurements are facial and palatal lengths whereas the negatively loaded measurements were neurocranial dimensions. The two highest positively loaded dimensions for PC 2 are the palatal and superior facial lengths (ol-sta and bas-pros), scoring 0.24 and 0.20. Working against these cranial variables are negatively loaded measurements such as the parietal and occipital sagittal chords (br-lam and lam-opn), cranial height (bas-br), frontal sagittal chord (nas-br) and cranial vault length (g-o), which achieve variable loadings of -0.43, -0.36, -0.27, -0.26 and -0.22, respectively. The variable loadings for PC 3 have mixed values. The positively loaded dimensions were palatal and facial breadths while the negatively loaded variables are a mix of facial and neurocranial breadth and length measurements. Positively loaded variables included palatal (ol-sta), interentoglenoid (bien) and bizygomaxillare superior

315 Figure 6.28: Hominoidea scatterplot of PC 1 & 2, PC 2 & 3 and PC 1 & 3 for Ln transformed generic means. Hy. - Hylobates; Po. - Pongo; Go. - Gorilla; Pan; and Ho. - Homo. Palatal & Palatal and Facial -2 0.3 lengths Interentoglenoid breadths 0.2 Po.

0.1

-3 Po. Hy. Pan 0 Go. PC 3: PC 2: 1.63% Ho. 13.38% Increase in body size -0.1 Pan

-0.2 -4 Hy. Parietal & Nasal & -0.3 Facial Occipital Palatal & Sagittal Chords Facial Ho. lengths -0.4 lengths Go.

20 21 22 23 24 25 -4 -3 -2 PC 1: 84.24% PC 2: 13.38% Parietal & Occipital Inferior breadth of the Nasal Bones Sagittal Chords

% of PCA PC Eigenvalue Var. Cum. %. scores Axis 1 Axis 2 Axis 3 1 3.11 84.24 84.24 Hy. 20.26 -3.08 -0.22 2 0.49 13.38 97.62 Po. 24.00 -2.95 0.23 3 0.06 1.63 99.25 Go. 24.96 -3.17 -0.45 Pan 23.40 -3.06 -0.08 Table 6.15: Inter-generic Euclidean distances based on PCA Ho. 23.12 -4.63 -0.07 scores from PC 1-5 per genus from Ln transformed generic Mean 23.15 -3.38 -0.12 means. Max. 24.96 -2.95 0.23 Hy. 0 Mean 2.45 Min. 20.26 -4.63 -0.45 Po. 3.77 0 Range 3.9 Range 4.71 1.68 0.67 Go. 4.71 1.20 0 SD 1.76 0.70 0.25 Pan 3.16 0.80 1.65 0 CV 7.62 20.81 208.31 Ho. 3.26 1.92 2.38 1.62 0 Hy. Po. Go. Pan Ho.

0.3 Table 6.16: Intra-generic Euclidean distances based Palatal and Interentoglenoid on Ln transformed data. 0.2 Po. breadths Mean Max Min Range 0.1 Hy. 1.12 2.36 0.40 1.96

Po. 1.06 2.00 0.48 1.52 0 Go. 1.07 2.16 0.33 1.83 PC 3: 1.63% Pan 0.76 1.46 0.36 1.10 -0.1 Ho.Pan Ho. 0.76 1.31 0.36 0.95 Mean Max Min Range -0.2 Hy. Hy. 1.12 2.36 0.40 1.96 Nasal & -0.3 Po. 1.06 2.00 0.48 1.52 Facial lengths Go. 1.07 2.16 0.33 1.83 -0.4

Pan 0.76 1.46 0.36 1.10 Go. Ho. 0.76 1.31 0.36 0.95 20 21 22 23 24 25 PC 1: 84.24% 316 Table 6.17: Variable loadings for Hominoidea Ln transformed generic means. PC loadings Component 1 Component 2 Component 3 nas-br 0.08 br-lam -0.43 inbrnabo -0.51 bizs 0.09 lam-opn -0.36 zs-zygi -0.26 inbrnabo 0.11 bas-br -0.27 g-o -0.23 bifmo 0.11 nas-br -0.26 nas-br -0.23 rhi-ns 0.12 g-o -0.22 bas-nas -0.21 bas-nas 0.13 inbrnabo -0.18 nas-rhi -0.19 bien 0.13 bien -0.16 bifmo -0.18 bienm 0.13 bienm -0.14 ol-sta -0.16 zs-zygi 0.13 biast -0.13 maxalvlen -0.11 g-o 0.14 bifmo -0.11 iv-pms -0.08 biecm 0.14 biaur -0.07 bifmt -0.08 biaur 0.15 bipor -0.07 nas-ns -0.06 bifmt 0.15 biecm -0.06 bas-pros -0.06 bas-br 0.15 bizi -0.05 pros-o -0.06 maxalvlen 0.15 pros-o -0.05 maxnawi -0.05 bipor 0.15 bifmt -0.05 zs-zi -0.05 biast 0.15 maxnawi -0.04 ol-pms -0.01 bas-pros 0.16 bizs -0.03 palhei -0.01 bicanex 0.16 rhi-ns -0.02 br-lam 0.05 bicanin 0.16 bas-nas -0.02 bipor 0.05 i1i2 0.17 bizygo 0.00 biast 0.09 ol-sta 0.17 zs-zygi 0.03 biaur 0.09 bizygo 0.17 zs-zi 0.04 lam-opn 0.09 biseptal 0.17 nas-pros 0.08 bicanex 0.11 br-lam 0.17 nas-ns 0.09 bizygo 0.12 bizi 0.17 iv-pms 0.13 biecm 0.12 pros-o 0.17 bicanin 0.14 nas-pros 0.13 maxnawi 0.18 nas-rhi 0.14 biseptal 0.14 palhei 0.18 biseptal 0.15 bas-br 0.15 zs-zi 0.19 palhei 0.15 i1i2 0.16 ol-pms 0.20 bicanex 0.18 bicanin 0.16 lam-opn 0.20 i1i2 0.19 bizi 0.17 iv-pms 0.20 maxalvlen 0.20 rhi-ns 0.19 nas-ns 0.22 ol-pms 0.20 bizs 0.21 nas-pros 0.25 bas-pros 0.20 bien 0.22 nas-rhi 0.31 ol-sta 0.24 bienm 0.24

317 (bizs) breadths with variable loadings of 0.24, 0.22 and 0.21, respectively. Juxtaposed against these cranial measurements are negatively loaded dimensions such as the inferior breadth of the nasal bones (inbrnabo), the maximum length of the zygomatic bone (zs-zgyi), cranial vault length (g-o), cranial height (bas-br) and inferior cranial length (bas-nas), with variable loadings of -0.51, -0.26, -0.23, -0.23 and -0.21, correspondingly. The scatterplots for PCs 1 & 2, PCs 2 & 3 and PCs 1 & 3 all revealed interesting configurations between hominoid genera (Figure 6.28). Table 6.15 provides the inter-generic Euclidean distance matrix produced from PCA object scores per genus from the first to fifth principal components based on Ln transformed generic means. The average distance between hominoid genera was 2.5 (Range 3.9). The largest distance produced was between Gorilla and Hylobates at 4.7 while the smallest distance was between Pan and Pongo with 0.80. For comparison, Table 6.16 provides intra-generic Euclidean distances based on Ln transformed data. The largest mean intra-generic Euclidean distance was produced by Hylobates with 1.12 (followed by Gorilla and Pongo with 1.07 and 1.06). This was due to the size differences between the smaller gibbons and the larger siamang. The smallest mean intra-generic Euclidean distance was produced by Pan and Homo each with 0.76. In summary, PCA of Ln transformed generic means revealed the incredible distinctiveness of Homo and Hylobates and the similarity in size between Pongo, Gorilla and Pan (Figure 6.28); although the scatterplot for PCs 1 & 3 revealed a close relationship between Pan and Homo. Viscerocranial dimensions were still amongst the higher variable loadings but so too were neurocranial dimensions (Table 6.17).

6.5.2 PCA results for MSV: By subjecting the generic mean MSV for the Hominoidea to PCA based on the variance-covariance matrix, the first (Eigenvalue - 0.92; 77.18%) and second (Eigenvalue - 0.18; 15.41%) PCs accounted for 92.59% of variation within the superfamily (5.03% less variation explained than PCs 1 & 2 for Ln transformed generic means but 77.58% more than PCs 2 & 3 for Ln transformed generic means; Figure 6.29). One hundred percent of the variation was explained by the four PCs. The variable loadings for the first PC were not all positive and so cannot be interpreted as size differences between genera; all PCs had mixed values. Table 6.18 lists the variable loadings for cranial MSV. Again, similar to results for the other primate groups studied,

318 Figure 6.29: Hominoidea scatterplot of PC 1 & 2 for generic mean MSV. Hy. - Hylobates; Po. - Pongo; Go. - Gorilla; Pan; and Ho. - Homo.

Facial length & 0 Table 6.18: Variable loadings for Hominoidea Occipital -0.1 Sagittal Chord -0.2 generic mean MSV. -0.3 Po. PC loadings -0.4 -0.5 Go. Ho. PC 1 PC 2 -0.6 bas-pros -0.32 nas-br -0.40 -0.7 Pan -0.8 ol-sta -0.19 bas-nas -0.31 PC 2: -0.9 maxalvlen -0.16 bifmo -0.29 -1 ol-pms -0.13 bizs -0.23 15.41% -1.1 -1.2 nas-pros -0.13 zs-zygi -0.18 -1.3 bicanex -0.11 bas-pros -0.16 -1.4 Hy. -1.5 Parietal Sagittal nas-ns -0.09 g-o -0.16 -1.6 Chord & Cranial nas-rhi -0.09 maxalvlen -0.14 -1.7 -1.8 Superior Facial length Vault length biseptal -0.07 bifmt -0.12 Frontal Sagittal -1.9 iv-pms -0.06 ol-sta -0.09 Chord & Inferior 2 3 4 5 bicanin -0.06 inbrnabo -0.05 Cranial Length i1i2 -0.05 rhi-ns -0.05 PC 1: 77.18% bizygo -0.04 biecm -0.04

% of zs-zi -0.03 biaur -0.04 PC Eigenvalue Var. Cum. %. palhei -0.03 bicanex -0.03 1 0.92 77.18 77.18 zs-zygi -0.02 bien -0.03 2 0.18 15.41 92.59 maxnawi 0.01 bienm -0.02 3 0.06 5.32 97.90 rhi-ns 0.01 bicanin -0.01 bas-nas 0.02 bipor -0.01 PCA bizs 0.03 i1i2 0.00 scores Axis 1 Axis 2 inbrnabo 0.03 biseptal 0.00 Hy. 2.39 -1.37 bizi 0.04 palhei 0.00 Po. 1.80 -0.28 biecm 0.04 maxnawi 0.03 Go. 1.99 -0.43 bifmt 0.06 biast 0.04 Pan 2.10 -0.63 bienm 0.06 zs-zi 0.05 Ho. 4.16 -0.47 bipor 0.08 iv-pms 0.06 Mean 2.49 -0.64 pros-o 0.08 ol-pms 0.07 Max. 4.16 -1.37 biaur 0.09 bizygo 0.10 Min. 1.80 -0.28 bifmo 0.13 bas-br 0.12 Range 2.36 1.09 bien 0.13 bizi 0.14 SD 0.96 0.43 biast 0.13 pros-o 0.15 CV 38.50 67.26 lam-opn 0.31 nas-ns 0.17 nas-br 0.32 nas-rhi 0.22 bas-br 0.36 br-lam 0.22 g-o 0.40 lam-opn 0.32 br-lam 0.42 nas-pros 0.41

319 the cranial MSV procure low to moderate variable loadings. Evidence for the removal and control of size from the datasets can be seen in the PC object scores per genus. Homo received the highest, 4.16, and Pongo received the least, 1.80; Gorilla, Pan and Hylobates had figures between these extremes. The first PC is dominated by positively loaded neurocranial dimensions. These include parietal sagittal chord (lam-opn), cranial vault length (g-o), cranial height (bas- br), frontal sagittal chord (nas-br) and occipital sagittal chord (lam-opn), with variable loadings of 0.42, 0.42, 0.36, 0.32 and 0.31, respectively. Working against these MSV is the superior facial length (bas-pros) with a negative loading of -0.32 (Table 6.18). The second PC contained an interesting assortment of length and breadth dimensions. Positively loaded MSV include the superior facial height (nas-pros), occipital and parietal sagittal chords (lam-opn and br-lam) and the sagittal length of the nasal bones and nasal height (nas-rhi and nas-ns) scoring 0.41, 0.32, 0.22 and 0.22, correspondingly. Contrasting with these cranial dimensions were negatively loaded MSV such as the frontal sagittal chord (nas-br), inferior cranial length (bas-nas), bifrontomalarorbitale breadth (bifmo) and bizygomaxillare superior breadth (bizs) procuring -0.40, -0.31, -0.29 and -0.23, respectively. The scatterplot for PC 1 & 2 for MSV (Figure 6.29) is very similar to the scatterplot for Ln transformed data. Table 6.19 provides the inter-generic Euclidean distance matrix produced from PCA object scores per genus from the first to fifth principal components based on generic mean MSV. The average distance between hominoid genera was 1.4 (Range 1.86). Now, the largest distance produced was between Homo and Pongo at 1.5 whereas the smallest distance was between Pan and Gorilla with 0.53. For comparison, Table 6.20 provides intra-generic Euclidean distances based on MSV. The largest mean intra- generic Euclidean distance was again produced by Hylobates at 0.79 (again, followed by Gorilla, 0.78). The smallest mean intra-generic Euclidean distance was produced by Pan at 0.58 (succeeded by Homo, 0.66).

Hy. 0 Mean 1.39 Table 6.19: Inter-generic Euclidean distances Po. 1.27 0 Range 1.86 based on PCA scores from PC 1-5 per genus Go. 1.11 0.72 0 from generic mean MSV. Pan 0.89 0.73 0.53 0

Ho. 1.99 2.39 2.20 2.09 0 Hy. Po. Go. Pan Ho.

320 Mean Max Min Range Hy. 0.79 1.25 0.36 0.89 Table 6.20: Intra-generic Po. 0.72 1.21 0.38 0.83 Euclidean distances based on MSV. Go. 0.78 1.31 0.34 0.97

Pan 0.58 1.01 0.29 0.72 Ho. 0.66 1.00 0.36 0.63

To summarize, results for hominoid MSV were similar to results for Ln transformed data although neurocranial dimensions figured more prominently. The scatterplot for PCs 1 & 2 based on MSV is very comparable to PCs 1 & 2 for Ln transformed data, but orientated differently.

6.5.3 CVA results for Ln transformed data: By subjecting the entire Hominoidea (n = 143) Ln transformed pooled sex multivariate dataset to MANOVA and CVA the first (Eigenvalue - 40.15; 45.99%) and second (Eigenvalue - 28.43; 32.57%) canonical variate axes were found to account for 78.56% of the variation within the superfamily (Figure 6.30). Table 6.22 lists the CVA object scores per genus and Table 6.23 lists the variable loadings for the cranial measurements. Again, on average, Homo had the greatest CVA object scores for CV axis 1 at 9.64 (Min. - Max., 9.42 - 9.77; followed by Gorilla with 8.98 (Min. - Max., 8.71 - 9.44)), whereas Hylobates scored the least with 7.59 (Min. - Max., 7.35 - 7.83). The variable loadings, like PCA, were low to moderate. Examination of the scatterplot for CV axes 1 & 2 reveals the distinctiveness of Hylobates and Homo but there is considerable overlap of Pan, Pongo and Gorilla (Figure 6.30). In order to test for significance of results, CVA object scores per genus (for both Ln transformed and MSV multivariate datasets) from the first ten CV axes, which together account for more than 95% of the variation, were subjected to a two-sample multivariate t-test using StatistiXL (version 1.5). For each pair wise generic comparison the null hypothesis that there is no significant difference was rejected (p-value < 0.0001). For CV axis 1 the highest positively loaded dimension was cranial height (bas- br), 0.44, followed by parietal sagittal chord (br-lam), maxillo-alveolar breadth (biecm), occipital sagittal chord (lam-opn), biauriculare breadth (biaur), interentoglenoid breadth (bien) and cranial vault length (g-o) with variable loadings of 0.27, 0.27, 0.24, 0.23, 0.23 and 0.22, correspondingly (Table 6.23). Juxtaposed against these measurements were negatively loaded dimensions such as superior facial length (bas-pros), external bicanine breadth (bicanex), and palatal length A (ol-sta) which achieved variable loadings of -0.37, -0.31 and -0.22, respectively.

321 Figure 6.30: Hominoidea scatterplot of canonical variate axes 1 & 2 of Ln transformed pooled sex data with 95% confidence ellipses. Hylobates - dot ; Pongo - cross +; Gorilla - square ; Pan - x; and Homo – open circle o.

Cranial height, 20 Parietal Sagittal 19 Chord & Biecm 18

CV 16 Axis 2: 32.57% 15

14 13

12 Superior Facial length & External Bicanine 11 Cranial height & Nasal Aperture Occipital Sagittal Chord height 8 9 10 CV Axis 1: 45.99%

MANOVA Wilks's Pillai lambda: 0.00 trace: 3.73 df1: 144.00 df1: 144.00 df2: 416.90 df2: 428.00 F: 51.38 F: 41.33 p(same): 0.00 p(same): 0.00 CVA Eigenvalue 1: 40.15 Percent: 45.99 Eigenvalue 2: 28.43 Percent: 32.57 Total %: 78.56

Table 6.21: Inter-generic Euclidean distances based on mean CVA scores from axes 1-5 per genus from Ln transformed data. Hy. 0 Mean 1.91 Po. 2.89 0 Range 3.07 Go. 3.64 0.95 0 Pan 2.52 0.57 1.21 0 Ho. 2.67 1.48 1.83 1.35 0 Hy. Po. Go. Pan Ho.

322 Table 6.22: Hylobates spp. (n=31; 16, 15). CVA scores Axis 1 Axis 2 Table 6.23: Variable loadings for Hominoidea Ln transformed pooled sex data. Mean 7.59 11.54 Max. 7.83 12.33 CVA Min. 7.35 10.76 Eigen- Eigen- vectors Axis 1 vectors Axis 2 Range 0.48 1.57 bas-pros -0.37 rhi-ns -0.20 SD 0.12 0.38 bicanex -0.31 nas-br -0.14 CV 1.53 3.27 ol-sta -0.22 zs-zygi -0.14 Pongo pygmaeus (n=26; 14, 12). maxalvlen -0.07 bienm -0.06 CVA scores Axis 1 Axis 2 i1i2 -0.06 ol-sta -0.06 Mean 8.62 14.16 bizygo -0.04 bizs -0.05 Max. 8.81 14.63 zs-zygi -0.03 biaur -0.02 Min. 8.35 13.66 iv-pms -0.03 bien -0.01 Range 0.46 0.98 zs-zi -0.02 lam-opn 0.01 SD 0.12 0.31 bipor -0.02 zs-zi 0.02 CV 1.40 2.19 Gorilla gorilla (n=34; 17, 17). palhei -0.02 bas-nas 0.03 CVA scores Axis 1 Axis 2 nas-ns -0.02 inbrnabo 0.04 Mean 8.98 14.78 bicanin 0.00 pros-o 0.06 Max. 9.44 15.32 ol-pms 0.00 maxalvlen 0.06 Min. 8.71 14.30 rhi-ns 0.02 bas-br 0.06 Range 0.74 1.01 biseptal 0.02 bizygo 0.07 SD 0.20 0.31 bizs 0.03 biast 0.07 CV 2.18 2.08 nas-pros 0.05 bifmo 0.07 Pan spp. (n=33; 19, 14) maxnawi 0.06 biecm 0.09 CVA scores Axis 1 Axis 2 bas-nas 0.06 bicanin 0.11 Mean 8.55 13.85 nas-rhi 0.07 br-lam 0.11 Max. 8.71 14.22 biast 0.07 g-o 0.11 Min. 8.39 13.53 bifmt 0.08 iv-pms 0.12 Range 0.32 0.69 inbrnabo 0.09 palhei 0.12 SD 0.08 0.18 bizi 0.10 bizi 0.13 CV 0.93 1.30 nas-br 0.15 bifmt 0.15 Homo sapiens (n=20; 11, 9). bifmo 0.15 bicanex 0.18 CVA scores Axis 1 Axis 2 pros-o 0.17 nas-ns 0.21 Mean 9.64 13.16 bienm 0.19 nas-pros 0.22 Max. 9.77 13.49 g-o 0.22 bipor 0.25 Min. 9.42 12.77 bien 0.23 ol-pms 0.26 Range 0.35 0.72 biaur 0.23 i1i2 0.27 SD 0.10 0.21 lam-opn 0.24 biseptal 0.28 CV 1.02 1.61 biecm 0.27 maxnawi 0.28 br-lam 0.27 bas-pros 0.30 bas-br 0.44 nas-rhi 0.44

323 The highest positively loaded measurement for CV 2 was the sagittal length of the nasal bones (nas-rhi), 0.44 (Table 6.21). Cranial height was followed by the superior facial length (bas-pros), maximum width of the nasal aperture (maxnawi), biseptal breadth (biseptal), alveolar length (i1i2), palatal length B (ol-pms), biporionic breadth (bipor), superior facial and nasal heights (nas-pros and nas-ns) which achieved variable loadings of 0.30, 0.28, 0.28, 0.27, 0.26, 0.25, 0.22 and 0.21, respectively. The largest negatively loaded measurement contrasting with the positively loaded dimensions was the sagittal height of the nasal aperture (rhi-ns), -0.20. The scatterplot for CV axes 1 & 2 (Figure 6.30) revealed the distinctiveness of Hylobates and Homo, which do not overlap with any other genus, but Pan, Pongo and Gorilla overlap considerably. Table 6.21 provides the inter-generic Euclidean distance matrix produced from mean CVA object scores per genus from the first to fifth canonical variate axes based on Ln transformed pooled sex data. The average distance between hominoid genera was 1.9 (Range 3.07). The largest distance produced was between Gorilla and Hylobates at 3.64 while the smallest distance was between Pan and Pongo with 0.57. In summary, CVA results for hominoid Ln transformed data strengthen results from PCA. Neuro- and viscerocranial dimensions equally contributed to segregating genera. Despite size differences, on average Homo produced the largest CVA object scores followed by Gorilla.

6.5.4 CVA results for MSV: After converting the raw data of the Hominoidea (n=143) into MSV and subjecting the entire multivariate dataset to MANOVA and CVA the first (Eigenvalue - 36.07; 51.70% of Var.) and second (Eigenvalue - 18.54; 26.58% of Var.), canonical variate axes were found to account for 78.28% of the variation (0.28% less variation explained than Ln transformed pooled sex datasets) within the superfamily (Figure 6.31). Tables 6.25 lists the CVA object scores per genus and Table 6.26 lists the low to moderate variable loadings for cranial MSV. Again, on average, Homo scored the largest CVA score for CV axis 1 at 4.06 (Min. - Max., 3.76 - 4.25; followed by Hylobates at 2.70 (Min. - Max., 2.45 - 2.97)) but Gorilla scored the least with 2.31 (Min. - Max., 2.11 - 2.62).

324 Figure 6.31: Hominoidea scatterplot of canonical variate axes 1 & 2 of pooled sex MSV with 95% confidence ellipses. Hylobates - dot ; Pongo - cross +; Gorilla - square ; Pan - x; and Homo – open Facial circle o. length & Lower facial breadth 0.6 0.5 0.4 0.3 0.2

0.1

CV 0 Axis 2: -0.1 32.57% -0.2 -0.3 -0.4 -0.5

-0.6 -0.7 Facial & Nasal lengths Cranial height, Occipital & -0.8 Parietal sagittal chords Length of Zygomatic 2 3 4 5 &Biorbital CV Axis 1: 45.99% breadth

MANOVA Wilks's Pillai lambda: 0.00 trace: 3.69 df1: 144.00 df1: 144.00 df2: 416.90 df2: 428.00 F: 41.26 F: 34.75 p(same): 0.00 p(same): 0.00 CVA Eigenvalue 1: 36.07 Percent: 51.71 Eigenvalue 2: 18.54 Percent: 26.58 Total %: 78.28

Table 6.24: Inter-generic Euclidean distances based on mean CVA scores from axes 1-5 per genus from MSV. Hy. 0 Mean 1.15 Po. 1.03 0 Range 1.52 Go. 0.90 0.63 0 Pan 0.70 0.65 0.43 0 Ho. 1.69 1.95 1.80 1.70 0 Hy. Po. Go. Pan Ho.

325 Table 6.25: Hylobates spp. (n=31; 16, 15). Table 6.26: Variable loadings for Hominoidea CVA scores Axis 1 Axis 2 pooled sex MSV. Mean 2.70 -0.56 Max. 2.97 -0.80 CVA Min. 2.45 -0.34 Eigen- Eigen- vectors Axis 1 vectors Axis 2 Range 0.52 0.45 bas-pros -0.34 zs-zygi -0.38 SD 0.15 0.11 nas-rhi -0.22 bifmo -0.30 CV 5.55 19.47 Pongo pygmaeus (n=26; 14, 12). bicanex -0.20 bas-nas -0.28 CVA scores Axis 1 Axis 2 biseptal -0.13 rhi-ns -0.25 Mean 2.32 0.35 bipor -0.12 nas-br -0.24 Max. 2.66 0.50 maxnawi -0.09 maxalvlen -0.20 Min. 2.15 0.16 ol-pms -0.06 bizs -0.17 Range 0.52 0.34 maxalvlen -0.01 ol-sta -0.14 SD 0.12 0.08 i1i2 -0.01 bicanin -0.11 CV 5.38 23.39 ol-sta -0.01 inbrnabo -0.09 Gorilla gorilla (n=34; 17, 17). bicanin -0.01 g-o -0.08 CVA scores Axis 1 Axis 2 iv-pms -0.01 bipor -0.07 Mean 2.31 0.12 bifmt 0.00 bifmt -0.03 Max. 2.62 0.4 bas-nas 0.01 bas-pros -0.02 Min. 2.11 -0.15 nas-ns 0.01 bicanex 0.00 Range 0.51 0.55 nas-pros 0.01 bien 0.00 SD 0.12 0.10 zs-zygi 0.05 bienm 0.01 CV 5.10 83.43 pros-o 0.05 biaur 0.01 Pan spp. (n=33; 19, 14). zs-zi 0.06 iv-pms 0.02 CVA scores Axis 1 Axis 2 bifmo 0.06 biast 0.03 Mean 2.42 0.03 biast 0.06 palhei 0.03 Max. 2.60 0.16 bizygo 0.07 zs-zi 0.04 Min. 2.23 -0.06 bizi 0.08 biecm 0.06 Range 0.37 0.22 palhei 0.08 nas-ns 0.08 SD 0.10 0.06 bizs 0.11 pros-o 0.09 CV 4.32 169.81 rhi-ns 0.17 nas-rhi 0.13 Homo sapiens (n=20; 11, 9). g-o 0.18 biseptal 0.15 CVA scores Axis 1 Axis 2 inbrnabo 0.20 br-lam 0.16 Mean 4.06 0.24 bien 0.21 bizygo 0.17 Max. 4.25 0.44 biaur 0.21 ol-pms 0.17 Min. 3.76 0.11 biecm 0.23 i1i2 0.19 Range 0.49 0.33 nas-br 0.26 lam-opn 0.21 SD 0.12 0.10 bienm 0.27 bas-br 0.22 CV 2.92 39.81 br-lam 0.28 maxnawi 0.22 lam-opn 0.32 bizi 0.23 bas-br 0.38 nas-pros 0.26

326 Again, very similar to results for Ln transformed data, examination of the scatterplot for CV axes 1 & 2 revealed the distinctiveness of Hylobates and Homo but there was considerable overlap of Pan, Pongo and Gorilla (Figure 6.31). The first three largest positively loaded MSV for CV 1 were neurocranial dimensions (Table 6.26). For example, cranial height (nas-br), and the occipital and parietal sagittal chords (lam- opn and br-lam) procured variable loadings of 0.38, 0.32 and 0.27, respectively. These MSV were followed by palatal breadth (bienm), frontal sagittal chord (nas-br), and maxillo-alveolar, biauriculare, interentoglenoid breadths (biecm, biaur, bien) with positive variable loadings of 0.23, 0.21 and 0.21, correspondingly. Juxtaposed against these positively loaded measurements were negatively loaded dimensions such as the superior facial length (bas-pros), sagittal length of the nasal bones (nas-rhi) and external bicanine breadth (bicanex), which achieved variable loadings of -0.34, -0.22 and -0.20, respectively. The largest positive and negative variable loadings for CV 2 were a mix of cranial breadth and length MSV (Table 6.26). On the positive side, the superior facial height (nas-pros) and bizygomaxillare inferior breadth (bizi) produced variable loadings of 0.26 and 0.23. These were closely followed by the maximum width of the nasal aperture (maxnawi), cranial height (bas-br) and the occipital sagittal chord (lam-opn), with variable loadings of 0.22, 0.22 and 0.21, respectively. Contrasting with these MSV were negatively loaded dimensions like the maximum length of the zygomatic bone (zs- zgyi), bifrontomalarorbitale breadth (bifmo), inferior cranial length (bas-nas), sagittal height of the nasal aperture (rhi-ns), frontal sagittal chord (nas-br) and maxillo-alveolar length (pros-dm3) which scored variable loadings of -0.38, -0.30, -0.28, -0.25, -0.24 and -0.20, correspondingly. The scatterplot for CV axes 1 & 2 based on MSV is very similar to that of Ln transformed data only orientated differently. However, Pan, Gorilla and Pongo still overlap whereas Homo and Hylobates were clearly separated. Table 6.24 is an inter-generic Euclidean distance matrix produced from mean CVA object scores per genus from the first to fifth canonical variate axes based on pooled sex MSV. The average distance between hominoid genera was 1.2 (Range 1.5). The largest distance produced was between Homo and Pongo at 1.95 whereas the smallest distance was between Pan and Gorilla with 0.43. For comparison, Table 6.27 provides Cherry et al’s (1978) M distance between hominoid genera. The average M distance between genera was 2.0 (Range 2.34). The largest morphological divergence

327 Hy. 0 Mean 2.0 Table 6.27: Inter-generic Cherry et Po. 1.56 0 Range 2.34 al’s M distance between hominoid Go. 1.44 1.23 0 genera. Pan 1.31 1.08 1.06 0

Ho. 2.42 3.4 3.19 3.34 0 Hy. Po. Go. Pan Ho. was achieved by Homo and Pongo, 3.4 (followed by Homo and Pan at 3.34); the smallest was between, Pan and Gorilla, 1.06 (succeeded by Pan and Pongo at 1.08). In summary, CVA results for hominoid MSV confirm those of PCA. Hylobates and Homo were well separated from Pongo, Gorilla and Pan, whereas the latter three overlap considerably. Again, neuro- and viscerocranial dimensions equally contributed to segregating genera.

6.6 Limb proportions: Published data on postcranial elements and limb proportions for the hominoids (Tables 6.1-6.5) was gathered from Schultz (1930, 1933, 1936a, 1936b, 1969, 1973), Napier & Napier (1967), Groves (1989 & 2001a), Aiello & Dean (1990), Jenkins (1990), Sarmiento (1994 & 1998) and Whitehead et al (2005). Limb proportions of the extant hominoids, particularly the intermembral index, clearly demonstrate the arboreal disposition of the apes in stark contrast to Homo (Tables 6.1 to 6.5). Hylobates and Pongo showed extreme arboreal adaptations with intermembral indices ranging from 130 to 150%. Gorilla and Pan were less, but still greater than 100%. On average, modern Homo had an intermembral index of 72-74%, which is highly derived compared to the other apes. Homo also had on average a rather low brachial index, 76%. Interestingly, all apes, excluding Pongo, had very similar crural indices, ranging from 83 to 86%; the orangutan scored 90%.

6.7 Genetics of the Hominoidea: Because of their close relationship to humans and historical interest (both scientific and social) the genetic differences (including macromolecules and sequenced nucleotides) between the apes and humans have received considerable attention (e.g. Gagneux et al, 1999; Fujiyama et al, 2002; DiVore & Gagneux, 2007). In fact, this large genetic research campaign served as the main impetus for the research herein (e.g. King & Wilson, 1975; Bruce & Ayala, 1978; Caccone & Powell, 1989; Rogers, 1993; Tamura & Nei, 1993; Chaline et al, 1996; Goodman et al, 1998; Dover, 1999; Arnason et al, 2000; Page & Goodman, 2001; Chen & Li, 2001; Chen et al, 2001; Fujiyama et al, 2002; Olson & Varki, 2003; Anzai et al, 2003; Curnoe & Thorne, 2003; Clark et al,

328 2003; Wildman et al, 2003; Hellmann et al, 2003; Uddin et al, 2004). In recent years, there have been two major developments. First, the diploid number and divergence estimates of the hylobatids have swayed some to employ this data in their classification schemes. Second, chimps and humans are genetically more close to one another than either is to the gorilla or orangutan. Thus, the African apes (Gorilla, Homo and Pan) share a more recent common ancestry to the exclusion of Pongo, the only remaining Asian great ape. Despite this, all great apes have a diploid number of 48, while the only biped, Homo, has 46. At present, the gibbons, like the cercopithecins, have varied diploid numbers. Firstly, Hylobates (Hylobates) 2n = 44; secondly, Hylobates (Hoolock) 2n = 38 (see Mootnick & Groves, 2005); thirdly, Hylobates (Nomascus) 2n = 52; and lastly, Hylobates (Symphalangus) 2n = 50 (Geissmann, 2002; Chatterjee, 2006). Furthermore, these and other genetic analyses suggest the four subgenera successively diverged from one another >1-10 Mya and are genetically as distinct as the extant hominids (Hayashi et al, 1995; Arnason et al, 1996; Goodman et al, 1998 & 2001; Roos & Geissmann, 2001). The diversity of the remaining four Great apes pales in comparison to the diversity of apes during the Miocene (Andrews, 1992). Now with the completion of the human, chimp and macaque genomes a much clearer and exciting picture is emerging from genetic studies (International Human Genome Sequencing Consortium, 2001; Chimpanzee Sequencing and Analysis Consortium, 2005; Rhesus Macaque Genome Sequencing and Analysis Consortium, 2007). Most striking perhaps are instances where humans and chimps share a particular mutation different to that of gorillas and the other apes (Wimmer et al, 2002) or human specific mutations which must have occurred after the chimpanzee and human divergence (Chou et al, 1998). Still, Tarzami et al, (1997) and Deinard & Kidd (1999) report genetic evidence linking all the Great apes. These particular features may be analyzed cladistically, alongside robust statistical support (Andrews, 1986; Andrews & Martin, 1987; Williams & Goodman, 1989; Ruvolo, 1997a; Page & Goodman, 2001; Wildman et al, 2004; Raaum et al, 2005). However, even though humans and chimps may be genetically very similar, recent genetic research suggests the speciation process for these taxa was a very complex event. For example, work by Patterson et al (2006) and Navarro & Barton (2003) suggest that during the initial speciation of these taxa, there was still some degree of genetic exchanges which may have continued for many generations before complete species

329 divergence. However, Szamalek et al (2007) recently analysed inverted chromosomal regions and their results suggest that these areas have not experienced major evolutionary divergence. In other words, inverted chromosomal regions have not necessarily undergone any tremendous sequence divergence in comparison to non- inverted regions. In addition, Bakewell et al (2007) have recently reported that chimpanzees have experienced greater genomic positive selection in comparison to humans. Table 6.28 provides the inter-generic genetic distances between hominoid genera from Page et al (1999) and Page & Goodman (2001).

Ho. 0 Mean 0.032 Table 6.28: Inter-generic Uncorrected ‘p’ Pan 0.0165 0 Range 0.028 genetic distances between hominoid Go. 0.019 0.0195 0 genera from Page et al, 1999. Po. 0.031 0.032 0.033 0 Hy. 0.042 0.0415 0.044 0.041 0 Ho. Pan Go. Po. Hy.

6.8 Discussion and Conclusion: From the forgoing analysis many sound inferences and conclusions may be drawn. 6.8.1 Question 1: Are the extant Catarrhini genera adequately defined and do they compose an ‘adaptively coherent’ grouping of one or more species which occupy an ‘adaptive zone’, as defined by Wood & Collard (1999a)? All extant hominoid genera (Hylobates, Pan, Pongo, Gorilla and Homo) are adequately defined, each adaptively coherent and occupying different adaptive zones. However, the only genus which deserves further attention is Hylobates because it is unclear whether or not the subgenera (Hylobates, Hoolock, Nomascus and Symphalangus) should be accorded with full generic status but all species share very similar adaptive strategies (Bartlett, 2007), although, generic boundaries could be defined by chromosome number.

330 to a separate family is justified, as is the creation of a family containing all Asian and African apes” (p. 43). Hylobates spp.: Overall, very small compared to the other apes. Some cranial proportions overlap with those of Homo (Figures 6.22 bizi/nas-pros; and 6.25 nas- pros/g-o). Sexual dimorphism is greatly reduced; males and females both have large canines. On average, palatal length (ol-sta) less than 60 mm whereas palatal breadth (bienm) less than 30 mm. Cranial height was on average less than 70 mm whereas superior facial height and length less than 50 mm and 90 mm, respectively. The nuchal crest is weakly defined and sagittal crests do not occur. Pongo spp.: Extreme sexual dimorphism. Sagittal crests and large face are common in adult males. Cranium is Airorhynchous. The nuchal crest is highly developed and robust, less in females. Pongo is unique in the relative proportion of superior facial breadth (bifmt) to bizygomaxillare inferior breadth (bizi). On average this index is less than 100% for the orangutan, whereas this proportion in other apes, including Homo, is greater than 100%. Additionally, the relative proportion of inferior cranial length to superior facial length is on average less than 70% for Pongo and greater than 70% for other apes. Gorilla spp.: Extreme sexual dimorphism. Sagittal crests, continuous and robust supraorbital torus and large viscerocranium are common in adult males. The nuchal crest is highly developed and robust, less in females. Gorillas on average have a larger inferior cranial length (bas-nas) than all other apes. Maxillo-alveolar breadth (biecm) on average less than 80% of palatal length (ol-sta); other apes are greater than 80%. Pan spp.: Sagittal crests are not common for chimpanzees, although some may develop strong temporal lines. Sexual dimorphism is not as great as Gorilla or Pongo. Pan is different from the other apes in the relative proportion of biasterionic breadth (biast) to biauriculare breadth (biaur). In the chimpanzee, this index is less than 80%, whereas the other apes produced greater than 80%. Modern Homo sapiens: Extreme encephalization, relatively large neurocranium and relatively small face. Sexual dimorphism is greatly reduced compared to Gorilla or Pongo. Cranium generally lacks strong muscle markings. The cranium base is absolutely and relatively wide in relation to inferior cranial length (bas-nas) and superior facial length (bas-pros). Maxillo-alveolar breadth (biecm) is relatively greater than 100% of palatal length (ol-sta) (Table 6.10, no. 26). Cranial height (bas-br) is on average greater than 100% of inferior cranial length (bas-nas), superior facial length

331 (bas-pros), and biasterionic, biauriculare and bizygomatic breadths (biast and biaur) (Table 6.10, no. 1, 2, 3, 4 and 6).

Question 3: How much morphological variation is encompassed within a Catarrhini genus? Ranges or variation for the hominoids were larger for than the Cercopithecidae, certainly the result of body size increase and extreme sexual dimorphism. Morphological variation, either singular variables or indices, for hominoid genera can generate standard deviations and coefficients of variation less than 5 to greater than 20 (Table 6.6). Highly variable measurements were associated with the nasal and facial region. In contrast, some neurocranial variables produced much less variation. Intra- generic Euclidean distances based on Ln transformed data were on average 0.95 (Max. - Min., Hylobates, 1.12 - Pan and Homo, 0.76) while those based on MSV were 0.71 (Max. - Min., Hylobates, 0.79 - Pan, 0.58) (Tables 6.16 and 6.20).

Question 4: How much has one genus morphologically and genetically diverged from another? With regard to morphological divergence, by measuring size (i.e. Ln transformed data) differences, not surprisingly but consistently, Gorilla and Hylobates produced the largest morphological distances while the smallest distances were between Pan and Pongo (Tables 6.15 and 6.21). However, when measuring shape differences (i.e. MSV) the greatest distances were produced between Homo and Pongo (Tables 6.19 and 6.24). Shape differences were confirmed by Cherry et al’s M distance. Again, the greatest distance produced was between Homo and Pongo whereas the least was between Pan and Gorilla (Table 6.27). The average M distance between genera was 2.0 (Range 2.34). In contrast, genetic comparative analyses placed Homo and Pan (0.0165) as the closest (Table 6.28, from Page et al, 1999 and Page & Goodman, 2001) while the greatest distance was between Hylobates and Gorilla (0.044).

Question 5: Do cranial morphometric similarities and/or differences reflect adaptive zones? Cranial morphometrics and/or dispositions which may indicate the adaptive zones of these genera include the following. Hylobates spp.: Gibbons are highly arboreal, rarely coming to the ground and travel by means of brachiation and the vertebral column is nearly vertical either sitting or while in motion. Associated with this is a cranium that lacks major muscle markings,

332 the neurocranium is globular. Similar to, Miopithecus, the crania of Hylobates spp. gives no suggestion of intense mastication requirements as well as no strongly developed nuchal crest to aid in balancing and/or holding the cranium in place while in motion or sitting, like that seen in terrestrial genera, e.g. Papio or Gorilla (see also generic description for question 2 above and generic summaries below). Pongo spp.: Orangutans are highly arboreal, limited terrestrial behaviors and the locomotion is described as quadrumanous climbing, clambering and/or swinging. Associated with this is a highly distinct, yet variable cranium between and within species (Uchida, 1996). Like Gorilla, adult male Pongo can develop robust nuchal crest and plane for neck muscle attachments. The airorhynchous disposition of Pongo crania could be an adaptation to their locomotor and positional behaviours (see also generic description for question 2 above and generic summaries below). Gorilla spp.: Gorillas are extremely large, sexually dimorphic apes which travel and forage along the forest floor but arboreal resources are still sought after and exploited. Associated with this is a highly distinct and unmistakable, yet variable cranium between and within species (Uchida, 1996; Albrecht et al, 2003; Leigh et al, 2003; Stumpf et al, 2003). Furthermore, gorilla populations are genetically diverse (Jensen-Seaman et al, 2003; Clifford et al, 2003). Lastly, the sheer size of gorilla crania provides an indication that these species are terrestrial (see also generic description for question 2 above and generic summaries below). Pan spp.: Chimpanzees are large apes which travel and forage on the forest floor and amongst the trees. Associated with this is a highly distinct cranium with reduced sexual dimorphism (compared to Pongo or Gorilla) but like Gorilla and Pongo, there is considerable variability within and between species (Shea & Coolidge, 1988; Shea et al, 1993; Uchida, 1996). Furthermore, chimpanzee populations are genetically diverse (Kaessmann et al, 1999; Gagneux et al, 2001; Stone et al, 2002) (see also generic description for question 2 above and generic summaries below). Homo sapiens: Modern humans are currently globally distributed and habitual and obligate bipeds. Associated with this is a highly derived cranium with a relatively large globular neurocranium and relatively small viscerocranium (see also generic description for question 2 above and generic summaries below).

Question 6: What analogies may be drawn from extant Catarrhini genera for the interpretation of fossil hominin genera?

333 Several analogies and considerations have been and continue to be drawn upon from the extant hominoids and their generic arrangement for the interpretation of species-concepts, extant and fossil hominoid systematics and socioecology (e.g. Ghiglieri, 1987; Tutin et al, 1991; Morin et al, 1994; Sarmiento et al, 2002; Albrecht et al, 2003; Guy et al, 2003; Miller et al, 2004; Lockwood et al, 2004; Stanford, 2006).

6.8.2 Generic Summaries: Superfamily: Hominoidea - Curnoe et al (2006) produced estimates of the maximum median duration of the process of speciation in this primate group to be .66 Ma. Family: Hylobatidae (Lesser Apes) Genus: Hylobates Illiger, 1811. Four Subgenera: Hylobates Illiger, 1811; Gibbons, a polytypic subgenus comprising seven species; H. (H.) lar [type species of genus and subgenus; Lar or White-handed Gibbon], H. (H.) agilis (Agile Gibbon), H. (H.) albibarbis (Bornean White-bearded Gibbon), H. (H.) muelleri Muller’s Bornean Gibbon), H. (H.) moloch (Silvery Gibbon), H. (H.) pileatus (Pileated Gibbon) and H. (H.) klossi (Kloss Gibbon or Bilou). 2n = 44. Symphalangus Gloger, 1841. Siamang, a monotypic subgenus; H. (S.) syndactylus. 2n = 50. Hoolock (not “Bunopithecus”; see Mootnick & Groves, 2005); Hoolock Gibbon, a monotypic subgenus; H. (H.) hoolock. 2n = 38. Nomascus Miller, 1903. Crested Indochinese Gibbons, a polytypic subgenus comprising five species; H. (N.) concolor (Concolor or Black-crested Gibbon), H. (N.) hainanus (Hainan Gibbon), H. (N.) leucogenys (type species of subgenus; Northern White- cheeked Gibbon), H. (N.) siki (Southern White-cheeked Gibbon) and H. (N.) gabriellae (Red-cheeked Gibbon). 2n = 52. The monophyly of the Nomascus subgenus is supported by the recent mtDNA analyses by Monda et al (2007). The hylobatids have been the subjects of a long history of research (Groves, 1972; Frisch, 1973; Andrews & Simons, 1977; Andrews & Groves, 1976; Creel & Preuschoft, 1976; Myers & Shafer, 1979; Marshall & Sugardjito, 1986; Arnason et al, 1996a). The hylobatids are postcranially highly derived apes (Schultz, 1973) but do still share some primitive characteristics with the cercopithecines, such as ischial callosities and small body size (Rowe, 1996), yet cranially and dentally are nonetheless hominoids (Schultz, 1930 & 1933; Frisch, 1973). In recent years molecular studies have revealed a

334 much clearer picture of gibbon phylogeny and biogeography, which has led some to acknowledge either full generic or subgeneric rank (Hayashi et al, 1995; Hall et al, 1998; Roos & Geissmann, 2001; Geissmann, 2002). Although no ancient fossils have been allocated to this genus per se, many extinct Miocene genera from Europe and China have gibbon-like characteristics, e.g. Pliopithecus (Simons & Fleagle, 1973; Fleagle, 1984). Based on genetic data Goodman et al (1998 & 2001) suggest hylobatid lineages are ~7 to 8 million years old yet diverged from the other hominoids ~ 18 Ma. Interestingly, some female gibbons display sexual swelling while others do not (Cheyne & Chivers, 2006). Unlike the eye-orbits of the other apes, in Hylobates the eyes are surrounded by a “peculiar thickened rims” (Groves, 1989; p. 155). Hylobates also exhibit slim zygomatic arches and a narrow corpus of the mandible (Whitehead et al, 2005).

Family: Hominidae (Great Apes) Subfamily: Ponginae (Asian Apes) Pongo Lacepede, 1799; Orangutans. 2 species (see Xu & Arnason, 1996), P. pygmaeus (Borneo) [type species of genus] and P. abelii (Sumatra) with some subfossil evidence (e.g. Marcus, 1969). Yet, there are extinct fossil genera which show pongo- like characteristics, most notably Sivapithecus and Lufengpithecus (Andrews & Cronin, 1982; Shea, 1985; Schwartz, 1990; Kelley, 2002; Chaimanee et al, 2003; Fox et al, 2004). 2n = 48. Recent genetic and morphological data presented by Steiper (2006) confirms a clear distinction between P. abelii and P. pygmaeus, having estimated to be separated for ~ 2.7-5 million years. In contrast to the other Great apes which are klinorhyncous and possess a continuous supraorbital torus, Pongo is airorhynchous and possesses a discontinuous brow ridge (Whitehead et al, 2005). Furthermore, a particularly important feature differentiating the African and Asian apes can be found in the nasoalveolar region in sagittal section, which reveals the different orientation of the premaxilla and hard palate (Brown et al, 2005).

Subfamily: Homininae (African Apes) Based on genetic data Goodman et al (1998) suggest extant lineages emerged ~ 7 Ma. Tutin et al (1997), report the co-existence of Homo-Pan-Gorilla 60 Ka in Gabon. Gorilla I. Geoffroy, 1853; Gorillas. 2 species, G. gorilla (Western) [type species of genus] and G. beringei (Eastern), in total comprising four subspecies (G. g.

335 gorilla & G. g. diehli. and G. b. beringei & G. b. graueri) (see Tutin et al, 1991; Sarmiento, 1994; Garner & Ryder, 1996; Sarmiento & Oates, 2000; Jensen-Seaman et al, 2003; Stumpf et al, 2003; Leigh et al, 2003; Groves, 2003; Gorilla gorilla uellensis - Hofreiter et al, 2003; Miller et al, 2004; Robbins, 2007). 2n = 48. The gorillas, like the chimpanzees, hold a special place among the primates for many reasons, both historical and scientific. Recent genetic analysis confirms the divergence between the Eastern and Western species and subspecies (Clifford et al, 2003) but surprisingly both species have certain haplotypes in common (Jensen-Seaman et al, 2003), a feature not seen in the common and bonobo chimpanzees. In addition, gorilla subspecies can be easily distinguished from cranial and postcranial data (Leigh et al, 2003; Schultz, 1934). Moreover, Remis (2003) provides evidence to show gorillas are not merely herbivores but actively pursue fruits when available. Pan Oken, 1816; Chimpanzees. 2 species (see Fenart & Deblock, 1973;; McHenry & Corruccini, 1981; Coolidge & Shea, 1982; McHenry, 1984b; Vervaecke, & van Elsacker, 1992; Videan & McGrew, 2002; Williams et al, 2003), Pan troglodytes [type species of genus] and three subspecies, Pan t. troglodytes, Pan t. verus & Pan t. schweinfurthii; however, another has been described - Gonder et al, 1997, Pan t. vellerosus) and Pan panicus (Cramer, 1977; Zhilman et al, 1978; Zhilman & Cramer, 1978; Zhilman & Lowenstein, 1983); with very little fossil evidence (Mcbrearty & Jablonski, 2005; DeSilva et al, 2006). 2n = 48. The history of science and the chimpanzees is an interesting yet complicated one; namely due to a plethora of nomenclature synonyms and misidentifications with other primates (Hill, 1969a & b; Yerkes & Yerkes, 1970; Groves, 2001a). In the past some researchers placed the gorillas in Pan (e.g. Simpson, 1963) due to similarities such as knuckle-walking and gradistic concepts of evolutionary systematics. Moreover, Dainton & Macho (1999) have demonstrated that chimpanzee and gorilla knuckle-walking are different; they also suggest that knuckle-walking may have evolved independently in both lineages. In addition, numerous studies, both anatomical and molecular, have demonstrated that Pan and Homo form a clade to the exclusion of all other extant apes (Groves, 1986 & 1989; Williams & Goodman, 1989; Shoshani et al, 1996; Ruvolo, 1997a; Goodman et al 1998; Gibbs et al, 2002; Page & Goodman, 2002; Wimmer et al, 2002; Lockwood et al, 2004; Uddin et al, 2004). Lastly, the cultural and behavioural complexities of these species (and the other Great apes) have been studied in detail (McGrew et al, 1981; Collins & McGrew, 1988; Boesch & Boesch, 1989; Tutin et al, 1991; Boesch &

336 Tomasello, 1998; Whiten et al, 1999; Videan & McGrew, 2002; Boesch, 2003). Finally, chimpanzees are incapable of articulate speech (Duchin, 1990). Homo Linnaeus, 1758; Humans. At present, only one global-polytypic species surviving from many putative fossil ancestors (Weidenreich, 1943; Wood, 1991; Tobias, 1991; Grine et al, 1996; Wood & Collard, 1999 & 2001; Curnoe & Tobias, 2006; Rightmire et al, 2006). 2n = 46. Based on genetic data Goodman et al (1998) suggest Homo lineages diverged from our last common ancestor with the chimpanzees ~ 4-6 Ma. According to Groves (2001a) modern Homo sapiens is, “Based on humans from Europe, Africa, Asia, and America. No type specimen exists, and no lectotype has ever been proposed” p. 308; and detecting any modern human subspecies (if they exist) is “not resolvable” p.309. Paleontology, genetics and developmental biology have revealed a complex, rapid and fairly recent evolutionary history for modern Homo sapiens (Krings et al, 1997; Lieberman et al, 2002; Carrol, 2003; Bogin & Rios, 2003; Lovejoy et al, 2003; White et al, 2003; Dorus et al, 2004; McDougall et al, 2005). Furthermore, the major defining feature of this evolutionary history, and the clade to which humans belong, has been greatly influenced by the selection for efficient bipedalism, intra-specific cooperative behaviour and cognitive abilities; the former of which is substantiated by evidence from the fossil record and the latter two are determined by developmental, ecological, physical (i.e. anatomical and morphological) and mental attributes (Falk, 1985; Tobias, 1987; Aiello & Dunbar, 1993; Aiello & Wheeler, 1995; Gannon et al, 1998; Dainton & Macho, 1999b; Bingham, 1999 & 2000; Aiello & Wells, 2000; McBrearty & Brooks, 2000; McHenry & Coffing, 2000; Dean et al, 2001; Stiner, 2002; Asfaw et al, 2002; Semendeferi et al, 2002; Leonard et al, 2003; Holloway et al, 2003; Bramble & Lieberman, 2004; Jablonski & Chaplin, 2004; Bobe & Behrensmeyer, 2004; Coqueugniot et al, 2004; Curnoe & Tobias, 2006; Walker et al, 2006; Thorpe et al, 2007).

337 Chapter 7: Summary and Discussion of results for the extant Catarrhini genera 7.1 Introduction The purpose of this chapter is to briefly summarize and discuss the results of the previous four chapters dealing with the extant Catarrhini genera as a whole and outline a comparative framework with which to apply to fossil hominin genera and assess their biological comparability, usefulness and validity. The extant Parvorder or Infraorder Catarrhini (E. Geoffroy, 1812) (Figure 7.1; Old World monkeys and apes; see Delson & Andrews, 1975; Andrews, 1985; Andrews et al, 1996) contains 26 genera and 152 species (Groves, 2001a), many of which are sympatric yet competition is greatly reduced by niche separation (e.g. Clutton-Block, 1974; Dunbar & Dunbar, 1974; Curten, 1976; Peters & O’Brien, 1981; Coe, 1984; Mitani, 1991; Tutin et al, 1991; Wahungu, 1998). All Catarrhini species have a dental formula of 2.1.2.3; postorbital closure; generally the frontal and sphenoid bones articulate to form the sphenofrontal suture (instead of the zygomatic and parietal bones as in the Platyrrhini); and a bony tube leading to the external auditory meatus (all key features which differentiate catarrhines from the Platyrrhini (New World monkeys); Fleagle, 1998). The average number of species per genus is 5.9, ranging from one to 25 with a standard deviation of 6.5. However, if genera that contain more than ten species are excluded (i.e. Cercopithecus (n=25), Macaca (n=20), Trachypithecus (n=17), Presbytis (n=11) and Hylobates (n=14)) the average number of species per genus is reduced to 3.1, ranging from one to 9 with a standard deviation of 2.4. In comparison, the average number of species per genus for the Platyrrhini (New World Monkeys, 16 genera and 107 species; Groves, 2001a), sister-group to the Catarrhini (which together form the higher primates, Simiiformes, or the traditional Anthropoidea), yet having experienced separate evolutionary histories and trajectories (Ciochon & Chiarelli, 1980; Schneider et al, 1993; Horovitz et al, 1998), is 6.7, ranging from one to 17 with a standard deviation of 5.5. Again, however, if genera with ten or more species are excluded (i.e. Callithrix (n=17, including three subgenera), Saguinus (n=17), Callicebus (n=15) and Alouatta (n=10)), the average number per genus reduces to 4.0, ranging from one to eight with a standard deviation of 2.5. The origin of the catarrhines lies in the Eocene-Oligocene (Simons & Rasmussen, 1996; Kay et al, 1997). Please bear in mind, despite more than thirty million years of evolution, there are known wild and/or captive intergeneric and interspecific hybridizations within the primate groupings studied herein (Van Gelder, 1977; Myers & Shafer, 1978; Bernstein & Gordon, 1980; Struhsaker et al, 1988; Vervaecke & van Elsacker, 1992; Jolly et al, 1997; Moore et al, 1999; Detwiler, 2002; Ackermann et al, 2006; Arnold & Meyer, 2006); and has been suggested for hominin species as well (Duarte et al, 1999; Wolpoff et al, 2001; Hunt, 2003; Holliday, 2003; Henneberg & De Miquel, 2004; Trinkaus, 2007). The species of extant Catarrhini genera range in size from around one kilogram (Miopithecus) to > 100 kg (Pongo, Homo and Gorilla). Currently, the extant catarrhines (ex. humans) are unevenly distributed across Africa and Eurasia. In terms of genus and species numbers, cercopithecines dominate sub-Saharan Africa, whereas colobines outnumber cheek- pouch monkeys in South, East and Southeast Asia. In addition, these two regions each have two ape genera; Gorilla and Pan in Africa; and Hylobates and Pongo in East and Southeast Asia; and modern Homo is globally distributed.

New World Al. Mi. Cp. Ch. E. Cc. Mn. Mc. L. Pp. Th. C P N Hy. Po. Go. Pan Homo Monkeys

Cercopithecini Papionini

Hylobatidae Hominidae

Cercopithecinae Colobinae

Cercopithecidae

Cercopithecoidea Hominoidea

Platyrrhini Catarrhini

Figure 7.1 A taxonomic and phylogenetic Simiiformes diagram representing the likely relationships Haplorrhini within the extant Catarrhini based on molecular and morphological data. Primates

Generic Abbreviations: Al. - Allenopithecus; Mi. - Miopithecus; Cp. - Cercopithecus; Ch. - Chlorocebus; E. - Erythrocebus; Cc. - Cercocebus; Mn. - Mandrillus; Mc. - Macaca; L. - Lophocebus; Pp. - Papio; Th. - Theropithecus; C = Colobina (Co. - Colobus; Pi Piliocolobus. - & Pro. - Procolobus); P = Presbytina (Se. - Semnopithecus, Tr. - Trachypithecus & Pre. - Presbytis); N = Nasalina (Py. - Pygathrix, R. - Rhinopithecus, N. - Nasalis & Si. - Simias); Hy. - Hylobates; Po. - Pongo; & Go. - Gorilla.

339 7.2 Summary per catarrhine grouping: Important features which distinguish a genus from other closely allied genera included the following. 7.2.1 Tribe Cercopithecini: The five cercopithecin genera are a group of primates with a body mass of less than 25 kg and are restricted to tropical forests and savannas mosaics of sub-Saharan Africa. Cranial regions and measurements that were significant in sorting genera include: Nasal, Palatal, Cranial and Facial Lengths: Sagittal length of the nasal bones (nas-rhi); Palatal length A (ol-sta); Parietal sagittal chord (br-lam); Occipital sagittal chord (lam-opn) and Superior facial height nas-pros). Nasal, Facial, Palatal and Cranial breadths: Inferior breadth of the nasal bones (inbrnabo); Bizygomaxillare superior breadth (bizs); Bizygomaxillare inferior breadth (bizi); Palatal breadth (bienm); Maxillo-alveolar breadth (biecm) and Biasterionic breadth (biast). These cranial dimensions and their relative contribution and/or proportion to other cranial dimensions proved useful in distinguishing papionin genera but more importantly, many produced statistical significance. Bizs/bizi x 100 Bizs/zs-zgyi x 100 Nas-rhi/nas-pros x 100 Zs-zgyi/ol-sta x 100

340 7.2.2 Tribe Papionini: The majority of papionins, ranging in size from ~5 kg up to 35-40 Kg, also inhabit sub-Saharan Africa. The genus Macaca enjoys a large geographic range in South, East and Southeast Asia. Cranial regions and measurements that were significant in sorting genera include: Nasal, Palatal, Facial and Cranial Lengths: Sagittal length of the nasal bones (nas-rhi); Palatal length A (ol-sta); Superior facial height (nas-pros) and Occipital sagittal chord (lam-opn). Nasal, Facial and Cranial Breadths: Bizygomaxillare superior breadth (bizs); Bizygomaxillare inferior breadth (bizi); and Biasterionic breadths (biast). These cranial dimensions and their relative contribution and/or proportion to other cranial dimensions proved useful in distinguishing papionin genera but more importantly, many produced statistical significance. Biast/lam-opn x 100 Bifmt/bizi x 100 Bifmt/nas-pros x 100 Bizi/nas-pros x 100 Zs-zgyi/ol-sta x 100

341 7.2.3 Subfamily Colobinae: The colobines are complex-stomach monkeys which vary in body size from <5 kg up to 35-40kg, and are unevenly distributed - three genera occur in Africa whereas seven are distributed across East, South and Southeast Asia. Cranial regions and measurements that were significant in sorting genera include: Nasal, Facial and Palatal Lengths: Sagittal length of the nasal bones (nas-rhi); Inferior cranial length (bas-nas); Superior facial height (nas-pros); Superior facial length (bas-pros); Palatal length A (ol-sta) and Maximum length of the zygomatic bone (zs-zgyi). Facial, Nasal, Palatal and Cranial breadths: Superior facial breadth (bifmt); Bizygomaxillare superior breadth (bizs); Bizygomaxillare inferior breadth (bizi); Palatal breadth (bienm) and Maxillo-alveolar breadth (biecm) These cranial dimensions and their relative contribution and/or proportion to other cranial dimensions proved useful in distinguishing colobine genera but more importantly, many produced statistical significance. Bas-br/bas-pros x 100 Biecm/ol-sta x 100 Bifmt/nas-pros x 100 Ol-sta/bas-pros x 100

342 7.2.4 Superfamily Hominoidea: The remaining five genera are hominoids and they range in size from >5 Kg to greater than 100 kg, and are morphologically and behaviorally diverse. In addition, the “morphological gap” (see, Mayr, 1969; and Simpson, 1963) between hominoid genera is greater than that seen in either the Cercopithecinae or Colobinae. Cranial regions and measurements that were significant in sorting genera include: Facial, Cranial and Palatal Lengths: Inferior cranial length (bas-nas); Parietal sagittal chord (br-lam); Occipital sagittal chord (lam-opn); Superior facial length (bas-pros); Maximum length of the zygomatic (zs-zgyi); Zygomatico-maxillary suture length (zs-zi); Superior facial height (nas-pros) and Palatal length A (ol-sta). Facial and Palatal Breadths: Maxillo-alveolar breadth (biecm); Palatal breadth (bienm); Biporionic breadth (bipor); Biasterionic breadth (biast) and Interentoglenoid breadth (bien). These cranial dimensions and their relative contribution and/or proportion to other cranial dimensions proved useful in distinguishing hominoid genera but more importantly, many produced statistical significance. Bas-nas/bas-pros x 100 Biecm/zs-zgyi x 100 Bipor/g-o x 100 Nas-pros/g-o x 100

343 7.3 Multivariate Statistics: The results from multivariate statistics were both statistically robust and reproduced results between primate groupings studied herein and from the published literature. Both PCA and CVA generally resulted in neurocranial dimensions being juxtaposed against facial and nasal lengths. Overall, variable loadings were low to moderate which is related to dataset homogeneity and data transformation prior the multivariate analysis (Tabachnick & Fidell, 2007). To identify the underlying structure (i.e. the cranial variables which account for the most variation between genera) of the entire Catarrhini (n=26) multivariate dataset based on Ln transformed generic means and generic mean MSV, PCA was employed, as in the previous chapters. Figures 7.2 and 7.4 provide the results of applying PCA to all the catarrhines, both Ln transformed generic means and generic mean MSV. Table 7.1 provides the variable loadings for Ln transformed data, whereas Table 7.2 provides the variable loadings for MSV.

7.3.1 Catarrhini Ln transformed data: PCA derived from the Ln transformed generic means based on the variance- covariance matrix resulted in the first (Eigenvalue - 3.36; 87.34%), second (Eigenvalue - 0.29; 7.49%) and third (Eigenvalue - 0.06; 1.58%) PCs accounting for 96.41% of variation within the parvorder (Figure 7.2). The low to moderate variable loadings for the first PC are all positive but PCs 2 and 3 have mixed values (Table 7.1). The first PC is dominated by nasal and palatal dimensions. These include sagittal length of the nasal bones (nas-rhi), superior facial height (nas-pros), maximum width of the nasal aperture (maxnawi), length between incisivion and the palatomaxillary suture (iv-pms), palatal length B (ol-pms), nasal height (nas-ns), zygomatico-maxillary suture length (zs-zi) and palatal length A (ol-sta) with variable loadings of 0.26, 0.23, 0.22, 0.21, 0.20, 0.20, 0.20 and 0.20, respectively. Measurements which are positively loaded for the second PC include three nasal length dimensions, sagittal length of the nasal bones (nas-rhi), nasal height (nas-ns) and superior facial height (nas-pros) with variable loadings of 0.54, 0.30 and 0.20, correspondingly. Juxtaposed against these dimensions are negatively loaded measurements such as bizygomaxillare superior breadth (bizs), parietal sagittal chord (br-lam), occipital sagittal chord (lam-opn) and cranial height (bas-br) with variable loadings of -0.26, -0.24, -0.22 and -0.21, respectively.

344

Figure 7.2: Catarrhini (n=26) scatterplot of PC 1 & 2 and PC 2 & 3 for Ln transformed generic means.

Nasal Pp. length & PC 2: Increase in terrestrialism Mn. % of Cum. height 7.49% PC Eigenvalue Var. %. 0.6 Th. Mc. 1 3.36 87.34 87.34 Al. 2 0.29 7.49 94.83 L. Cc. 0.3 Si. 3 0.06 1.58 96.41 Cp. N. E. Semiterrestrial Ch. 0 /Arboreal Pi. Mi. Co. Pro. Se. -0.3 PCA Tr. Pan Po. Go. Arboreal Py. scores PC 1 PC 2 PC 3 Hy. Pre. -0.6 Mean 20.89 -4.61 1.56 R. Max. 25.08 -3.63 1.96 Increase in body size -0.9 Min. 17.51 -6.11 0.77 Range 7.58 2.48 1.19 SD 1.83 0.54 0.25 -1.2 Bizygomaxillare Superior, Parietal CV 8.77 11.63 15.83 -1.5 & Occipital Ho. Nasal & Sagittal Chords Facial -1.8 lengths

-4 -3 -2 -1 0 1 2 3 4

PC 1: 87.34%

PC 3: Ho. 0.72 1.58%

0.6 Bizygomaxillare Superior & External Bicanine 0.48 breadths

0.36

0.24 Mi. Length of Mc. Cp. Nasal Bones, E. Al. 0.12 Occipital & Si. Th. Parietal Cc. Pre. Pro. Ch. Pp. N. L. 0 Sagittal Go. Chords Mn.

-0.12 Bizygomaxillare Pi. Superior, Parietal & Tr. Nasal Pan Se. -0.24 length & Occipital Sagittal Hy.Py. Chords height -0.36 Po. PC 2: 7.49% Co. R. -1.8 -1.5 -1.2 -0.9 -0.6 -0.3 0 0.3 0.6

345 Table 7.1: Variable loadings for Catarrhini Ln transformed generic means.

PC loadings PC 1 PC 2 PC 3 bizs 0.10 bizs -0.26 nas-rhi -0.35 nas-br 0.11 br-lam -0.24 lam-opn -0.34 bifmo 0.12 lam-opn -0.22 br-lam -0.33 bien 0.12 bas-br -0.21 inbrnabo -0.31 bas-nas 0.12 biast -0.19 bas-br -0.18 bifmt 0.13 nas-br -0.18 zs-zi -0.17 g-o 0.13 bifmo -0.18 g-o -0.14 zs-zygi 0.13 bifmt -0.15 nas-br -0.13 biaur 0.14 g-o -0.15 nas-ns -0.09 rhi-ns 0.14 biaur -0.14 bienm -0.08 biecm 0.14 bipor -0.13 nas-pros -0.05 bizygo 0.15 bien -0.12 biast -0.05 br-lam 0.15 maxnawi -0.12 pros-o -0.03 bas-br 0.15 bizi -0.12 palhei -0.02 bipor 0.15 bienm -0.10 zs-zygi -0.01 palhei 0.15 bas-nas -0.09 maxnawi 0.00 bienm 0.15 bizygo -0.09 bipor 0.02 maxalvlen 0.16 biecm -0.07 bien 0.03 biast 0.16 bicanin -0.04 biecm 0.03 bizi 0.16 inbrnabo -0.04 biaur 0.04 bicanex 0.16 bicanex -0.03 bifmo 0.06 bas-pros 0.17 pros-o -0.02 bas-nas 0.07 biseptal 0.17 zs-zygi 0.01 bizi 0.08 pros-o 0.17 biseptal 0.02 bifmt 0.09 i1i2 0.17 rhi-ns 0.03 rhi-ns 0.11 bicanin 0.17 palhei 0.03 i1i2 0.13 inbrnabo 0.17 bas-pros 0.06 bicanin 0.13 lam-opn 0.18 i1i2 0.09 bizygo 0.14 ol-sta 0.20 zs-zi 0.09 iv-pms 0.14 zs-zi 0.20 iv-pms 0.10 biseptal 0.15 nas-ns 0.20 maxalvlen 0.12 ol-pms 0.17 ol-pms 0.20 ol-pms 0.15 maxalvlen 0.20 iv-pms 0.21 ol-sta 0.17 bas-pros 0.22 maxnawi 0.22 nas-pros 0.20 ol-sta 0.22 nas-pros 0.23 nas-ns 0.30 bicanex 0.22 nas-rhi 0.26 nas-rhi 0.54 bizs 0.29

346 Positively loaded measurements for the third PC include bizygomaxillare superior (bizs) and external bicanine breadths (bicanex), palatal length A (ol-sta) superior facial length (bas-pros) and maxillo-alveolar length (pros-dm3) with variable loadings of 0.29, 0.22, 0.22, 0.22 and 0.20, correspondingly. Contrasting with these dimensions are negatively loaded measurements such as sagittal length of the nasal bones (nas-rhi), occipital sagittal chord (lam-opn), parietal sagittal chord (br-lam) and inferior breadth of the nasal bones (inbrnabo) with variable loadings of -0.35, -0.34, - 0.33 and -0.31, respectively. The scatterplots for PCs 1 & 2 and PCs 2 & 3 each reveal the dramatic separation of modern humans from the other catarrhines. Figure 7.3 is Non-metric MDS scatterplot of the Euclidean distance matrix based on the PCA object scores from PCs 1 to 5 from Catarrhini Ln transformed generic means. The average Euclidean distance based on Ln transformed generic means between catarrhine genera was 2.29 (n=325; Range 7.32; SD 1.5); not surprisingly, the largest distance produced was between the largest and smallest extant catarrhines, Gorilla and Miopithecus (7.58); whereas the smallest distance was between a cercopithecine and colobine, Chlorocebus and Simias (0.27).

347 Figure 7.3: Non-metric multi-dimensional scaling of the Euclidean distance matrix based on the PCA scores per genus for PC 1-5 from Ln transformed generic means (Stress: 0.04).

Mi.

0.24

0.2

0.16

0.12

Coordinate 2 0.08 Pro.Pre. Ho.

0.04 Tr. Go. Po. Si. Al. Ch. Pan 0 R. Cp. Py. Pi.

-0.04 Hy. Pp. Co. Mn. Se. Th. N. -0.08 E. L. Cc.Mc.

-0.16 -0.08 0 0.08 0.16 0.24 0.32 0.4 0.48

Coordinate 1

Mean Max. Min. (n=325) (Go.-Mi.) (Ch.-Si.) Range SD 2.29 7.58 0.27 7.32 1.5 R%:319 MI:28

348 7.3.2 Catarrhini generic mean MSV: PCA derived from the generic mean MSV based on the variance-covariance matrix resulted in the first (Eigenvalue – 0.50; 56.79%) and second (Eigenvalue - 0.14; 15.96%) PCs accounting for 72.75% of variation within the parvorder (22.08% less variation explained than PCs 1 & 2 for Ln transformed generic means but 63.68% more variation explained than PCs 2 & 3 for Ln transformed generic means; Figure 7.4). Both PCs 1 and 2 have mixed values and, again, the variable loadings are low to moderate (Table 7.2). Positively loaded measurements for the first PC include cranial vault length (g- o), frontal sagittal chord (nas-br), bifrontomalarorbitale and bifrontomalartemporale breadths (bifmo and bifmt), cranial height (bas-br) and bizygomaxillare superior breadth (bizs) which procure variable loadings of 0.28, 0.27, 0.25, 0.22, 0.22 and 0.20, respectively. Juxtaposed against these dimensions are negatively loaded measurements such as superior facial height (nas-pros), nasal height (nas-ns), sagittal length of the nasal bones (nas-rhi) and palatal length A (ol-sta) with variable loadings of -0.33, -0.33, -0.32 and -0.22, correspondingly. The two largest positively loaded measurements for the second PC include the parietal and occipital sagittal chords (br-lam and lam-opn) with variable loadings of 0.26 and 0.25. Contrasting with these dimensions are negatively loaded measurements such as superior facial length (bas-pros), bizygomatic breadth (bizygo), inferior cranial length (bas-nas), maxillo-alveolar length (pros-dm3) and bifrontomalarorbitale and bifrontomalartemporale breadths (bifmo and bifmt) which achieve variable loadings of - 0.42, -0.39, -0.27, -0.26, -0.22 and -0.22, respectively. The scatterplot for PCs 1 & 2 again exhibits the impressive shape differences of modern humans in comparison to the remaining catarrhines but also Miopithecus. Figure 7.5 is a Non-metric MDS scatterplot of the Euclidean distances matrix based on the PCA object scores from PCs 1 to 5 from generic mean MSV. The average Euclidean distance based on generic mean MSV between catarrhine genera was 1.10 (n=325; Range 3.22; SD 0.63). The largest distance produced was between Homo and Papio (3.31); the smallest distance was between Cercocebus and Lophocebus (0.09).

349 Figure 7.4: Catarrhini (n=26) scatterplot of PC 1 & 2 for generic mean MSV.

-4 PC 2: 15.96% Parietal & Occipital Mi. Sagittal Chords

Ho.

Long viscerocranium Globular neurocranium, -5 Very little facial projection

Go. Pan Po.Cc.L. Cp. Mc. E. Al. Ch. Pp. Mn. Hy. Th. Si. Pro. Pre. Cranial Superior N. Pi. Co. length & Superior Facial Se. Tr.Py. Facial & height length & Nasal R. Bizygomatic height PC 1: 56.79% breadth 1 2 3 4

% of PC Eigenvalue Var. Cum. % 1 0.50 56.79 56.79 2 0.14 15.96 72.75 3 0.10 11.22 83.96

PCA scores PC 1 PC 2 Mean 2.27 -5.40 Max. 3.58 -4.22 Min. 0.54 -5.86 Range 3.04 1.64 SD 0.70 0.37 CV 31.12 6.91

350 Table 7.2: Variable loadings for Catarrhini generic mean MSV.

PC loadings PC 1 PC 2 nas-pros -0.33 bas-pros -0.42 nas-ns -0.33 bizygo -0.39 nas-rhi -0.32 bas-nas -0.27 ol-sta -0.22 maxalvlen -0.26 bas-pros -0.16 bifmt -0.22 ol-pms -0.16 bifmo -0.22 maxalvlen -0.12 ol-sta -0.18 iv-pms -0.09 biaur -0.17 zs-zi -0.08 bien -0.17 pros-o -0.08 bizs -0.16 i1i2 -0.03 zs-zygi -0.16 biseptal -0.03 bizi -0.16 bicanin -0.01 nas-ns -0.14 bicanex -0.01 pros-o -0.13 rhi-ns -0.01 rhi-ns -0.12 palhei -0.01 nas-br -0.11 maxnawi -0.01 bicanex -0.11 inbrnabo 0.00 ol-pms -0.11 zs-zygi 0.01 biecm -0.09 bienm 0.03 bipor -0.08 biecm 0.05 nas-pros -0.08 bizi 0.09 iv-pms -0.05 lam-opn 0.11 biseptal -0.05 bipor 0.13 i1i2 -0.02 bien 0.14 g-o -0.02 biast 0.14 bicanin -0.02 bizygo 0.15 nas-rhi -0.01 bas-nas 0.17 palhei 0.00 br-lam 0.18 bienm 0.00 biaur 0.19 zs-zi 0.01 bizs 0.20 maxnawi 0.02 bas-br 0.22 inbrnabo 0.02 bifmt 0.22 biast 0.07 bifmo 0.25 bas-br 0.15 nas-br 0.27 lam-opn 0.25 g-o 0.28 br-lam 0.26

351 Figure 7.5: Non-metric multi-dimensional scaling of the Euclidean distance matrix based on the PCA scores per genus for PC 1-5 from generic mean MSV (Stress: 0.08).

0.36 Ho.

0.3

0.24

0.18 R.Pre.

Mi. 0.12 Tr.Py. Pro.

Coordinate 2 0.06 Hy. Se. Pi. 0 Co. Si. Ch. N. Po. -0.06 Cp. Pan Al. E. Go. -0.12 L. Cc. Mc. Th. Mn. Pp. -0.18

-0.18 -0.12 -0.06 0 0.06 0.12 0.18 0.24 0.3 0.36

Coordinate 1

Mean Max. Min. (n=325) (Ho.-Pp.) (Cc.-L.) Range SD 1.10 3.31 0.09 3.22 0.63 R%:292 MI:36

7.4 Morphological and Genetic Distances: Morphological distances are useful because the can reduce the differences of many variables to a single figure. In addition, morphological distances can highlight genera or species that are dramatically different as well as reveal similarities between closely related genera. Tables 7.3 and 7.4 list the mean, maximum and minimum Euclidean distances for the primate groupings studied herein. Figure 7.3 is a Non- metric Multi-dimensional scaling (NM-MDS) scatterplot based on Euclidean distance matrix from Ln transformed data. Figure 7.5 is a NM-MDS scatterplot based on Euclidean distance matrix from MSV. For comparison, Figure 7.4 is a NM-MDS

352 Figure 7.6: Non-metric multi-dimensional scaling of the genetic distance matrix from Page et al, 1999 & Page & Goodman, 2001; Catarrhini generic means only (Stress: 0.09).

Cpo 0.18 10-9 Ma Tob Nla Cgu 0.15 Colobinae

0.12

0.09

Cercopithecoidea 14 Ma 0.06 25-28 Ma Coordinate 2 0.03 18 Ma Hla 0 Hominoidea

7 Ma Ppy -0.03 Cercopithecinae GgoPan Hsa 5-6 Ma Cae Cce -0.06 Epa 7 Ma Cga Mn. Mc.TgeLat Pcy -0.09

-0.36 -0.3 -0.24 -0.18 -0.12 -0.06 0 0.06 0.12

Coordinate 1

scatterplot based on genetic distance matrix from Page et al (1999) and Page & Goodman (2001). Table 7.3: Euclidean distances produced from the generic mean PCA object scores from PCs 1-5 based on Ln transformed data (see Tables 3.15, 4.16, 5.21 & 6.16). Euclidean Distances Mean Max. Min. based on Ln Distance Distance Distance transformed data: Cercopithecini 1.57 3.41 0.52 Papionini 1.51 2.81 0.38 Colobinae 1.02 1.71 0.40 Hominoidea 2.45 4.71 0.80 Catarrhini 2.29 7.58 0.27

353 Table 7.4: Euclidean distances produced from the generic mean PCA object scores from PCs 1-5 based on MSV (see Tables 3.19, 4.20, 5.24 & 6.19). Euclidean Distances Mean Max. Min. based on MSV: Distance Distance Distance Cercopithecini 0.69 1.18 0.37 Papionini 0.87 1.46 0.24 Colobinae 0.67 1.10 0.29 Hominoidea 1.39 2.39 0.53 Catarrhini 1.10 3.31 0.09

Three observations emerged by tallying the mean, maximum and minimum Euclidean distances between genera from Ln transformed data and MSV. First, distances between monkey genera were similar. For size, Cercopithecini and papionini were very similar, whereas for shape, Cercopithecini and Colobinae were very similar. Second, distances based on shape (MSV) were much smaller than those based on size (Ln transformed). Third, distances between the hominoids were greater than cercopithecoids.

7.5 Summary of findings: 7.5.1 Size of the cranium: It has been found here that the cranium size of adults is a very important distinguishing factor between catarrhine genera, intimately related to body size (i.e. mass). In all groups studied, facial and nasal dimensions were responsible for most of the variation and were the main distinguishing features between genera. In addition, the generic means for many cranial dimensions were found to be significantly correlated to generic mean body weights in linear regression (Tables 3.14, 4.15, 5.19 & 6.14; Figures 3.28, 4.27, 5.27, 6.27). Body size, for catarrhines, also provides indications of possible preferred habitat types, i.e. small body size = likely arboreal, limited use of open, terrestrial niches, selective feeders; medium or larger body size = likely some semiterrestrial behaviors or increase in terrestrialism and increase in dietary choices (Figures 7.2 & 7.4). Cranial regions and measurements which were useful in delineating genera include: Facial and Nasal regions: Superior facial height and length (nas-pros and bas-pros);

354 Facial breadths (bifmt, bizs, bizi and bizygo) and Length of the Zygomatic (zs-zgyi). Length and Breadth of the Palate: Palatal length A (ol-sta) and Maxillo-alveolar and Palatal breadths (biecm and bienm). Neurocranial lengths and breadths and associated proportions: Cranial, Frontal, Parietal and Occipital Sagittal Chords (g-o, nas-br, br-lam and lam-opn) and Biasterionic and Biporionic breadths (biast and bipor).

7.5.2 Shape of the cranium: When size is removed from analyses, cranial shape reveals stark contrasts, important phylogenetic inferences as well as reducing the variation between different species, and sexually dimorphic males and females. The following were found to be significant distinguishers of catarrhine genera: Cranial height and facial lengths (bas-br, nas-pros, bas-pros) / cranial breadths (biast and bipor); Facial breadths (bifmt or bifmo) / facial lengths or breadths (nas-pros and bizi) and Palatal length (ol-sta) / Palatal breadths (biecm, bienm) and facial lengths (nas-pros, nas-pros, zs-zgyi and zs-zi).

7.5.3 Number of species per genus: Although the number of species per genus ultimately rests with an organisms’ evolutionary history, adaptive radiation, being preserved and later discovered in the fossil record, as well as a taxonomist’s judgment, there are nonetheless trends and thresholds in extant catarrhine genera irrespective of hierarchical ranking. Table 7.5 provides the average number of species per genus with the standard deviation (second column), and the average number of species per genus but excluding genera with ten or more species with the standard deviation (third column; however, please note, for the Presbytina, Presbytis spp. were included with the third column figures but has eleven species, instead Trachypithecus was excluded because it has 17 species; and for the Hylobatidae, Hylobates (Hylobates) were excluded from the third column figures but has only seven species). For out-group comparison, the Platyrrhini were also listed.

355 The most striking result of this summary of species per genus is the uniformity across most genera. The largest average number of species per genus was for the members of Presbytina with 11.7. However, this figure is skewed because Trachypithecus has 17 species. But even if Trachypithecus is removed and the average for Semnopithecus (7 species) and Presbytis (11 species) is calculated, the mean still remains the highest of all groups at 9.0. But most importantly, extant Great apes are not particularly speciose. In fact, the hominids produced the smallest average number of species per genus of all primate groups. 2. Average No. of 3. Average No. of species 1. Hierarchical Rank species per genus per genus minus genera with >10 species Cercopithecinae (n=11) ¯x 6.5 8.2 (n=9) ¯x 3.0 2.1 Table 7.5: Average no. of species per genus for Groves, Cercopithecini (n=5) ¯x 7.0 10.3 (n=4) ¯x 2.5 2.4 2001a; those in bold represent Papionini (n=6) ¯x 6.2 7.0 (n=5) ¯x 3.4 2.1 the maximum and minimum for both the average ( ¯x ) and Colobinae (n=10) ¯x 5.9 5.2 (n=8) ¯x 3.9 3.3 standard deviation () per Colobina (n=3) ¯x 5.0 4.0 (n=3) ¯x 5.0 4.0 column. Presbytina (n=3) ¯x 11.7 5.0 (n=2) ¯x 9.0 2.8 Nasalina (n=4) ¯x 2.3 1.5 (n=4) ¯x 2.3 1.5 Cercopithecoidea (n=21) ¯x 6.2 6.8 (n=17) ¯x 3.4 2.5 Hominoidea (n=5) ¯x 4.2 5.5 (n=4) ¯x 1.8 0.5 Hylobatidae (n=4) ¯x 3.5 3.0 (n=3) ¯x 2.3 2.3 Hominidae (n=4) ¯x 1.8 0.5 (n=4) ¯x 1.8 0.5 Catarrhini (n=26) ¯x 5.9 6.5 (n=21) ¯x 3.1 2.4 Platyrrhini (n=16) ¯x 6.7 5.4 (n=12) ¯x 4.0 2.4 Simiiformes (n=42) ¯x 6.2 6.1 (n=33) ¯x 3.4 2.4 Range of ¯x : 9.9 Range of ¯x : 7.2 Range of : 9.8 Range of : 8.5

7.6 Conclusion and Application to Hominins: With this comparative framework in place the fossil hominin genera can be assessed. Admittedly though, little can be said about the cranial morphology and disposition of Orrorin and Ardipithecus. The extant catarrhines are both phenotypically and behaviorally diverse yet do share several morphological features and are the most appropriate analogs to compare and judge the fossil record and biological classification of hominin and/or human evolution.

356 Chapter 8: Results for genera of the Hominina

Pan Sahelanthropus Ardipithecus Australopithecus Paranthropus Homo

? Panina Hominina

Hominini Homininae Hominidae Hominoidea Figure 8.1: A taxonomic and phylogenetic Catarrhini diagram representing the possible Simiiformes relationships within the Hominini based on Haplorrhini biomolecular and morphological data. Primates 8.1 Introduction: The purpose of this chapter is to examine and report the results for genera of the Subtribe Hominina (Gray, 1825) by implementing similar methods as the previous five chapters for the extant catarrhines. The fossil record bearing on hominin evolution is quite diverse, both temporally and geographically, having been recovered from Late Miocene to Pleistocene deposits in Central, Eastern and Southern Africa. The paleohabitat reconstructions for these early hominin deposits indicate moist, wooded environments near water (Pickford & Senut, 2001; Woldegabriel et al, 2001; Vignaud et al, 2002; White et al, 2007). However, later hominin genera and species have been recovered from more open and drier paleohabitats (Stanley, 1992; Suwa et al, 1997; Reed, 1997). By using the “partial” taxonomy of the Hominoidea and fossil hominins from Tuttle (2006), the average number of species for seven genera is 3.0, ranging from one to seven with a standard deviation of 2.5 (Tuttle, 2006; p. 251); and seven species are listed within Homo. However, if monotypic genera are excluded (i.e. Sahelanthropus, Orrorin and Kenyanthropus), the average number of species per genus increases to 4.5, ranging from two to seven with a standard deviation of 2.4. Interestingly, the hominin taxonomy reviewed and proposed by Wood & Richmond (2000), prior to the announcements of Sahelanthropus, Orrorin, Kenyanthropus and A. kadabba, the average number of species for four genera was 4.5 with a standard deviation of 2.9; and eight species are listed within Homo. Lastly, using the first (FAD) and last appearance dates (LAD) provided by Foley (1999a, p. 331, table 23-1), on average, four genera and eleven species occur in the African Plio-Pleistocene fossil record for only a brief period of time, ~ 0.7 million years (Myr). Furthermore, this figure does not change drastically if only the FAD and LAD of genera is considered; Ardipithecus, 0.7 Myr; Australopithecus, 0.87 Myr; Paranthropus, 0.7 Myr; and early Homo, 0.50 Myr. For thoroughness, original family and generic diagnoses of hominines and hominins are located in the appendix. Body weight estimates (based on regression models from fossil cranial and postcranial remains) for some of these hominins vary from >20 up to < 80-100 kg (Jungers, 1988; McHenry, 1988; Aiello & Wood, 1994; Foley, 1999a & 1999b; Collard, 2002; Spocter & Manger, 2007). Please note, taxonomy within the Hominidae and/or Hominini continues to be contentious and other generic and specific names are available for fossil material (Groves, 1999). Genera and species of hominins currently used in the paleoanthropological literature include the following: Sahelanthropus tchadensis (Brunet et al, 2002 & 2005; Vignaud et al, 2002;) is the binomial name given to fossil material from Late Miocene deposits, ~6-7 Ma in Chad, in Central Africa. The holotype (TM 266-01-060-1) is a nearly complete but distorted cranium, which has been virtually reconstructed (Zollikofer et al, 2005). Features which Brunet et al (2002) cite as important characteristics include, the short face crowned by a robust brow ridge; large compound temporal-nuchal crest; small canines (which they argue is evidence for a non-honing C-P3 complex) and small cheek teeth; wide interorbital pillar; a sagittal crest is present; and the face is less prognathic than Pan or Australopithecus. Guy et al’s (2006) morphometric analysis of TM 266-01-060-1, based the reconstruction (Zollikofer et al, 2005), reveal similarities to both Pan and Australopithecus (discussed more fully below).

358 Orrorin tugenensis (Senut et al, 2001; Sawada et al, 2002; Pickford et al, 2002; Galik et al, 2005) is the binomial name given to fossil material from Late Miocene deposits, ~6 Ma in Kenya, Eastern Africa. The holotype and paratypes include both dental and postcranial fossil remains. Differential characteristics of particular note are the smaller jugal teeth compared to Australopithecus; greater enamel thickness than Ardipithecus; and proximal femora morphology more human-like than australopithecines or African apes (Senut et al, 2001). However, other morphological features of the humerus (distal shaft exhibits a straight lateral crest for the insertion of the m. brachioradialis) and a curved proximal phalanx, suggest climbing adaptations as seen in chimps and A. afarensis specimens (Senut et al, 2001). Nakatsukasa et al (2007) estimate the body weight a young adult is ~35-50 kg with a stature of 1.1-1.2 m tall based on linear regression analyses from femoral dimensions and comparisons to extant skeletal material. Ardipithecus ramidus (White et al, 1994 & 1995; type species of genus) and A. kadabba (Haile-Selassie, 2001; Haile-Selassie et al, 2004) are the binomial names for two fossil species described from specimens recovered from Late Miocene-Early Pliocene deposits in the Afar depression of Ethiopia, Eastern Africa (but the former was originally announced as a species of Australopithecus; and the latter was originally described as a subspecies of the former but was later elevated to full species status). White et al (1994) argue for the inclusion of the Aramis fossils in the hominin clade due to two derived characters which all hominins possess; 1) “anterior placement of the foramen magnum”; and 2) “a more incisiform canine with reduced sexual dimorphism” (p. 312). However, White et al (1994) also note the primitive nature of the fossil material and its very close to resemblance to chimpanzees, particularly in tooth size and enamel thickness. The genus Australopithecus Dart, 1925 currently encompasses many fossil species restricted to sub-Saharan Africa which have a geological timespan of just over four million years ago to a little more than two million years ago with a geographic range from Central Africa to Eastern Africa and to Southern Africa (Robinson, 1954, 1956 & 1972; Foley, 1999a; Brunet et al, 1995; Kimbel, 1995). Characteristics which typify this genus include, a form of bipedalism (not exactly like modern humans) yet a postcranial skeleton suggesting continued use of arboreal niches, dramatic reduction in the sexual dimorphism of canines but some increase in cranial capacity, with thick tooth enamel, large post-canine

359 dentition, anterior pillars and generally lacking sagittal crests (excluding AL-444-2) (Wolpoff & Lovejoy, 1975; Walker, 1976; Grine, 1981 & 1986; Rak, 1983; McHenry, 1984a; Wood & Chamberlain, 1986; Grine & Kay, 1988; Grine & Martin, 1988; McKee, 1989; Kimbel et al, 2004). The first appearance of this genus is the material allocated to A. anamensis in northeastern Ethiopia at ~4.2 Mya and similarly aged deposits in Kenya (Leakey et al, 1995 & 1998; Ward et al, 1999 White et al, 2006). Kimbel et al (2004) have presented compelling evidence of mandibular morphology demonstrating A. anamensis was ancestral to A. afarensis (see also White et al, 2006). The A. afarensis hypodigm has caused controversy since its inception (e.g. Day et al, 1980). Many argued it was only a variant or subspecies of A. africanus in Southern Africa (e.g. Tobias, 1980). Additionally, the locomotor repertoire of this fossil species has been interpreted as nearly human-like bipedalism (Lovejoy et al, 1973 & 2002), to a form of bipedalism that still allowed the exploitation of arboreal niches (Susman et al, 1984 & 1985; Hunt, 1998; Ward, 2002). Finally, very recently Rak et al (2007) have presented convincing evidence of mandibular morphology linking A. afarensis to the ‘robust australopiths’. Other species of this genus include A. bahrelghazali (Brunet et al, 1995) for a mandibular fragment from Chad with an estimated age of 3.0-3.5 Ma, which is very similar to A. afarensis; A. garhi (Asfaw et al, 1999; Strait & Grine, 2001) for a partial skull and skeleton associated with stone tools and butchered animal bones from Ethiopia dated to ~2.5 Ma; and finally, A. africanus [type species of genus] from the paleo-cave sites in Southern Africa with an approximate temporal range of ~3.0 to >2.0 Ma. (Ahern, 1998; Lockwood & Tobias, 1999 & 2002). Objections to this genus stem from cladistics analyses of primarily craniodental features by Strait et al (1997; see also Strait & Grine, 2005). Their results have argued that the inclusion of A. afarensis within the genus Australopithecus results in paraphyletic trees. Thus they recommend its removal from Australopithecus and instead be placed in Praeanthropus; the original genus name for the Garusi maxillary fragment (see Puech et al, 1986). Wood & Collard’s (1999a) review and analysis of the genus category agreed with the results of Strait et al (1997). Kenyanthropus platyops (Leakey et al, 2001; Lieberman, 2001; Cameron, 2003; K. rudolfensis? Cela-Conde & Ayala, 2003), or the flat-faced man of Kenya, is the binomial

360 name for a fossil genus and species from mid-Pliocene deposits, ~3.5 Ma, in Kenya, Eastern Africa. The holotype is a distorted cranium (White, 2003), KNM-WT 40000; a specimen with both primitive and derived characters. Interestingly, KNM-WT 40000, possesses several features seen in another fossil cranium, KNM-ER 1470 (which was traditionally assumed to be a member of the genus Homo; e.g. Leakey, 1973b), while differing markedly from the species of Paranthropus. Some features which KNM-WT 40000 and KNM-ER 1470 share to the exclusion of Paranthropus include thin palate; a flat nasoalveolar clivus with a stepped nasal cavity entrance; a low and curved zygomaticoalveolar crest; a flat midface transverse contour and tall malar region; and moderate post-orbital constriction (Leakey et al, 2001, p. 434, table 2). This led to Cela- Conde & Ayala (2003) to suggest KNM-WT 40000 be placed in the genus Homo, which would make the first appearance date for the genus at 3.5 Ma. Paranthropus spp. is the generic name given to fossils recovered initially by Broom (1938, 1954, 1956 & 1972) in the 1930’s, which were commonly referred to as the ‘robust australopithecines’ (e.g. Day, 1969; McHenry, 1988; Turner & Wood, 1993; McCollum, 1999). The first appearance of this genus is in Eastern Africa ~ 2.5-2.7 Ma with P. aethiopicus (Walker et al, 1986; Kimbel, 1995; however, the binomial nomenclature of this fossil taxon is in need of refinement; Groves, 1999) and the last appearance date is ~ 1 Ma with P. boisei (original genus name, “Zinjanthropus”; see Tobias, 1967 and Constantino & Wood, 2007) and P. robustus (Klein, 1988; Foley, 1994, 1999a & 1999b; Wood & Strait, 2004). The type species of the genus is P. robustus. The defining features of this genus include a broad yet flat and dished face with wide zygomatic arches, robust masticatory cranial (e.g. sagittal crest and strong musculature markings) and mandibular (e.g. large mandibular corpus; Hylander, 1988) features and very large postcanine dentition, including molarization of the premolars, with very thick tooth enamel (Grine, 1981 & 1986; Rak, 1983; McHenry, 1984a & 1988; Clarke, 1985; Grine & Martin, 1988). The monophyletic status of this genus has been both supported (Turner & Wood, 1993; Strait et al, 1997) and questioned (Wood, 1988; McCollum, 1999; McHenry et al, 2007). Despite questions of monophyly, it is important to note that Susman (1988a & 1988b) has demonstrated that P. robustus had the manual dexterity to make and use tools. These

361 observations place the bone-tool findings of Backwell & d’Errico (2001) in a very interesting light, although more than one species is known from Swartkrans. The genus of modern humans, Homo, has both historical and biological importance. Historically the genus Homo is significant because by giving humans a binomial name, Linnaeus put humans firmly within the biological world. Biologically the genus Homo is essential because field research and laboratory analysis has unmistakably demonstrated that modern humans were once one of many, though different, adapted species (Wood, 1991 & 1992; Dunsworth & Walker, 2002). Some of the oldest fossils that have been attributed to the genus Homo have been reported by Hill et al (1992), Schrenk et al (1993), Kimbel et al, (1996) and Prat et al (2005). Fossil species generally allocated to this genus and used in the paleoanthropological literature include: H. rudolfensis (Lieberman et al, 1996; Wood, 1999; Kennedy, 1999); H. habilis (Leakey, 1966; Tobias, 1966; Stringer, 1986; Rightmire, 1993; Grine et al, 1996); H. australis (Curnoe, in press); H. ergaster (Groves & Mazak, 1975); H. erectus (Weindenreich, 1939, 1943 & 1946; Santa Luca, 1980; Anton, 2002; Manzi et al, 2003; Asfaw et al, 2002; Kidder & Durband, 2004); H. georgicus (Gabunia et al, 2000; Vekua et al, 2002; Lordkipanidze et al, 2006; Rightmire et al, 2006); H. antecessor (Arsuaga et al, 1997); H. hiedelbergensis (Rightmire, 1996 & 2004) H. cepranensis (Manzi et al, 2001); H. neanderthalensis (Krings et al, 1997; Schillaci & Froehlich, 2001; Williams et al, 2003; Harvati, 2003; Thackeray et al, 2003; Noonan et al, 2006; Green et al, 2006; Macchiarelli et al, 2006; Weaver et al, 2007; Trinkaus, 2007); H. sapiens idaltu (White et al, 2003);

362 H. floresiensis (Brown et al, 2004; Morwood et al, 2005; Weber et al, 2005; Falk et al, 2005a, 2005b & 2006; Jacob et al, 2006; Martin et al, 2006a & 2006b; Hershkovitz et al, 2007) and H. sapiens Linnaeus, 1758 [type species of genus] (Liebermann et al, 2002; Carroll, 2003)

8.2 Descriptive Statistics and Univariate Analyses for cranial variables: Tables 8.1 to 8.15 provide the descriptive statistics for the Hominin genera cranial variables. Craniometric data of the fossil hominin genera were complied from published paleoanthropological literature (primarily Wood, 1991; Tobias, 1991; Kimbel et al, 2004; Lordkipanidze et al, 2006) original fossil crania (Sts 5 and SK 48) and fossil casts (e.g. OH 24, KNM-ER 1470, KNM-ER 1813, KNM-WT 17000). Australopithecus spp. (Au.): The sample for Australopithecus includes two species, A. afarensis and A. africanus; Paranthropus spp. (Para.): The sample for Paranthropus includes three species, P. aethiopicus, P. boisei and P. robustus; Homo1: The sample for Homo1 includes all extinct fossil species traditionally attributed to Homo (H. habilis, H. rudolfensis, H. erectus, H. ergaster, H. hiedelbergensis and H. neanderthalensis); Au.+hab: The sample for Au.+hab includes Australopithecus species and H. habilis and H. rudolfensis as Wood & Collard (1999a) suggested; Homo2: The sample for Homo2 contains all extinct fossils species of Homo (H. erectus, H. ergaster, H. hiedelbergensis, and H. neanderthalensis) excluding H. habilis and H. rudolfensis; H. sap (modern Homo sapiens): and finally, the last sample is only modern H. sapiens, the same sample used in chapter six. These arrangements were devised to simulate and test various hypotheses including that of Wood & Collard (1999a). However, consideration must be given to for fossil preservation, possible deformation, measurement estimates and measurements from fossil casts. Thus, results for the hominins should be considered as approximates. Despite Schultz’s admonishment,

363 Tables 8.1 – 8.15: Descriptive Statistics of cranial variables for the fossil hominin genera (Au. – Australopithecus spp.; Para. – Paranthropus spp.; Homo1 – H. habilis, H. rudolfensis, H. erectus, H. ergaster, H. heidelbergensis and H. neanderthalensis; Au.+hab – Australopithecus spp., H. habilis and H. rudolfensis; Homo2 – H. erectus, H. ergaster, H. heidelbergensis and H. neanderthalensis; H.sap. – modern H. sapiens).

8.1 G-o Au. Para. Homo1 Au.+hab Homo2 H.sap. N 3 5 44 6 41 20 Mean 147.6 156.2 190.8 150.5 193.5 174.8 Median 146.8 163.0 193.5 146.9 195.5 175.7 Max. 167.0 173.0 219.0 167.0 219.0 190.7 Min. 129.0 130.0 146.0 129.0 153.0 160.5 Range 38.0 43.0 73.0 38.0 66.0 30.2 SD 19.0 18.2 16.9 14.5 13.6 8.5 CV 12.9 11.7 8.8 9.6 7.0 4.9

8.2 Biast Au. Para. Homo1 Au.+hab Homo2 H.sap. N 10 5 51 10 45 20 Mean 84.9 91.2 117.0 88.5 120.0 105.3 Median 80.8 93.0 119.5 85.0 120.0 105.1 Max. 103.0 96.5 142.0 112.0 142.0 120.3 Min. 74.0 81.0 80.0 74.0 92.0 96.8 Range 29.0 15.5 62.0 38.0 50.0 23.5 SD 10.6 6.5 13.5 13.1 10.6 5.7 CV 12.5 7.1 11.5 14.8 8.9 5.4

8.3 Bipor Au. Para. Homo1 Au.+hab Homo2 H.sap. N 3 5 19 8 15 20 Mean 98.3 118.4 120.3 108.1 123.1 110.1 Median 100.0 115.4 122.6 102.5 124.0 109.9 Max. 100.0 134.4 134.0 132.0 134.0 121.7 Min. 95.0 103.0 100.0 95.0 106.5 98.1 Range 5.0 31.4 34.0 37.0 27.5 23.7 SD 2.9 12.1 9.7 13.7 6.9 7.1 CV 3.0 10.3 8.1 12.7 5.7 6.5

8.4 Bizygo Au. Para. Homo1 Au.+hab Homo2 H.sap. N 3 4 8 5 6 20 Mean 139.4 160.1 133.6 130.6 138.9 122.6 Median 126.0 158.3 134.0 125.2 141.0 123.9 Max. 167.0 179.0 150.0 167.0 150.0 140.5 Min. 125.2 145.0 117.0 117.0 121.6 99.7 Range 41.8 34.0 33.0 50.0 28.4 40.9 SD 23.9 16.1 13.8 20.7 11.4 9.7 CV 17.1 10.1 10.4 15.9 8.2 7.9

364 8.5 Bifmt Au. Para. Homo1 Au.+hab Homo2 H.sap. N 3 4 10 7 6 20 Mean 97.2 109.1 113.4 101.4 119.4 102.4 Median 93.5 111.0 113.4 101.0 119.5 102.8 Max. 106.0 115.4 129.3 115.0 129.3 116.6 Min. 92.0 99.1 100.0 92.0 111.0 88.9 Range 14.0 16.3 29.3 23.0 18.3 27.7 SD 7.7 7.7 10.2 7.7 7.2 6.8 CV 7.9 7.1 9.0 7.6 6.0 6.6

8.6 Bizs Au. Para. Homo1 Au.+hab Homo2 H.sap. N 2 2 2 - 2 20 Mean 54.0 57.1 84.3 - 84.3 56.0 Median 54.0 57.1 84.3 - 84.3 55.9 Max. 58.0 61.3 85.0 - 85.0 67.7 Min. 50.0 53.0 83.5 - 83.5 45.5 Range 8.0 8.3 1.5 - 1.5 22.2 SD - - - - - 6.2 CV - - - - - 11.1

8.7 Zs-zgyi Au. Para. Homo1 Au.+hab Homo2 H.sap. N 1 2 1 - 1 20 Mean 55.6 65.0 61.0 - 61.0 52.2 Median - 65.0 - - - 53.1 Max. - 65.9 - - - 61.2 Min. - 64.0 - - - 42.8 Range - 1.9 - - - 18.4 SD - - - - - 4.4 CV - - - - - 8.5

8.8 Bas- pros Au. Para. Homo1 Au.+hab Homo2 H.sap. N 1 4 7 3 5 20 Mean 127.0 128.6 112.6 108.0 118.2 96.2 Median - 136.0 116.0 105.0 118.0 94.9 Max. - 147.8 121.2 127.0 121.2 109.7 Min. - 98.8 92.0 92.0 116.0 87.5 Range - 44.9 29.2 35.0 5.2 22.2 SD - 20.2 10.5 17.7 2.3 5.9 CV - 15.7 9.3 16.4 2.0 6.1

365 8.9 Nas- pros Au. Para. Homo1 Au.+hab Homo2 H.sap. N 3 4 9 8 5 20 Mean 79.7 94.8 83.2 80.8 88.4 64.3 Median 77.0 93.5 82.0 80.5 82.0 65.1 Max. 91.0 112.0 132.0 100.0 132.0 72.8 Min. 71.0 80.0 66.0 66.0 69.0 53.5 Range 20.0 32.0 66.0 34.0 63.0 19.3 SD 10.3 13.9 20.1 12.5 25.0 5.4 CV 12.9 14.7 24.1 15.4 28.2 8.4

8.10 Nas-br Au. Para. Homo1 Au.+hab Homo2 H.sap. N 2 5 35 5 32 20 Mean 82.7 85.4 108.3 84.5 110.4 109.4 Median 82.7 85.0 111.0 89.0 111.7 108.1 Max. 92.3 97.0 123.0 92.3 123.0 119.5 Min. 73.0 73.0 77.0 73.0 98.0 103.5 Range 19.3 24.0 46.0 19.3 25.0 16.0 SD - 10.6 9.9 8.8 7.0 4.9 CV - 12.4 9.1 10.5 6.3 4.4

8.11 Br-lam Au. Para. Homo1 Au.+hab Homo2 H.sap. N 5 4 47 12 40 20 Mean 74.0 78.4 99.2 77.5 102.6 109.5 Median 73.5 79.4 101.0 77.0 103.5 108.7 Max. 77.0 80.0 119.0 91.0 119.0 126.7 Min. 70.3 75.0 68.0 68.0 82.0 96.2 Range 6.7 5.0 51.0 23.0 37.0 30.5 SD 2.6 2.4 12.0 6.5 9.2 7.3 CV 3.6 3.0 12.1 8.4 9.0 6.7

8.12 Lam- opn Au. Para. Homo1 Au.+hab Homo2 H.sap. N 4 4 37 10 31 20 Mean 58.8 60.0 85.1 68.7 87.0 97.4 Median 58.0 60.0 85.0 66.7 86.0 95.0 Max. 62.0 64.0 107.0 98.9 107.0 117.3 Min. 57.0 56.0 65.4 57.0 71.0 83.3 Range 5.0 8.0 41.6 41.9 36.0 34.1 SD 2.4 3.7 9.6 12.6 8.0 8.9 CV 4.0 6.1 11.3 18.3 9.2 9.1

366 8.13 Biecm Au. Para. Homo1 Au.+hab Homo2 H.sap. N 3 4 7 7 3 20 Mean 74.7 74.0 66.5 70.3 66.0 62.9 Median 79.0 74.4 66.0 67.0 66.0 64.5 Max. 82.0 81.0 70.0 82.0 66.8 69.5 Min. 63.0 66.0 65.0 63.0 65.0 55.7 Range 19.0 15.0 5.0 19.0 1.8 13.8 SD 10.2 6.5 1.7 7.3 0.9 3.9 CV 13.7 8.8 2.6 10.4 1.4 6.1

8.14 Bienm Au. Para. Homo1 Au.+hab Homo2 H.sap. N 2 3 11 4 9 20 Mean 39.2 35.6 38.5 37.1 39.3 40.1 Median 39.2 37.0 39.0 37.7 39.5 40.8 Max. 41.0 38.2 44.0 41.0 44.0 44.7 Min. 37.4 31.6 32.0 32.0 34.0 35.1 Range 3.7 6.6 12.0 9.0 10.0 9.6 SD - 3.5 3.5 3.7 3.1 2.8 CV - 9.9 9.2 10.1 7.9 6.9

8.15 Ol-sta Au. Para. Homo1 Au.+hab Homo2 H.sap. N 2 3 2 - 2 20 Mean 70.4 69.6 57.0 - 57.0 46.0 Median 70.4 71.4 57.0 - 57.0 45.3 Max. 75.0 79.1 59.0 - 59.0 55.0 Min. 65.9 5.3 55.0 - 55.0 39.4 Range 9.1 20.8 4 - 4 15.6 SD - 10.5 - - - 4.2 CV - 15.1 - - - 9.1

“Fossil finds usually represent at best very inadequate samples of a population so that it is impossible to determine whether a given specimen stands near the average of its species or happens to be an extreme variation” (Schultz 1963: p.85)1, they are [the fossils] nonetheless the only evidence we have of evolutionary events from the ancient past; and must be interpreted accordingly (e.g. Aiello et al, 2000; Smith, 2005; Wolpoff & Lee, 2006).

1 Similarly, “However, we are forced to interpret fossils in terms of the familiar forms alive today and these might look very different from the fossils that confront us. Given this difficulty, it is best not to make the act of interpretation harder by expecting fossils to conform to preconceived stories about the course that evolution may have taken. As far as we are able, we should try to see fossils as they are, not as we want them to be” (Gee, 2000: p. 64).

367 Furthermore, when dealing with the fossil record, large sample sizes are generally rare and normal distributions are difficult to achieve for statistical analyses. Hominin samples variation, as shown by standard deviations and coefficients of variation values, are within range of that recorded for extant genera (monkeys or apes). In fact, most estimates of variation are small in comparison. Cranial variables with low variation include biporionic breadth (Table 8.3, bipor, CV 2.9%) for the Australopithecus sample, or the superior facial breadth (bifmt) and length (bas-pros) for the Homo2 sample (Tables 8.5 and 8.8, CV 6.0% and 2.0%, respectively). In contrast, cranial variables with inflated variation include bizygomatic breadth (bizygo) for the Australopithecus sample, and the superior facial length (Table 8.8, bas- pros; CV 17.1%) for Paranthropus and Homo1 sample; although each is derived from small sample sizes. Interestingly, bizygomatic breadth for the Au.+hab sample also generates inflated variation (Table 8.4, bizygo, CV 20.7%). Figures 8.2 to 8.13 are box-plots of cranial variables for the hominins. Similar to previous box-plot for other catarrhine groups, some figures for the hominins also feature cranial variables that increase with body weight. For example, Figures 8.9, 8.10 and 8.11 are box-plots of the frontal (nas-br), parietal (br-lam) and occipital (lam-opn) sagittal chords for the hominins which demonstrate a positive scaling trend with increasing body size (see below). Figures 8.4, 8.6 and 8.13 are box-plots of biporionic (bipor), bifrontomalartemporale (bifmt) and palatal breadths for the hominins which demonstrate some overlap of samples. For biporionic breadth (Figure 8.4), Paranthropus, Homo1 and Homo2 samples overlap, whereas, the Australopithecus, Au.+hab and H.sap. samples overlap. This arrangement is similar to bifrontomalartemporale breadth (bifmt), although Paranthropus overlaps with all other hominin genera except Australopithecus. In contrast, there were also some cranial variables which displayed a decrease in size with an increase in body weight. For example, Figures 8.5, 8.7, 8.8, 8.12 and 8.13 are box-plots of bizygomatic breadth (bizygo), superior facial length (nas-pros) and height (nas-pros), maxillo-alveolar (biecm) and palatal breadth (bienm) for the hominins which demonstrate a negative (or inverse) scaling with increasing body size. Overlap of genera can be observed in cranial variables such as biporionic (Figure 8.4) and palatal breadths (Figure 8.13).

368

Box Plots for cranial variables of fossil hominin genera:

Homo1 Homo2 220

210

200 H.sap. 19 0

18 0 Para. Au. Au.+hab. mm 17 0

16 0

15 0

14 0

13 0

1 2 3 4 5 6 7 Figure 8.2: Cranial vault length, Glabella to opisthiocranion.

Homo1 Homo2

14 0

13 0 H.sap.

12 0

110 mm

10 0 Para.

90 Au. Au.+hab.

70 1 2 3 4 5 6 7 Figure 8.3: Biasterionic Breadth.

369 Para. Homo1 Homo2 Au.

13 0 Au.+hab. H.sap.

12 0 mm

110

10 0

1 2 3 4 5 6 7 Figure 8.4 Biporionic Breadth.

Para. 18 0 Au. 17 0

16 0 Homo1 Homo2 15 0 H.sap.

14 0 mm Au.+hab. 13 0

12 0

110

10 0

1 2 3 4 5 6 7 Figure 8.5 Bizygomatic Breadth.

370 Homo1 Homo2 13 0

12 0 H.sap. Para. Au.+hab.

110 mm

10 0 Au.

1 2 3 4 5 6 7 Figure 8.6 Bifrontomalartemporale Breadth.

Para.

14 0

13 0 Au.+hab. Homo1 Homo2

12 0

mm H.sap.

110

10 0

1 2 3 4 5 6 Figure 8.7 Superior Facial Length, Basion to prosthion.

371 Homo1 Homo2

13 0

12 0 Para.

110 Au. Au.+hab. 10 0

mm 90

80 H.sap.

1 2 3 4 5 6 7 Figure 8.8 Superior Facial Height, Nasion to prosthion.

Homo1 Homo2 H.sap. 12 0

110

10 0 Para.

mm Au. Au.+hab.

70 1 2 3 4 5 6 7 Figure 8.9 Frontal Sagittal Chord, Nasion to bregma.

372 H.sap. 13 0

2 Homo1 Homo 12 0

110

10 0

mm Au.+hab.

90 Para.

80 Au.

1 2 3 4 5 6 7 Figure 8.10 Parietal Sagittal Chord, Bregma-lambda.

12 0 H.sap.

2 110 Homo Homo1 Au.+hab. 10 0

90 mm

70 Para. Au.

50 1 2 3 4 5 6 7 Figure 8.11 Occipital Sagittal Chord, Lambda-opisthion.

373 90

Au. Au.+hab. Para.

H.sap. 1 70 Homo 2 mm Homo

1 2 3 4 5 6 7 Figure 8.12 Maxillo-alveolar Breadth, Biecm.

48 Homo2 H.sap. 46 Homo1 44 Au. Au.+hab. 42 Para. 40 mm

30 A B E F C D

Figure 8.13 Palatal Breadth, Bienm.

374 8.2.1 Kruskal-Wallis and Mann-Whitney results for cranial variables: Table 8.16 provides the Kruskal-Wallis and post-hoc Mann-Whitney results for the Hominin cranial variables. Many significant and non-significant differences were revealed between hominin genera. Most comparisons of fossil hominin samples to the H.sap. sample were significantly different. However, generic sample comparisons that were not significantly different to H.sap. included; Cranial vault length (g-o, Table 8.16, no. 1) for Paranthropus; Biporionic breadth (bipor, no. 3) of Australopithecus, Paranthropus and Au.+hab; Bizygomatic breadth (bizygo, no. 4) for Australopithecus, Homo1 and Au.+hab; Bifrontomalartemporale breadth (bifmt, no. 5) for all hominin genera except Homo1; Superior facial length (bas-pros, no. 6) for Au.+hab; frontal sagittal chord (nas-br, no. 8) for Homo1 and for Homo2; and Maxillo-alveolar breadth (biecm, no. 11) for Australopithecus and Homo2. Additionally, all cranial variables for the Homo1 and Homo2 samples were not significantly different to each other, except for the occipital sagittal chord (lam-opn, Table 8.16, no. 10). There were no significant differences between all hominin genera with regard to palatal breadth (bienm, no. 12). Australopithecus and Paranthropus were not significantly different with respect to their cranial vault length (g-o), biporionic and bifrontomalartemporale breadths (bipor and bifmt), superior facial height (nas-pros), parietal and occipital sagittal chords (br-lam) and maxillo-alveolar and palatal breadths (biecm and bienm). Interestingly, the cranial length (g-o) sample for the Australopithecus was significantly different to Homo1 but not the Au.+hab sample (Table 8.16, no. 1). This outcome is similar to comparisons for bifrontomalartemporale breadth (bifmt, Table 8.16, no. 5). Again, the sample for the Australopithecus was significantly different to Homo1 but not from the Au.+hab sample for this variable.

375 Table 8.16: Kruskal-Wallis results for cranial variables of hominin genera with Mann- Whitney pairwise comparisons (p (same)).

Au. Para. Homo1 Au.+hab. Homo2 H.sap. 1. G-o - 0.551 0.008 0.897 0.006 0.020 Au. Hc: 50.8 - 0.001 0.648 0.000 0.027 Para. p=<0.000 - 0.000 0.601 0.000 Homo1 - 0.000 0.002 Au.+hab. - 0.000 Homo2 - H.sap. 2. Biast - 0.061 0.001 0.885 0.001 0.002 Au. Hc: 46.3 - 0.003 0.061 0.003 0.002 Para. p=<0.000 - 0.001 0.997 0.000 Homo1 - 0.001 0.002 Au.+hab. - 0.000 Homo2 - H.sap. 3. Bipor - 0.178 0.123 0.471 0.110 0.230 Au. Hc: 19.7 - 0.749 0.391 0.458 0.110 Para. p=0.01 - 0.134 0.521 0.002 Homo1 - 0.064 0.561 Au.+hab. - 0.000 Homo2 - H.sap. 4. Bizygo - 0.216 0.760 0.456 0.897 0.294 Au. Hc: 15.16 - 0.051 0.066 0.110 0.002 Para. p=0.010 - 0.608 0.478 0.088 Homo1 - 0.235 0.865 Au.+hab. - 0.010 Homo2 - H.sap. 5. Bifmt - 0.112 0.052 0.569 0.028 0.338 Au. Hc: 19.64 - 0.480 0.186 0.110 0.112 Para. p=0.001 - 0.036 0.303 0.015 Homo1 - 0.008 0.561 Au.+hab. - 0.000 Homo2 - H.sap.

376 Au. Para. Homo1 Au.+hab. Homo2 H.sap. 6. Bas-pros ------Au. Hc: 20.52 - 0.156 0.216 0.270 0.006 Para. P<0.000 - 0.820 0.465 0.004 Homo1 - 0.551 0.254 Au.+hab. - 0.000 Homo2 - H.sap. 7. Nas-pros - 0.471 0.537 0.552 1 0.005 Au. Hc: 27.74 - 0.190 0.203 0.391 0.002 Para. P<0.000 - 0.847 0.842 0.000 Homo1 - 0.884 0.002 Au.+hab. - 0.002 Homo2 - H.sap. 8. Nas-br ------Au. Hc: 25.28 - 0.001 0.754 0.000 0.000 Para. P<0.000 - 0.001 0.551 0.979 Homo1 - 0.000 0.000 Au.+hab. - 0.554 Homo2 - H.sap. 9. Br-lam - 0.066 0.000 0.246 0.000 0.000 Au. Hc: 55.31 - 0.005 0.431 0.001 0.002 Para. P<0.000 - 0.000 0.261 0.001 Homo1 - 0.000 0.000 Au.+hab. - 0.009 Homo2 - H.sap. 10. Lam-opn - 0.773 0.014 0.104 0.001 0.002 Au. Hc: 45.93 - 0.014 0.157 0.001 0.002 Para. P<0.000 - 0.212 0.01 0.005 Homo1 - 0.000 0.000 Au.+hab. - 0.000 Homo2 - H.sap. 11. Biecm - 0.860 0.494 0.732 0.663 0.132 Au. Hc: 16.44 - 0.073 0.450 0.157 0.005 Para. P=0.006 - 0.553 0.732 0.014 Homo1 - 0.494 0.014 Au.+hab. - 0.132 Homo2 - H.sap. 12. Bienm - 0.3865 0.9214 0.817 0.9062 0.6073 Au. Hc: 6.77 - 0.2129 0.5959 0.1391 0.0613 Para. P>0.24 - 0.5139 0.6485 0.1797 Homo1 - 0.3545 0.1123 Au.+hab. - 0.4367 Homo2 - H.sap.

377 8.2.2 Summary of cranial variables In summary, despite small sample sizes and a small number of cranial variables, there are some clear differences in size between hominin genera, as well as visual positive and negative scaling trends (Figures 8.2 to 8.13). In addition, the levels of variation produced by these hominin samples are overwhelmingly within the range of extant catarrhines (Tables 8.1 to 8.15).

8.3 Descriptive Statistics and Univariate Analyses for cranial indices: Very few fossils were complete enough to generate cranial indices with reasonable sample sizes. Tables 8.17 to 8.19 provide the descriptive statistics for cranial indices for three Hominin genera. Similar to cranial variables, the available cranial indices also produced low to moderate levels of variation. The two largest standard deviation and coefficients of variation values were produced by the Paranthropus, Au.+hab and Homo2 samples. The former produced a CV of 14.4% for the relative proportion of superior facial height (nas-pros) to biporionic breadth (bipor, Table 8.18); the second generated a CV of 15.6% for the relative proportion of superior facial height (nas-pros) to biasterionic breadth (biast, Table 8.17); and the latter produced a CV of 14.7% for the relative proportion of superior facial breadth (bifmt) to bizygomaxillare inferior breadth (bizi, Table 8.19). Figures 8.14 to 8.16 are box-plots for three cranial indices for the hominin genera. Figure 8.14 displays the relative proportion of superior facial height (nas-pros) to biasterionic breadth (biast). Figure 8.15 displays the relative proportion of superior facial height (nas-pros) to biasterionic breadth (biast). Finally, Figure 8.16 displays the relative proportion of superior facial breadth (bifmt) to bizygomaxillare inferior breadth (bizi). Figure 8.14 is interesting because like some cranial variables (see above), there is a decrease in the relative proportion of superior facial height to biasterionic breadth (biast) with an increase in body size. In contrast, there is a slight increase in the relative proportion of superior facial breadth to bizygomaxillare inferior breadth with increasing body. There was overlap between the Homo samples for each index.

378 Tables 8.17 – 8.19: Descriptive Statistics of cranial indices for the fossil hominin genera (Au. – Australopithecus spp.; Para. – Paranthropus spp.; Homo1 – H. habilis, H. rudolfensis, H. erectus, H. ergaster, H. heidelbergensis and H. neanderthalensis; Au.+hab – Australopithecus spp., H. habilis and H. rudolfensis; Homo2 – H. erectus, H. ergaster, H. heidelbergensis and H. neanderthalensis; H.sap. – modern H. sapiens).

8.17 Bas-pros/ biast Au. Para. Homo1 Au.+hab Homo2 H.sap. N 2 4 8 4 6 20 Mean 125.2 110.6 94.2 106.2 91.0 90.5 Median 134.7 113.6 101.0 134.7 101.0 105.0 Max. 115.7 109.9 82.6 95.7 82.6 81.5 Min. 19.0 3.7 18.3 39.0 18.3 23.4 Range 13.4 1.6 6.0 18.4 6.6 6.9 SD 10.7 1.5 6.5 16.6 7.2 7.6 CV 8.55 1.36 6.90 15.63 7.91 8.40

8.18 Bas-pros/ bipor Au. Para. Homo1 Au.+hab Homo2 H.sap. N 2 5 5 4 3 20 Mean 99.2 81.8 89.3 91.0 89.3 86.2 Median 106.3 89.3 91.2 106.3 91.2 102.1 Max. 92.0 66.2 80.7 84.8 80.7 75.5 Min. 14.3 23.2 10.6 21.6 10.6 26.6 Range 10.1 9.3 4.4 9.2 5.6 7.9 SD 10.2 11.8 5.0 9.9 6.5 9.0 CV 10.28 14.43 5.60 10.88 7.28 10.44

8.19 Bifmt/ bizi Au. Para. Homo1 Au.+hab Homo2 H.sap. N 3 4 7 7 3 20 Mean 95.8 91.4 105.2 116.3 111.8 105.7 Median 98.2 101.9 116.3 129.6 112.3 115.4 Max. 88.2 85.8 85.7 99.0 85.7 91.5 Min. 9.9 16.1 30.6 30.6 26.6 23.9 Range 5.2 7.3 10.4 11.2 15.2 7.4 SD 5.5 7.9 9.9 9.9 14.7 7.1 CV 5.74 8.64 9.41 8.51 13.15 6.72

379 Box Plots for cranial indices of fossil hominin genera:

Au. Au.+hab.

13 0

12 0 Para.

% 110 H.sap. Homo1 Homo2

10 0

1 2 3 4 5 6 7 Figure 8.14: Bas-pros/biast x 100.

110 Au. Au.+hab. H.sap.

10 0

Homo1 Homo2 Para. 90 %

1 2 3 4 5 6 7 Figure 8.15: Bas-pros/bipor x 100.

380 Au.+hab. 13 0

1 12 0 Homo H.sap. Homo2

110 % Para. Au. 10 0

1 2 3 4 5 6 7 Figure 8.16: Bifmt/bizi x 100.

8.3.1 Kruskal-Wallis and Mann-Whitney results for cranial indices: Table 8.20 provides the Kruskal-Wallis and Mann-Whitney results for hominin cranial indices. The H.sap sample is significantly different from the Australopithecus and Paranthropus samples in regard to the relative proportion of superior facial height (nas- pros) to biasterionic breadth (biast) but not significantly different to the Homo1 and Homo2 samples (Table 8.20, no. 1). All generic comparisons for the second index (bas-pros/bipor, Table 8.20, no. 2) were not significantly different except for the Australopithecus versus Au.+hab samples. Similarly, all generic comparisons for the last index (bifmt/bizi, Table 8.20, no. 3) were not significantly different except for, the Au.+hab versus Australopithecus samples; the Au.+hab versus Paranthropus samples; and the modern H.sap versus both Homo1 and Homo2 samples.

8.3.2 Summary of cranial indices To summarize, with the few cranial indices presented, little can be said of relative proportions for fossil hominins. Still, the differences produced from the Australopithecus and Au.+hab were interesting. For the first index (Figure 8.14, bas-pros/biast), the

381

Table 8.20: Kruskal-Wallis results for cranial indices of hominin genera with Mann- Whitney pairwise comparisons (p (same)).

Au. Para. Homo1 Au.+hab. Homo2 H.sap. 1. Bas-pros/ biast ------Au. Para. Hc: 15.37 - 0.008 0.885 0.014 0.002 p=0.004 - 0.075 0.747 0.492 Homo1

- 0.070 0.014 Au.+hab. 2 - 0.927 Homo

- H.sap. 2. Bas-pros/ bipor ------Au. Hc: 5.60 - 0.144 0.037 0.371 0.096 Para. p=0.231 - 0.270 0.882 0.973 Homo1 - 0.377 0.300 Au.+hab. - 0.891 Homo2 - H.sap. 3. Bifmt/ bizi - 0.596 0.111 0.023 0.663 0.032 Au. Hc: 12.25 - 0.156 0.030 0.596 0.022 Para. 1 p=0.032 - 0.201 0.909 0.934 Homo

- 0.362 0.081 Au.+hab. - 0.964 Homo2 - H.sap.

Australopithecus sample was clearly separated from the other genera but with the inclusion of H. habilis sensu lato the amount of variation increases dramatically. In addition, for the third index (Figure 8.16, bifmt/bizi), again the Australopithecus sample produced some of the smaller percentages but with the inclusion of H. habilis sensu lato the amount of variation increased.

8.4 Bivariate Results: Table 8.21 provides the bivariate (regression) results of plotting generic hominin medians for some cranial dimensions against generic mean body weight. Figure 8.17 is a bivariate plot of the generic median for the frontal sagittal chord (nas-br; Table 8.10; Figure 8.9) against generic mean body weight estimates (Collard & Wood, 1999). Six out of fifteen cranial variables were significantly correlated with body size with an adjusted R 382 squared value >0.60 and a p-value of <0.05; cranial vault length (g-o), biasterionic breadth (biast), frontal, parietal and occipital sagittal chords (nas-br, br-lam and lam-opn) and palatal length A (ol-sta). These results were very similar to the bivariate results for the Colobinae. This is because small hominins (i.e. Australopithecus and Paranthropus) have large cranial dimensions (particularly facial dimensions), whereas the larger hominins (i.e. Homo1, Homo2 and H.sap.) have small cranial dimensions (again, particularly facial and palatal dimensions). However, it is important to note that the cranial variables that were significantly correlated with body weight were neurocranial dimensions (excluding palatal length). This would suggest that while Australopithecus and Paranthropus had large facial features, they still experienced some increase in neurocranial dimensions or encephalization like that witnessed in the lineage leading to modern humans (see also Elton et al, 2001). Also of particular note is the distance between Australopithecus, Paranthropus and Au.hab from the Homo samples. In general, body size has been an important feature between catarrhine genera for this study.

383 Table 8.21: Results of generic mean body weight estimates (Ln (g)) plotted against the generic median for particular cranial dimensions (Ln (mm)) those in bold represent cranial variables with an adjusted R squared value >0.60 and a p-value <0.05).

Adjusted P-value/ Mean Generic Body Weight (g) Ln (mm) R squared Significance F (Collard & Wood, 1999): G-o 0.69 0.03 Au. - 45466 Biast 0.69 0.03 Para. - 42133 Bipor 0.25 0.18 Homo1 - 58500 Bizygo -0.21 0.76 Au.hab - 42688 Bifmt 0.10 0.28 Homo2 - 67100 Bizs 0.18 0.27 H.sap - 63500 Zs-zgyi -0.19 0.59 Bas-pros -0.14 0.53 Nas-pros 0.05 0.32 Nas-br 0.86 0.004 Br-lam 0.88 0.004 Lam-opn 0.82 0.008 Biecm 0.34 0.13 Bienm 0.44 0.09 Ol-sta 0.64 0.06

4.75 2 Figure 8.17: Adjusted R2: 0.86 Homo1 Homo 4.70 Significance F & P-value 0.004 H.sap 4.65 Ln y = 0.6365x - 2.3391 Frontal 4.60 Sagittal 4.55 Chord 4.50 Au. + hab (nas-br) 4.45 Para. Au. 4.40

4.35 10.6 10.7 10.8 10.9 11 11.1 11.2

Ln Body Weight (g)

384 8.5 Multivariate Statistics: 8.5.1 PCA results for hominins (Au., Para., Homo1 and H. sap), Ln transformed data: This multivariate analysis reflects the tradition taxonomic scheme with habilis/rudolfensis combined with other extinct species (H. erectus, H. ergaster, H. hiedelbergensis, and H. neanderthalensis) generally attributed to Homo. To identify the underlying structure (i.e. the cranial variables which account for the most of the variation between genera) of the Hominin multivariate dataset derived from Ln transformed generic medians, PCA was employed. By doing so, PCA based on the variance-covariance matrix resulted in the first (Eigenvalue – 0.05, 70.91%), second (Eigenvalue – 0.02, 24.28%) and third (Eigenvalue – 0.003, 4.81%) PCs accounting for 95.19% of the variation within the tribe (Figures 8.18 and 8.19); 100% of the variation was explained by the first three PCs. Table 8.22 lists the low to moderate variable loadings and PCs 1 to 3 all have positive and negative loadings (fifteen measurements); as such PC 1 can not be interrupted as size differences but shape. For the first PC the largest positively loaded cranial variables were palatal length A (ol-sta) and superior facial length (bas-pros) at 0.40 and 0.26, respectively. Contrasting with these were negatively loaded cranial variables such as the occipital and parietal sagittal chords (lam-opn and br-lam) at -0.50 and -0.37, respectively (Table 8.22). The largest positively loaded cranial variable for the PC 2 was palatal breadth (bienm) at 0.13. Juxtaposed against this cranial variable were negatively loaded cranial variables such as bizygomaxillare superior breadth (bizs) and superior facial height (nas- pros) at -0.55 and -0.41, correspondingly (Table 8.22). The third PC’s largest positively loaded cranial variable was bizygomaxillare superior (bizs) with 0.62. Contrasting with this dimension were negatively loaded cranial variables such as bizygomatic and bifrontomalartemporale breadths with -0.54 and -0.28, respectively. In both scatterplots, PCs 1 & 2 and PCs 2 & 3, all four generic samples were well separated from each other (Figures 8.18 and 8.19).

385 Palatal breadth

Bizs & superior facial height

Occipital & Parietal Palatal length & superior sagittal chords facial length

Figure 8.18: Hominin scatterplot of PC 1 & 2 of Ln transformed generic medians for conventional taxonomic scheme.

Bizygomaxillare Superior

Bizygo- matic Bizs & breath superior facial height Palatal breadth

Figure 8.19: Hominin scatterplot of PC 2 & 3 of Ln transformed generic medians for conventional taxonomic scheme.

386

Table 8.22: PCA results for hominin genera Ln transformed generic medians for conventional taxonomic scheme.

Eigenvalue %Variance Cum. % PC1 0.045628 70.912 70.91 PC2 0.015622 24.279 95.19 PC3 0.003095 4.8096 100 PC4 5.06E-18 7.87E-15

PC1 PC2 PC3 g-o -0.2033 -0.2094 -0.03825 biast -0.2976 -0.2817 -0.0429 bipor -0.09025 -0.2448 -0.1711 bizygo 0.09937 -0.255 -0.5372 bifmt -0.07028 -0.2504 -0.276 bizs -0.218 -0.5571 0.6262 zs-zygi 0.06538 -0.2782 -0.2247 bas-pros 0.26 -0.1975 0.139 nas-pros 0.1809 -0.4093 -0.2248 nas-br -0.3111 -0.07582 0.103 br-lam -0.3787 0.004182 -0.171 lam-opn -0.5013 0.07256 -0.06254 biecm 0.1934 0.0306 0.1194 bienm -0.041 0.1257 0.05729 ol-sta 0.4055 -0.2591 0.17

In summary, reminiscent of previous results for the catarrhines, PCA resulted in a contrast between facial and palatal dimensions (with positive loadings for PC 1) with neurocranial measurements (with negative loadings PC 1, Table 8.22). While Australopithecus and Paranthropus species have relatively wide and long faces, the Homo species display smaller facial features combined with large neurocranial lengths.

8.5.2 PCA results for hominins (Au., Para., Homo1 and H. sap), MSV: PCA based on the variance-covariance resulted in the first (Eigenvalue – 0.24, 83.18%) and second (Eigenvalue – 0.03, 9.34%) PCs accounting for 92.52% of the variation within the tribe (2.67% less variation explained than PCs 1 & 2 for Ln transformed data but 63.43% more variation explained than PCs 2 & 3 for Ln transformed data; Figure 8.20); 100% of the variation was explained by the first three PCs. The variable loadings for the first PC were positive and negative (Table 8.23., fifteen measurements),

387 Biecm breadth & Occipital sagittal chord

Occipital & Parietal Bizs sagittal Superior facial chords length

Figure 8.20: Hominin scatterplot of PC 1 & 2 of generic median MSVs for conventional taxonomic scheme. Table 8.23: PCA results for hominin genera Ln transformed generic medians for taxonomic scheme placing H. habilis/rudolfensis specimens in Australopithecus.

Eigenvalue %Variance Cum. % PC1 0.242482 83.181 83.18 PC2 0.0272227 9.3384 92.52 PC3 0.0218067 7.4806 100 PC4 1.31536E-17 4.5122E-15

PC1 PC2 PC3 g-o -0.278 -0.1792 -0.00115 biast -0.2643 -0.2887 0.02321 bipor -0.03165 -0.1591 -0.1389 bizygo 0.2694 -0.1833 -0.6949 bifmt -0.003335 -0.1621 -0.242 bizs -0.1055 -0.5171 0.4925 zs-zygi 0.09125 -0.08904 -0.1063 bas-pros 0.4286 0.1891 0.2321 nas-pros 0.2353 -0.2827 -0.1491 nas-br -0.2919 0.1311 0.07329 br-lam -0.3549 0.2598 -0.2131 lam-opn -0.4033 0.3007 -0.1465 biecm 0.1994 0.404 0.1259 bienm -0.001774 0.25 0.01391 ol-sta 0.3274 0.0865 0.1428

388 which again vary from low to moderate, and thus cannot be interpreted as size differences but instead shape dissimilarities; the second PC also had mixed loading polarities. The largest positively loaded cranial variables for PC 1 were superior facial length (bas-pros) and palatal length A (ol-sta) at 0.42 and 032, respectively. Contrasting with these were negatively loaded cranial variables such as occipital and parietal sagittal chords (lam- opn and br-lam) with -0.40 and -0.35, correspondingly. The largest positively loaded cranial variables for PC 2were maxillo-alveolar breadth (biecm) and the occipital sagittal chord (lam-opn) at 0.40 and 0.30. Juxtaposed against these was the negatively loaded bizygomaxillare superior breadth (bizs) with -0.52. The scatterplot for PCs 1 & 2 (Figure 8.20) is very similar to PCs 1& 2 for Ln transformed data (Figure 8.18). In summary, PCA resulted in contrasting facial and palatal dimensions (with positive loadings for PC 1) with neurocranial measurements (with negative loadings PC 1; Table 8.23). Thus, shape variables find the same situation as those including size (Ln).

8.5.3 PCA results for hominins (Au.+hab, Para., Homo2 and H. sap), Ln transformed data: This multivariate analysis reflects the suggestions and conclusions reached by Wood & Collard (1999a) which places habilis/rudolfensis in Australopithecus. To identify the underlying structure (i.e. the cranial variables which account for the most of the variation between genera) of the Hominin multivariate dataset derived from Ln transformed generic medians, PCA was employed. By doing so, PCA based on the variance-covariance matrix resulted in the first (Eigenvalue – 0.03, 70.58%), second (Eigenvalue – 0.01, 27.72%) and third (Eigenvalue – 0.00, 1.69%) PCs accounting for 100% of variation within the tribe (Figures 8.21 and 8.22). Table 8.24 lists the low to moderate variable loadings and all PCs have mixed polarities (twelve measurements). The largest positively loaded cranial variables for PC 1 were the occipital and parietal sagittal chords (br-lam and lam-opn) at 0.56 and 0.46, respectively. Juxtaposed against these were negatively loaded cranial dimensions such as superior facial height and length (nas-pros and bas-pros) with -0.31 and -0.20, correspondingly.

389 Superior facial length & Bizygo

Superior facial height & Occipital & Parietal length sagittal chords

Figure 8.21: Hominin scatterplot of PC 1 & 2 of Ln transformed generic medians for taxonomic scheme placing H. habilis/rudolfensis specimens in Australopithecus.

Bizygo & Parietal sagittal chord

Superior facial height & Superior facial nas-br length & Bizygo

Figure 8.22: Hominin scatterplot of PC 2 & 3 of Ln transformed generic medians for taxonomic scheme placing H. habilis/rudolfensis specimens in Australopithecus.

390

Table 8.24: PCA results for hominin genera generic median MSVs for taxonomic scheme placing H. habilis/rudolfensis specimens in Australopithecus.

Eigenvalue %Variance Cum. % PC1 0.027327 70.582 70.58 PC2 0.010733 27.723 98.3 PC3 0.000656 1.6945 100 PC4 5.47E-18 1.41E-14

PC1 PC2 PC3 g-o 0.2357 0.338 0.02806 biast 0.3131 0.379 -0.1598 bipor 0.06848 0.3181 0.02179 bizygo -0.1501 0.3974 0.5455 bifmt 0.03065 0.3141 -0.2444 bas-pros -0.2034 0.453 -0.04125 nas-pros -0.3131 0.3691 -0.3769 nas-br 0.3497 0.1008 -0.3607 br-lam 0.4553 0.1252 0.4221 lam-opn 0.5616 -0.06871 -0.01232 biecm -0.1372 0.1105 0.4045 bienm 0.1225 -0.00732 0.009972

The largest positively loaded cranial variables for PC 2 were superior facial length (bas-pros) and bizygomatic breadth (bizygo) at 0.45 and 0.40, respectively. Contrasting with these is one very low negatively loaded dimension, the occipital sagittal chord (lam- opn) with -0.07. The third PC’s largest positively loaded cranial variables were bizygomatic breadth (bizygo) and parietal sagittal chord (br-lam) at 0.55 and 0.42, respectively. Juxtaposed against these were negatively loaded dimensions such as superior facial height (nas-pros) and frontal sagittal chord (nas-br) with -0.38 and -0.36, correspondingly. The scatterplots for PCs 1 & 2 and PCs 2 & 3 (Figures 8.22 and 8.23) are very similar to those for Ln transformed data (Figures 8.21 and 8.22). Again, all four genera are well separated from each other. Contrasts in face and neurocranium are clear.

391 8.5.4 PCA results for hominins (Au.+hab, Para., Homo2 and H. sap), MSV: PCA based on the variance-covariance resulted in the first (Eigenvalue – 0.13, 89.24%) and second (Eigenvalue – 0.01, 7.60%) PCs accounting for 96.84% of variation within the tribe (1.46% less variation explained than PCs 1 & 2 for Ln transformed data but 67.43% more variation explained than PCs 2 & 3 for Ln transformed data; Figure 8.23). One hundred percent of the variation is explained by the first three PCs. The low to moderate variable loadings for the first PC showed both positive and negative loadings (Table 8.25, twelve measurements). The largest positively loaded cranial variables for PC 1 were the occipital and parietal sagittal chords at 0.41 and 0.36, respectively. Contrasting with these were negatively loaded measurements such as bizygomatic breadth (bizygo) and superior facial length (bas-pros) with -0.44 and -0.42, correspondingly. The largest positively loaded cranial variables for PC 2 were cranial vault length (g- o) and biasterionic breadth (biast) at 0.60 and 0.53, respectively. Juxtaposed against these were negatively loaded measurements such as maxillo-alveolar breadth (biecm) and the occipital sagittal chord (lam-opn) with -0.34 and -0.23, correspondingly. The scatterplot for PCs 1 & 2 (Figure 8.23) is nearly identical to PCs 1& 2 for Ln transformed data (Figure 8.21).

392 Cranial vault length & Biast

Biecm & Occipital sagittal chord Occipital & Parietal sagittal chords

Bizygo & Superior facial length Figure 8.23: Hominin scatterplot of PC 1 & 2 of generic median MSVs for taxonomic scheme placing H. habilis/rudolfensis specimens in Australopithecus. Table 8.25: PCA results for hominin genera generic median MSVs for taxonomic scheme placing H. habilis/rudolfensis specimens in Australopithecus.

Eigenvalue %Variance Cum. % PC1 0.134155 89.244 89.24 PC2 0.011428 7.6019 96.84 PC3 0.004742 3.1542 100 PC4 7.54E-18 5.01E-15

PC1 PC2 PC3 g-o 0.1982 0.5975 -0.05424 biast 0.1942 0.5302 0.1291 bipor -0.06757 0.1705 0.002624 bizygo -0.438 0.1917 -0.6739 bifmt -0.1051 0.117 0.2716 bas-pros -0.4177 0.2323 0.02147 nas-pros -0.3771 -0.08151 0.2693 nas-br 0.2721 -0.0779 0.2342 br-lam 0.3588 0.03296 -0.4909 lam-opn 0.4081 -0.2254 -0.2203 biecm -0.1706 -0.3432 -0.1874 bienm 0.02088 -0.2146 0.007721

393 8.5.5 PCA results for hominins (Au., Para., Homo1 and H. sap) and other catarrhine genera, Ln transformed data: This multivariate analysis reflects the tradition taxonomic scheme with habilis/rudolfensis combined with other extinct species (H. erectus, H. ergaster, H. hiedelbergensis, and H. neanderthalensis) generally attributed to Homo. PCA based on the variance-covariance matrix resulted in the first (Eigenvalue – 0.28, 87.16%), second (Eigenvalue – 0.02, 6.5%) and third (Eigenvalue – 0.0006, 1.98%) PCs accounting for 95.64% of the variation of the combined hominin and other catarrhine multivariate dataset (Figure 8.24). Table 8.26 lists the low to moderate variable loadings and all cranial variables (fifteen measurements); PCs 2 and 3 have mixed polarities. Variable loadings are moderate to low showing no clear single variable accounting for variance in these data. That is, they are all quite similar morphometrically. The largest positively loaded cranial variables for PC 1were superior facial height (nas-pros) and the occipital sagittal chord (lam-opn) at 0.33 and 0.31, correspondingly. The highest positively loaded cranial variables for PC 2 were bizygomaxillare superior breadth (bizs) and the occipital sagittal chord (lam-opn) at 0.30 and 0.22, respectively. Contrasting with these are negatively loaded dimensions such as palatal length A (ol-sta) and superior facial height (nas-pros) with -0.52 and -0.51, correspondingly. The largest positively loaded cranial variables for PC 3 were bizygomaxillare superior breadth (bizs) and palatal length A (ol-sta) at 0.37 and 0.27, respectively. Juxtaposed against these were negatively loaded measurements such as palatal breadth (bienm) and occipital sagittal chord (lam-opn) with -0.73 and -0.22, correspondingly. The scatterplot for PCs 1 & 2 (Figure 8.24) was very informative. On the first PC, from left to right there is an increase in body size. Interestingly, the Colobinae, Cercopithecini, Hylobates and the smaller papionins (Lophocebus, Cercocebus and Macaca) were closely grouped together. The larger papionins (Theropithecus, Mandrillus and Papio) were set apart from this cluster, as were the Great apes and Australopithecus and Paranthropus. The H.sap sample and fossil Homo1 were well separated from all other catarrhine genera, including the apes and other hominins (Au. and Para.).

394 Table 8.26: PCA results for hominin genera Ln transformed generic medians for taxonomic scheme not placing H. habilis/rudolfensis specimens in Australopithecus and all catarrhine genera.

Eigenvalue %Variance Cum. % PC1 0.282028 87.163 87.16 PC2 0.02103 6.4993 93.6 PC3 0.006414 1.9824 95.64 PC4 0.004287 1.3251 96.97

PC1 PC2 PC3 g-o 0.2521 0.1478 0.04014 biast 0.277 0.1586 0.1009 bipor 0.2526 0.07028 0.08691 bizygo 0.2432 -0.03113 0.2035 bifmt 0.2216 0.1098 0.2127 bizs 0.2176 0.2938 0.3705 zs-zygi 0.208 0.08691 0.05482 bas-pros 0.2383 -0.3419 0.1254 nas-pros 0.3279 -0.5052 -0.2173 nas-br 0.2013 0.2123 0.08945 br-lam 0.2877 0.3197 -0.1507 lam-opn 0.3106 0.2191 -0.221 biecm 0.2593 -0.03636 0.003501 bienm 0.2603 0.006888 -0.7324 ol-sta 0.2792 -0.522 0.2736

Parietal & Occipital sagittal chords

Facial height & Palatal Facial height & length Occipital sagittal chord

Figure 8.24: Scatterplot of PC 1 & 2 of Ln transformed generic medians for taxonomic scheme not placing H. habilis/rudolfensis specimens in Australopithecus and all catarrhine genera.

395 To summarize, PCA resulted in contrasting facial and palatal dimensions (with negative loadings for PC 2) with neurocranial measurements (with positive loadings PC 2; Table 8.23); bizygomaxillare superior also assisted in separated genera along PC 2. Homo is highly distinct compared to other catarrhines.

8.5.6 PCA results for hominins (Au., Para., Homo1 and H. sap) and other catarrhine genera, MSV: PCA based on the variance-covariance resulted in the first (Eigenvalue – 0.13, 53.34%) and second (Eigenvalue – 0.05, 18.4%) PCs accounting for 71.74% of the variation within the tribe (23.45% less variation explained than PCs 1 & 2 for Ln transformed data but 63.26% more variation explained than PCs 2 & 3 for Ln transformed data; Figure 8.25). The low to moderate variable loadings for the first PC were positive and negative (Table 8.27, 15 measurements). Again, low to moderate loadings showing no one cranial variable is accounting for the variation between genera. The largest positively loaded cranial variables for PC 1 were the superior facial length and height (bas-pros and nas-pros) at 0.53 and 0.52, respectively. Contrasting with these are negatively loaded dimensions such as cranial vault length (g-o) and frontal sagittal chord (nas-br) both with -0.24. The largest positively loaded cranial variables for PC 2 were the bizygomatic and bifrontomalartemporale breadths (bizygo and bifmt) at 0.43 and 0.34, correspondingly. Juxtaposed against these were negatively loaded measurements such as the occipital and parietal sagittal chords (lam-opn and br-lam) with -0.39 and -0.38, respectively. The scatterplot for PCs 1 & 2 (Figure 8.25) was similar to that for Ln transformed data (Figure 8.24). Interestingly, when only shape is examined, the Great apes, Australopithecus and Paranthropus lie much closer to the cluster of monkeys but still, the Homo samples (H.sap and Homo1) are distinctly separated. To summarize, again, PCA resulted in contrasting facial and neurocranial dimensions. However, with MSVs the facial dimensions received positive loadings whereas the neurocranial measurements had negative loadings (Table 8.27). Homo contrasts dramatically with all other catarrhines.

396 Table 8.27: PCA results for hominin genera generic median MSVs for taxonomic scheme not placing H. habilis/rudolfensis specimens in Australopithecus and all catarrhine genera and all catarrhine genera.

Eigenvalue %Variance Cum. % PC1 0.13669 53.339 53.34 PC2 0.04716 18.403 71.74 PC3 0.016412 6.4042 78.14 PC4 0.01315 5.1312 83.27

PC1 PC2 PC3 g-o -0.2435 -0.1058 0.3532 biast -0.1258 -0.1304 0.1961 bipor -0.06417 0.04093 0.1123 bizygo 0.04397 0.4286 0.2231 bifmt -0.1392 0.342 0.1341 bizs -0.1836 0.2057 0.1268 zs-zygi -0.07109 0.224 -0.3507 bas-pros 0.5181 0.3135 0.0018 nas-pros 0.5339 -0.3603 0.1516 nas-br -0.2401 0.1966 0.2284 br-lam -0.2318 -0.3823 0.1264 lam-opn -0.1278 -0.3903 0.06055 biecm 0.02808 -0.003487 -0.01647 bienm 0.01115 -0.1004 -0.6389 ol-sta 0.4225 -0.04071 0.3377

Bizygomatic & Bifmt breadths

Occipital & Parietal sagittal chords Cranial vault Superior Facial length length & height

Figure 8.25: Scatterplot of PC 1 & 2 of generic median MSVs for taxonomic scheme not placing H. habilis/rudolfensis specimens in Australopithecus and all catarrhine genera.

397 8.5.7 PCA results for hominins (Au.+hab, Para., Homo2 and H. sap) and other catarrhine genera Ln transformed data: This multivariate analysis reflects the suggestions and conclusions reached by Wood & Collard (1999a) which places habilis/rudolfensis in Australopithecus. PCA based on the variance-covariance matrix resulted in the first (Eigenvalue – 0.24, 90%), second (Eigenvalue – 0.01, 5.3%) and third (Eigenvalue – 0.005, 1.97%) PCs accounting for 97.25% of the variation of the Hominin and other catarrhine multivariate dataset (Figure 8.26). Table 8.28 lists the low to moderate variable loadings and all variables were positively loaded; PCs 2 and 3 have mixed loadings (twelve measurements). The largest positively loaded cranial variables for PC 1 were the superior facial height (nas-pros) and the occipital sagittal chord (lam-opn) at 0.36 and 0.35, respectively. The highest positively loaded cranial variables PC 2 were the parietal and occipital sagittal chords (br-lam and lam-opn) at 0.39 and 0.26, correspondingly. Contrasting with these are negatively loaded facial dimensions, superior facial height and length (nas-pros and bas-pros) with -0.66 and -0.46. The largest positively loaded cranial variable for PC 3 was palatal breadth (bienm) at 0.80. Juxtaposed against these were negatively loaded breadth measurements, bizygomatic and bifrontomalartemporale (bizygo and bifmt). The scatterplot for PCs 1 & 2 (Figure 8.26) did not alter dramatically from Ln transformed data (Figure 8.24) despite placing habilis/rudolfensis within Australopithecus. In summary, PCA with habilis/rudolfensis within Australopithecus did not radically alter the results. Likewise, PCA still resulted in contrasting facial dimensions with neurocranial dimensions.

398 Table 8.28: PCA results for hominin genera Ln transformed generic medians for taxonomic scheme placing H. habilis/rudolfensis specimens in Australopithecus and all catarrhine genera.

Eigenvalue %Variance Cum. % PC1 0.24 90.00 90.0 PC2 0.01 5.28 95.28 PC3 0.005 1.97 97.25 PC4 0.003 1.06 98.31

PC1 PC2 PC3 g-o 0.28 0.17 -0.1255 biast 0.31 0.16 -0.1517 bipor 0.28 0.06 -0.1369 bizygo 0.27 -0.08 -0.3009 bifmt 0.25 0.11 -0.268 bas-pros 0.25 -0.46 -0.211 nas-pros 0.36 -0.66 0.08284 nas-br 0.22 0.25 -0.1638 br-lam 0.32 0.39 0.1534 lam-opn 0.35 0.26 0.1906 biecm 0.27 -0.07 -0.07076 bienm 0.29 -0.07 0.7981

Parietal & Occipital sagittal chords

Superior facial height & length Superior facial height & Occipital sagittal chord

Figure 8.26: Scatterplot of PC 1 & 2 of Ln transformed generic medians for taxonomic scheme placing H. habilis/rudolfensis specimens in Australopithecus and all catarrhine genera.

399 8.5.8 PCA results for hominins (Au.+hab, Para., Homo2 and H. sap) and other catarrhine genera, MSV: PCA based on the variance-covariance resulted in the first (Eigenvalue – 0.09, 52.81%) and second (Eigenvalue – 0.04, 23.4%) PCs accounting for 76.25% of the variation within the tribe (19.0% less variation explained than PCs 1 & 2 for Ln transformed data but 69.0% more variation explained than PCs 2 & 3 for Ln transformed data; Figure 8.27). The low to moderate variable loadings for all PCs were positive and negative (Table 8.29; fifteen measurements). The largest positively loaded cranial variables for PC 1 were bizygomaxillare superior breadth (bizs) and the maximum length of the zygomatic (zs-zgyi) at 0.63 and 0.58, respectively. Contrasting with these are negatively loaded dimensions such as superior facial height (nas-pros) and the maxillo-alveolar breadth (biecm) with -0.27 and - 0.25, correspondingly. The largest positively loaded cranial variables for PC 2 were breadth dimensions, bizygomatic and bifrontomalartemporale (bizygo and bifmt) at 0.49 and 0.39, respectively. Juxtaposed against these were negatively loaded measurements such as frontal sagittal chord (nas-br) and superior facial height (nas-pros) with -0.32 and -0.28, correspondingly. The scatterplot for PCs 1 & 2 (Figure 8.27) was similar to that for Ln transformed data (Figure 8.26). However, one difference was the placement of Australopithecus, which with the inclusion of habilis/rudolfensis, lies closer to the Homo samples (H.sap and Homo2).

400 Table 8.29: PCA results for hominin genera generic median MSVs for taxonomic scheme placing H. habilis/rudolfensis specimens in Australopithecus and all catarrhine genera and all catarrhine genera.

Eigenvalue %Variance Cum. % PC1 0.094449 52.805 52.81 PC2 0.041928 23.442 76.25 PC3 0.010901 6.0946 82.34 PC4 0.009695 5.4206 87.76

PC1 PC2 PC3 g-o -0.2502 0.08783 -0.4641 biast -0.1218 -0.03213 -0.1145 bipor -0.04818 0.1229 -0.04529 bizygo 0.09416 0.4938 -0.05923 bifmt -0.1222 0.3988 -0.04466 bizs 0.6341 0.3312 -0.03704 zs-zygi 0.5808 -0.402 -0.3919 bas-pros -0.2398 0.3152 -0.1464 nas-pros -0.2701 -0.2792 -0.1847 nas-br -0.1567 -0.3245 -0.1021 br-lam 0.03789 0.05267 0.02133 lam-opn 0.02331 -0.1293 0.7366 biecm -0.2502 0.08783 -0.4641 bienm -0.1218 -0.03213 -0.1145 ol-sta -0.04818 0.1229 -0.04529

Bizygomatic & bifmt breadths

Frontal sagittal chord & superior facial height Superior facial Bizygomaxillare superior height & biecm breadth & Zygomatic length

Figure 8.27: Scatterplot of PC 1 & 2 of generic median MSVs for taxonomic scheme placing H. habilis/rudolfensis specimens in Australopithecus and all catarrhine genera.

401 8.6 Morphological distances: 8.6.1: Summary of Euclidean distances produced from PCs 1-5: To further analyze the distances among hominin genera within a comparative catarrhine framework, PCA object scores were converted to Euclidean distances (i.e. distances of dissimilarity among genera). Three sets of data are considered in this section: in the first, mean distances values for the first five PCs are employed; the second uses only object scores for PC1; and the final uses only PC object scores for PC 2. Separate analyses are presented because in PCA not all principal components (PCs) are of equal weight (or value). That is, PC 1 usually accounts for the majority of the variance (often ~70%) in the data, with PC 2 accounting for the second largest amount of variance (often ~20%). The results for all four taxonomic scenarios explored in this chapter through PCA are summarized: PCA 1 = conventional taxonomic scheme Ln; PCA 2 = conventional taxonomic scheme MSV; PCA 3 = taxonomic scheme with H. habilis/rudolfensis placed in Australopithecus, Ln; and PCA 4 = taxonomic scheme with H. habilis/rudolfensis placed in Australopithecus, MSV. A summary of distance data for the first five PCs is provided in Table 8.30. For PCA 1, the mean distance for hominins is very similar to the grand mean for catarrhines (excluding hominins). As indicated by standard deviation (SD) values, dispersion in the hominin sample is smaller than the grand mean and equivalent to the colobines. For PCA 2, the mean distance among hominins is about the same as that for papionins and almost twice as large as the grand mean. However, dispersion is large but similar to that seen for the papionins and larger than the grand SD. The mean hominin distance for all five PCs for PCA 3 is below the grand mean, about the same as the mean as that for the Cercopithecini and similar to other catarrhines. Dispersion in the hominin sample is low, being similar to the Colobinae, and well below all other samples. Finally, the mean distance for hominins from PCA 4 is large, being well above the grand mean, but still below the mean values for papionins.

402 Table 8.31 presents the results of Kruskal-Wallis tests of the difference between median values for catarrhine distances involving five PCs. Non-parametric tests were used because they make no assumptions about the data under study. All four PCAs show significant median differences. Post hoc Mann-Whitney tests show: PCA 1: hominin median is not significantly different compared to all other catarrhines; only the Colobinae show significant median distances (compared to papionins and hominoids). PCA 2: the hominin median is significantly different to medians for the Colobinae and Cercopithecini; the papionin median is significantly different to the colobine and cercopithecine medians. PCA 3: the hominin median is not significantly different compared to all other catarrhines; papionin and hominoid median distances are significantly different to the Colobinae median. PCA 4: the hominin median is significantly different to medians for the Colobinae and Cercopithecinae; the papionin median distances is significantly different to the colobine and cercopithecine medians; the hominoid median distance is significantly different to the Colobinae median.

403 Table 8.30: Average and standard deviation of Euclidean distances from PCs1-5.

PCA 1 PCA 2 PCA 3 PCA 4 Hominini (N=6) 0.318±0.10 0.646±0.29 0.260±0.07 0.479±0.21 Colobinae (N=45) 0.251±0.10 0.302±0.12 0.210±0.09 0.280±0.101 Cercopithecini (N=10) 0.362±0.23 0.345±0.15 0.30±0.20 0.237±0.134 Papionini (N=15) 0.441±0.17 0.654±0.25 0.359±0.16 0.513±0.22 Hominoidea (N=6) 0.671±0.43 0.452±0.22 0.620±0.40 0.439±0.19 (exc. Homo)

Grand Mean±SD 0.308±0.16 0.389±0.22 0.284±0.19 0.315±0.18 (excl. Hominini)

Table 8.31: Kruskal-Wallis results for Euclidean distances with Mann-Whitney pairwise comparisons (p (same)).

Hominin Colob. Cerco. Papio. Hominoid 1. PCA 1 - 0.148 0.957 0.150 0.174 Hominin Hc: 19.14 - 0.202 0.000 0.010 Colob. p=0.000 - 0.360 0.143 Cerco. - 0.371 Papio. - Hominoid

2. PCA 2 - 0.010 0.034 0.969 0.379 Hominin Hc: 29.84 - 0.401 0.000 0.059 Colob. p<0.000 - 0.002 0.303 Cerco. - 0.150 Papio. Hominoid

3. PCA 3 - 0.132 0.786 0.111 0.128 Hominin Hc: 17.6 - 0.234 0.000 0.004 Colob. p=0.001 - 0.524 0.143 Cerco. - 0.259 Papio. - Hominoid

4. PCA 4 - 0.003 0.034 0.669 0.933 Hominin Hc: 26.89 - 0.697 0.000 0.015 Colob. p<0.000 - 0.004 0.058 Cerco. - 0.559 Papio. - Hominoid

404 8.6.2: Summary of Euclidean distances produced from PC 1: A summary of distance data for the first PC is provided in Table 8.32. The coefficient of variation (CV) values are also provided as these analyses are likely to be more meaningful then those above (using all five PCs). For PCA 1, the mean distance for hominins is small (the smallest) and well below the grand mean. Moreover, the SD and CV values are well below all other samples and well below the grand SD and CV. For PCA 2, the mean distance among hominins is inflated compared the grand mean, being similar to the mean for the papionini. The SD value is virtually indistinguishable from the papionin value. However, the CV for the hominin sample is similar to most sample values, but below the grand CV. The mean hominin distance for PCA 3 is well below the grand mean, and well below mean values for all other catarrhines. SD and CV values are similarly very low. Finally, the mean distance for hominins from PCA 4 is large, being well above the grand mean, but still below the mean values for papionins. The SD value for the hominins ample is identical to the grand SD; the CV value is low compared to the grand CV and all samples. Table 8.33 presents the results of Kruskal-Wallis tests of the difference between median values for catarrhine distances from PC 1. All four PCAs show significant median differences. Post hoc Mann-Whitney tests indicate: PCA 1: the hominin median is significantly different to medians for the papionins and hominoids; papionins and hominoids exhibit medians which are significantly different to the median for the Colobinae. PCA 2: the hominin median is significantly different to the colobine median; the papionin median is significantly different to the colobine median. PCA 3: the hominin median is significantly different to the hominoid median distance; papionin and hominoid median distances are significantly different to the colobine distance. PCA 4: the hominin median is significantly different to medians for the Colobinae and Cercopithecini; the papionin median distances are significantly different to the colobine median.

405 Table 8.32: Average, standard deviation and CV values (parenthesis) of Euclidean distances from PC1.

PCA 1 PCA 2 PCA 3 PCA 4 Hominini (N=6) 0.129±0.07 0.496±0.32 0.133±0.07 0.381±0.17 (50.6) (63.8) (56.0) (44.7) Colobinae (N=45) 0.187±0.12 0.214±0.14 0.161±0.10 0.182±0.12 (66.4) (64.1) (63.5) (65.4) Cercopithecini (N=10) 0.309±0.25 0.246±0.19 0.275±0.21 0.174±0.16 (79.6) (77.9) (74.7) (89.1) Papionini (N=15) 0.311±0.18 0.452±0.31 0.267±0.16 0.356±0.24 (58.2) (67.9) (59.6) (68.0) Hominoidea (N=6) 0.654±0.43 0.292±0.19 0.600±0.40 0.259±0.17 (exc. Homo) (66.5) (65.6) (67.3) (64.2)

Grand Mean±SD 0.264±0.23 0.271±0.21 0.232±0.20 0.221±0.17 (excl. Hominini) (85.9) (77.5) (87.5) (76.8)

Table 8.33: Kruskal-Wallis results for Euclidean distances of PC1 with Mann-Whitney pairwise comparisons (p (same)).

Hominin Colob. Cerco. Papio. Hominoid 1. PCA 1 - 0.357 0.303 0.018 0.008 Hominin Hc: 14.3 - 0.234 0.022 0.006 Colob. p=0.006 - 0.890 0.143 Cerco. - 0.080 Papio. - Hominoid

2. PCA 2 - 0.042 0.143 0.559 0.230 Hominin Hc: 11.24 - 0.686 0.005 0.327 Colob. P=0.024 - 0.071 0.551 Cerco. - 0.331 Papio. - Hominoid

3. PCA 3 - 0.569 0.212 0.080 0.013 Hominin Hc: 14.79 - 0.100 0.021 0.004 Colob. p=0.005 - 0.988 0.116 Cerco. - 0.094 Papio. - Hominoid

4. PCA 4 - 0.003 0.037 0.821 0.273 Hominin Hc: 14.2 - 0.655 0.009 0.214 Colob. P=0.007 - 0.063 0.481 Cerco. - 0.414 Papio. - Hominoid

406 8.6.3: Summary of Euclidean distances produced from PC 2: A summary of distance data for the second PC is provided in Table 8.34. For PCA 1, the mean distance for hominins is large, being more than double the grand mean. Moreover, the SD value is also inflated; however, the hominin CV is similar to the grand CV. For PCA 2, the mean distance among hominins is highly inflated compared the grand mean, and well above all catarrhine means. The SD value is large but similar to that for the Hominoidea. The CV value is low compared to all samples, and about half of the grand CV. The mean hominin distance for PCA 3 is also inflated, being more than double the grand mean. The hominin SD is only slightly larger than other catarrhines. The hominin CV is, however, reduced compared to all samples. Lastly, the mean distance for hominins from PCA 4 is large, being almost three times the size distance of the grand mean. The SD value for the hominins ample is large, but below the value for the Hominoidea. The hominin CV is low compared to all samples. Table 8.35 presents the results of Kruskal-Wallis tests of the difference between median values for catarrhine distances from PC 1. All four PCAs show significant median differences. Post hoc Mann-Whitney tests indicate: PCA 1: the hominin median is not significantly different to any median; papionins exhibit a median which is significantly different to the median for the Colobinae. PCA 2: the hominin median is significantly different to all medians except for the hominoids; the papionin median is significantly different to the colobine median. PCA 3: the hominin median is significantly different only to the Colobinae; no other medians are significantly different. PCA 4: the hominin median is significantly different to medians for the Colobinae and Cercopithecini; the papionin median distance is significantly different to the colobine median.

407 Table 8.34: Average, standard deviation and CV values (parenthesis) of Euclidean distances from PC2.

PCA 1 PCA 2 PCA 3 PCA 4 Hominini (N=6) 0.220±0.15 0.286±0.15 0.177±0.09 0.190±0.11 (66.1) (53.3) (49.8) (57.7) Colobinae (N=45) 0.094±0.06 0.036±0.03 0.80±0.05 0.047±0.05 (64.0) (86.8) (66.6) (95.9) Cercopithecini (N=10) 0.111±0.08 0.057±0.04 0.078±0.07 0.061±0.04 (68.8) (72.4) (86.4) (65.5) Papionini (N=15) 0.153±0.09 0.090±0.06 0.108±0.07 0.126±0.08 (59.0) (62.2) (65.0) (65.2) Hominoidea (N=6) 0.075±0.05 0.136±0.12 0.089±0.06 0.156±0.14 (exc. Homo) (72.7) (88.5) (66.1) (89.8)

Grand Mean±SD 0.107±0.07 0.057±0.06 0.086±0.06 0.073±0.07 (excl. Hominini) (67.4) (100.5) (68.6) (101.5)

Table 8.35: Kruskal-Wallis results for Euclidean distances of PC2 with Mann-Whitney pairwise comparisons (p (same)).

Hominin Colob. Cerco. Papio Hominoid 1. PCA 1 - 0.068 0.175 0.293 0.174 Hominin Hc: 8.97 - 0.534 0.021 0.456 Colob. p=0.062 - 0.279 0.481 Cerco. - 0.067 Papio. - Hominoid

2. PCA 2 - 0.000 0.006 0.004 0.128 Hominin Hc: 24.7 - 0.252 0.000 0.063 Colob. p<0.000 - 0.488 0.357 Cerco. - 0.726 Papio. - Hominoid

3. PCA 3 - 0.009 0.058 0.094 0.093 Hominin Hc: 8.34 - 0.608 0.167 0.672 Colob p=0.08 - 0.360 0.786 Cerco - 0.723 Papio - Hominoid

4. PCA 4 - 0.001 0.020 0.331 0.936 Hominin Hc: 19.49 - 0.129 0.002 0.077 Colob. P<0.000 - 0.056 0.623 Cerco. - 0.559 Papio. - Hominoid

408 8.6.4 Euclidean distances summary: To summarize the above findings, hominin size variation, as revealed by distances from PCAs employing Ln-transformed data, is both reduced and far less variable than all other catarrhines. That is, the mean distances for hominins are well below many catarrhines and often below the grand mean in these analyses. In contrast, mean distances from MSV data are much larger in hominins compared with living catarrhines. From this it is concluded that the difference in size among hominin genera is much smaller than for extant catarrhines, contrasting with differences in cranial shape, which are equivalent to the largest or larger than those between genera of extant catarrhines. This pattern holds for comparison involving PCs1-5 and those only involving PC1 or PC2. In terms of the variability of distances, as indicated with CVs for distances from PC1 and PC2 object scores, hominins are characterized by substantially lower relative variation than extant catarrhines. This finding is striking as in almost all instances the hominin CV is below most catarrhine CVs and the grand CV. In some instances, CV values for catarrhines are more than twice as large as those for hominins. Crucially, analyses which tested different hominin taxonomic scenarios – those involving the removal of H. habilis/rudolfensis from Homo and its placement to Australopithecus – show no substantial differences. In other words, the removal of H. habilis/rudolfensis from Australopithecus appears to have little affected on the median difference among hominins nor variability within this tribe.

8.7 Reconstructed paleodiets and paleohabitats of fossil hominins: A complete review of the evidence pertaining to the reconstructed paleodiets and paleohabitats for the fossil hominins is beyond the scope of this thesis; however, some brief comments are possible. First, the earliest hominins have been recovered from deposits indicating wooded forests, not open savanna. Only much later hominins are recovered from deposits indicative of open, dry environments. Second, tooth microwear, dental eruption sequences and isotopic analyses have demonstrated clear differences between Australopithecus, Paranthropus and early Homo; presently, the diet of Ardipithecus, Orrorin and Sahelanthropus are unknown because no studies have yet been published. (See Grine, 1981 & 1986; Maier, 1984; Martin, 1984; Lucas et al, 1985; Smith, 1986,

409 1991, 1994a and 1994b; Cela-Conde, 1996; Sponheimer & Lee-Thorp, 1999; Wood & Richmond, 2000; Teaford & Ungar, 2000; Lee-Thorp & Sillen, 2001; Lee-Thorp et al, 2003; Ungar, 2004; Wood & Strait, 2004; Singleton, 2004; Pickering, 2006; Grine et al, 2006; Ungar et al, 2006; Sponheimer et al, 2006; Lee-Thorp & Sponheimer, 2006.)

8.8 Locomotion and Limb Proportions of fossil hominins: Again, complete review of the evidence pertaining to the locomotion, limb proportions and postcranial anatomy for the fossil hominins is beyond the scope of this thesis; however, some brief comments are possible. First, it is fair to say that although hominins of Australopithecus and Paranthropus were bipedal, although it was different from that of modern humans. Second, the picture emerging of fore- and hindlimb evolution for hominin is relatively complex (with the possibility of some genera and/or species experiencing evolutionary reversals) and it is difficult to make any broad generalizations. This is due the few postcranial fossils discovered and the fragmentary nature of most postcranial remains. Additionally, if postcranial remains are not found in direct association with craniodental fossil remains, and more than one hominin species is known from the fossil bearing strata, it is very difficult to ascertain who they belong to. Finally, McHenry et al (2007) have presented evidence from fossil ulnae of presumably Paranthropus species but there is marked variation in the sample perhaps suggesting paraphyly. (See Day, 1969 & 1971; Robinson, 1972; Oxnard, 1975a & 1975b; McHenry, 1978 & 1994; Fleagle et al, 1981; Aiello & Dean, 1982; Deloison, 1985; Jungers, 1988a; Berger & Tobias, 1996; Richmond & Strait, 2000; Wood & Richmond, 2000; Richmond et al, 2001 & 2002; Ward, 2002; Haeusler & McHenry, 2004 & 2007; Ohman et al, 2005; Reno et al, 2005; White, 2006; Carrier, 2007; Green et al, 2007.)

8.9 Inferences from genetic data of extant catarrhines: The use of comparative biomolecular and genetic data has a long history in primatology and anthropology in general. As discussed in Chapter 2, the work of Goodman (1963a & 1963b), his students and colleagues which began in the early 1960s spearheaded and continues to break new ground in primate comparative genetic research (e.g. Uddin et al, 2004). The most obvious genetic difference between humans and the Great apes is the

410 diploid number of these species. Humans have 46 while the Great apes have 48. The estimated time for this chromosomal fusion and rearrangement is posited to have occurred at the initial divergence of chimpanzees and humans (Yunis & Prakash, 1982; Mai, 1983; Chiarelli, 1985; Navarro & Barton, 2003; Ayala & Coluzzi, 2005) or perhaps as little as 400 Ka (Arnason et al, 2000). However, despite differences in chromosome number, the African Apes, including humans, are chromosomally and genetically very similar (Yunis & Prakash, 1982; Eastel et al, 1995; Jobling et al, 2004). However, unlike modern human populations which are globally distributed yet genetically homogenous across populations, the genomes of chimpanzees and gorillas are much more genetically diverse (Morin et al, 1994; Gagneux et al, 1999 & 2001; Kaessmann et al, 1999; Stone et al, 2002). Despite this, hominoids exhibit a slowing down of neutral genetic evolution (Koop et al, 1986; Bailey et al, 1991; Elango et al, 2006). Eckhardt (2000; p.157, figure 7.1) is quite perceptive is his analysis and discussion of the very small genetic distance, less than 2%, between modern humans and chimpanzees (e.g. King & Wilson, 1975; Bruce & Ayala, 1978; Andrews, 1986; Goodman et al, 1989 & 1990; Page & Goodman, 2001; Wildman et al, 2003; cf. Britten, 2002 and Glazko et al, 2005). As such, intuitively all hominin genera and species must than be accommodated within the approximately one percent genetic difference between humans and chimpanzees. This results in hominin species since the human-chimp divergence on average producing genetic distances between species ~ 0.07 (e.g. 1/15 = 0.0666). In their mtDNA analyses of Simias concolor and its closest colobine relative, Whittaker et al (2006) state, “…individuals differing less than 10% are generally considered congenerics…” (p. 890, table 2). If this criterion was adopted for the hominins, no more than two or three genera would probably be necessary (for similar argument see, Watson et al, 2001; for criticism, see Cameron, 2003). Perhaps the most useful caveat of primate genetic comparisons is providing temporal estimates as well as constraints for the cladogenetic event leading to the human and chimpanzee lineages. Although estimates vary, most genetic studies suggest sometime between five and ten million years ago the last common ancestor of humans and chimpanzees diverged (5 Ma - Sarich & Wilson, 1967; 6-8 Ma - Caccone & Powell, 1989; 4.9±0.2 Ma - Horai et al, 1995; 10.5-13 Ma - Arnason et al, 2000; ~5-6 - Ma Page &

411 Goodman, 2001; 5.4±0.6 Ma - Stauffer et al, 2001; 8.1 Ma - Raaum et al, 2005; 4.98-7.02 Ma - Kumar et al, 2005; <6.3 Ma - Patterson et al, 2006). These differences are due to different region(s) examined (i.e. nDNA or mtDNA; coding or noncoding) and inherent theoretical assumptions, such as, calibration points and sequence substitution and/or mutation rate (Britten, 2002; Kumar, 2005). Additionally, Stedman et al (2004) reported that inactivation (caused by a frameshifting mutation) of the human sacromeric myosin gene, MYH16, was involved with the reduction of large mastication muscles as well an aiding the increase in cranial capacity. More importantly, by using the coding sequence of this nuclear DNA region, they were able to construct a molecular clock to estimate when this mutation occurred. Their results suggest this frameshifting mutation occurred ~2.4 Ma, which is very close in time to the first appearance of fossils generally attributed to Homo. However, these results have been challenged by McCollum et al (2006).

412 Chapter 9: Conclusions “From the purely biological point of view man is certainly at least as different as a very good genus” (Mayr, 1950; p. 111).

9.1 Introduction: The genus and species categories, which initiated binomial nomenclature, are perhaps the most important contributions Linnaeus made to biological classification. This thesis examined the genus category and cranial morphometrics of the extant Catarrhini and applied these results as comparative benchmarks to some fossil hominin genera. From the forgoing generic analyses and comparisons many sound inferences about the genus category and fossil hominin genus taxonomy are possible. The purpose of this chapter is to summarize and discuss the results of this thesis and its implication to the fossil record and biological classification of human evolution. This topic is important because the taxonomic and systematic relationships of humans and their closest living relative, extant or extinct, are vital to understanding and discussing human evolution and classification. Very little attention and analysis have been given to the genus category. One of the main reasons taxonomic ranks above the species level do not gain attention like species concept is because most think that only species can be the unit of study in evolution (Ereshefsky, 1991, 1997 & 2001). Additionally, many feel taxonomy above the species level is arbitrary. However, it is the author’s opinion based on the available evidence, that the genus category can classify biological phenomenon in a very meaningful and coherent fashion. The main research question posed at the beginning of this thesis was; Are the recently proposed hominin genera, such as Sahelanthropus, Orrorin, Ardipithecus and Kenyanthropus, adequately defined and necessary when the extant Catarrhini genera are used as a comparative benchmark? As well as, questions 7-9 in chapter 2. Question 7: Are the recently proposed fossil hominin genera adaptively coherent occupying an adaptive zone different from that of other hominin genera based on comparisons with extant Catarrhini genera?; Question 8: Is the congeneric placement of humans and chimpanzees (Goodman et al, 1998 & 2001; Wildman et al, 2003) more parsimonious than each occupying separate genera?; and Question 9: Would the congeneric placement of humans and chimpanzees better explain and synthesize the available evidence or oversimplify evolutionary differences? Or put more simply, what types of evidence best delineate generic boundaries?

The approach adopted here was grounded upon a comparative framework of extant catarrhines, as well as taxonomic uniformitarianism and equivalence. Put differently, the evidence and justification for any biological or generic classification should be reasonably similar for most taxa and no taxon should be accorded with unique classifications or evolutionary scenarios, including that of humans. This analysis was original by using all the extant catarrhines as a comparative benchmark to interpret the generic taxonomy for fossil hominin genera. The catarrhines are the most appropriate comparative analogs because hominins are also catarrhines.

9.2 Summary of many findings: 9.2.1 Catarrhines: In general, the cranium and its various dimensions proved very useful in sorting genera. Overall, body size seems to be the most important distinguishing feature among catarrhine genera and appears to reflect best differences in their adaptive zones. Conversely, cranial shape differences play a relatively minor role in distinguishing them. Additionally, many cranial dimensions were significantly correlated to body size. This was made clear by the large number of cranial variables exhibiting statistically significant correlations with body size (mass) during regression analyses. The average number of species per genus for Catarrhines is actually rather low, usually less than ten. Measurements which sorted genera and generated significant results included, Facial lengths and breadths (nas-pros, bas-pros, bifmt, bizygo, etc.); Length and breadth of palate (ol-sta, bienm, biecm); Neurocranial lengths (g-o, nas-br, br-lam and lam-opn); and

414 Neurocranial breadths (biast and bipor). Likewise, cranial indices involving these measurements also sorted genera and generated significant results. Furthermore, limb proportions, particularly the intermembral index, can reveal stark differences between closely related genera and indicate locomotor behaviors.

9.2.2 Hominins: Despite small sample sizes, some clear differences emerged between hominin genera. Measurements which sorted genera and generated significant results included, Cranial vault length (g-o); Biasterionic Breadth (biast); Bizygomatic breadth (bizygo); Superior facial height (nas-pros); and The frontal, parietal and occipital sagittal chords (nas-br, br-lam and lam-opn). Additionally, the first two cranial indices each produced significant results (bas-pros/biast x 100 and bas-pros/bipor x 100).

9.2.3 Hominins compared to Catarrhines: When the hominins are compared to the extant catarrhines some interesting observations and analogies are possible. First, based on extant catarrhine systematics and species numbers there is no reason to believe a priori that the hominins were particularly speciose. Second, the absolute size of hominin crania also provides some indication of terrestriality, irrespective of the positioning for the foramen magnum. Third, there can be large phenotypic dissimilarities between closely related genera or species (e.g. Nasalis and Simias or Cercocebus and Mandrillus). Fourth, the most important contrasts in cranial form between catarrhine genera are between the neurocranium and viscerocranium: overall, a pattern is seen where a large viscerocranium is combined with a small neurocranium, and vice versa. Fifth, when considering the genetic distances of extant genera and species, the genetic distances between fossil hominins and modern humans was probably very small.

415 Sixth, the cranium of modern humans is highly distinct and derived, and has witnessed dramatic changes (in size and shape) not seen in any other primate. Seventh, when treated with multivariate statistics, the cranial dimensions of fossil hominins are generally more similar morphometrically to the Great apes than to modern humans. Eighth, beware of convergence (homoplasy; e.g. Papio and Mandrillus). Ninth, body size (i.e. mass) is an important determinant of an organism’s adaptive zone and provides some indication of preferred habitat. This study has found that size differences between hominin genera are low compared to most catarrhines. This find is stark and forms the basis of an important difference between members of the tribe Hominini and other catarrhines. Moreover, variability in size within hominin genera is very low compared to extant catarrhines. Tenth, in contrast, shape differences between hominin genera are much greater than seen between many extant catarrhines. However, variability in shape within hominin genera is low. Eleventh, as with other catarrhines (see point 4), the most important contrasts in cranial form between hominin genera are between the neurocranium and viscerocranium: overall, a pattern is seen where a large viscerocranium is combined with a small neurocranium (Australopithecus and Paranthropus), the obverse applying in Homo (small viscerocranium and large neurocranium). Twelfth, hominins appear to have occupied very similar habitats, in terms of both ecological differences between and within genera; as indicated by small differences between and low variability within genera in body size Thirteenth, niche separation among hominin genera was likely to have been subtle; it may be undetectable with contemporary techniques employed by paleoanthropologist. Fourteenth, the large shape differences seen among hominin genera results almost exclusively from the dramatic changes in cranial morphology associated with encephalization in Homo Fifteenth, at a minimum, the major morphological differences between Homo and other hominin genera warrant its classification in at least a separate genus; a case

416 could be made for its classification in a higher rank (?subtribe) compared to all other hominins Sixteenth, Australopithecus and Paranthropus are about as dissimilar in their cranial morphology as Pan, Gorilla and Pongo are from each other. This is remarkable given the short divergence time between these hominins (LCA ~2.5-3.0 Ma) and the long divergence times among these living apes (~8.0-15.0 Ma). And lastly, seventeenth, within a broad catarrhine framework, Australopithecus and Paranthropus are good genera and should be retained distinct form each other and from Homo

9.2.4 Wood and Collard’s genus argument: The use of the adaptive zone as a concept for the classification of hominins especially Homo as proposed by Wood & Collard (1999) has merit. Cranial (body) size seems to be the greatest indicator of differences in adaptive zone within the catarrhines but lees so in hominins. This suggests that linking adaptive zones to body size in hominins may be unwise. Moreover, as shape differences among catarrhine genera are generally of little utility in distinguishing them, shape is unlikely to be an important indicator of adaptive zone distinctions among most hominins: the exception is Homo which has a substantially modified cranial shape due to increased encephalization. On this basis, generic distinctions among the best known hominins (Australopithecus, Paranthropus and Homo) receive support. However, the present study neither supports nor refutes their proposal to reclassify H. habilis/rudolfensis in Australopithecus: analyses testing this and the more conventional taxonomy showed no substantial difference in size and shape differences or variability. Importantly, if cranial size and shape changes resulting from encephalization are taken as indicating a major adaptive shift (in zone) with the emergence of Homo, then it may be premature to exclude H. habilis/rudolfensis form this genus.

417 9.4 Implications for paleoanthropology: Paleoanthropology has been overly burdened with hominin taxonomy and differences in opinion (e.g. evolutionary systematics or cladistics) as to how and why hominins are classified (Andrews & Harrison, 2005). Marks (2005) stated that hominin taxonomy is in “worst shape” (p. 49) now as it was in the earlier part of the 20th century. Indeed, when examining the paleoanthropological literature one can find a variety of differing cladograms, taxonomic names (generic and specific) and evolutionary scenarios. This thesis attempted to answer questions such as, Should afarensis be removed from Australopithecus and instead placed in Praeanthropus (Strait et al, 1997)? Is Paranthropus a monophyletic genus (Wood, 1988; McCollum, 1999; McHenry et al, 2007) or should these robust hominins be included within Australopithecus (Tobias, 1967; Suwa et al, 1997; Alemseged et al, 2002)? Should humans and chimpanzees be placed in the same genus (Goodman et al, 1998)? The data presented herein can indirectly provide some answers.

9.4.1 Question 7: After reviewing and comparing the available evidence of the early hominin genera, as well as evaluating them from an extant catarrhine perspective, in the author’s opinion and based on the available evidence, it is difficult to argue that they all occupied an adaptive zone different from that of other hominin genera on the available evidence. First, based on published body weight estimates, all the early hominin genera are very similar; Orrorin 35-40 kg (Nakatsukasa et al, 2007); Ardipithecus 40 kg (Collard & Wood, 1999) (see also Haile-Selassie, 2001; Haile-Selassie et al, 2004); and by using the linear equations derived the bivariate analyses of the mean for hominoid cranial variables versus generic mean body weight, Sahelanthropus likely weighed ~ 40-50 kg (Brunet et al, 2002; Bipor = 102mm (est., p.149, table 4), y = 0.251x + 1.9484, with estimated body weight of ~43 kg; additionally, Bifmt = 102mm (est., p.150, table 5), y = 0.2402x + 2.0177, with estimated body weight of ~52 kg), which is very similar to chimpanzees. Secondly, when compared to the genus systematics of the extant catarrhines, the number of genera within the tribe Hominini is perhaps ‘overcrowded’, so to speak. Furthermore, when consideration is given to the time-scale of human evolution (~5-<10

418 Myr), as well as the number of genera used for the extant catarrhine subgroups, again, the number of genera is large. However, in addition, no paleoanthropologist would like their discovery to be named by another person or group; and naming new a genus or species will probably also provide funding and fame (Smith, 2005). Lastly, the use of Praeanthropus for the afarensis hypodigm should perhaps be used with caution because of recent findings and analyses. On the one hand, there is a link from anamensis to afarensis (Kimbel et al, 2006; White et al, 2006); and on the other, there is a link from afarensis to the robust hominins of Paranthropus (Rak et al, 2007). Despite this, Australopithecus will remain paraphyletic because most agree the genus Homo emerged from a species from the former and yet are not allocated to the same genus.

9.4.2 Question 8: The congeneric placement of humans and chimpanzees, in the author’s opinion and based on the available evidence, is not the most parsimonious generic arrangement for these two extant apes. The arguments put forward by Goodman et al (1998 & 2001) and Wildman et al (2003) are completely valid, congruent with cladistic theory and associated with robust statistical support but when we take into consideration the present diversity of apes and the lack of ancient hominin DNA, could they really produce any other conclusion? Furthermore, the genetic analyses of Wildman et al (2003) are comprised of +90 thousand sequenced nucleotides comprising 97 genes, which is impressive but still remains only a small fraction of the entire genomes for the primate species examined. For example, if there are roughly 35,000 genes in the human genome (Ewing & Green, 2000), the analyses by Wildman et al (2003) really only samples ~0.28% of human genes (e.g. 97/35000=0.002771; 0.002771*100=0.27771). Furthermore, the claim by Wildman et al (2003) that, “DNA evidence provides an objective non-anthropocentric view of the place of humans in evolution” (p. 7181) is difficult to substantiate considering human DNA is an integral part of the analyses. Additionally, recent genetic analyses have demonstrated many interesting differences between humans and chimpanzees (Enard et al, 2002; Britten, 2002; Clark et al, 2003; Glazko et al, 2005; Wooding et al, 2006; Bakewell et al, 2007). Lastly, if chromosome number is used sort genus and/or species-groups for Hylobates or Cercopithecus, the same should be accorded to humans (2n=46) and

419 chimpanzees (2n=48). Furthermore, many comparative genetic analyses use non-coding nucleotide sequences and neutral mutations to gauge and estimate divergence time (although, the analyses of Clark et al (2003) and Wildman et al (2003) did measure the ratio between synonymous and nonsynonymous substitutions). However, these regions have not been affected by natural selection. Why should neutral genetic evolution over time be more important than features (e.g. genetic, morphological or behavioral) which have experienced positive selection (e.g. Ackermann & Cheverud, 2004a; Bakewell et al, 2007)? 9.4.3 Question 9: In light of the evidence presented here and recent genetic analyses, the congeneric placement of humans and chimpanzees would, in the author’s opinion and based on the available evidence, oversimplify evolutionary differences between these two apes. Placing humans and chimpanzees in the same genus generates the impression that they are very similar and not having changed drastically since last sharing a common ancestor, which we know is not the case. Of course numerous field and laboratory studies of chimpanzees have demonstrated the cultural and behavioral complexity of these species but it must be admitted that their respective ‘adaptive zone’ (e.g. from morphology to diet to ontogeny and locomotion) is very different to that of modern humans. Additionally, one aspect from the writings of Simpson (1943, 1945, 1959, 1961 & 1963) and Mayr (1950, 1963, 1982; Mayr et al, 1953; Mayr & Bock, 2002) is that one purpose of taxonomy is to aid communication and discussion of biological phenomenon and classification between researchers. If chimpanzees and humans are placed in the same genus, there will be some confusion and ambiguity. Or perhaps worded better, “If classification is to serve primarily for communication and identification, utility is the principal criterion for choosing one system over another” (Raup & Stanley, 1978, p. 130). 9.5 Avenues for future research: As the main focus of this study were the catarrhines, applying these similar comparisons and statistical methods to other primate groupings, such as the Platyrrhini or Strepsirhini, as well as other mammalian orders, would be useful. Additionally, more measurements could also improve results and separate genera. It would also be worthwhile and interesting to examine if there are any significant correlations between morphological and genetic distance matrices.

420 References:

Abegg, C. & Thierry, B. 2002. Macaque evolution and dispersal in insular south-east Asia. Biol. J. of the Linn. Soc. 75, 555-576.

Ackermann, R.R. 2002. Patterns of covariation in the hominoid craniofacial skeleton: implications for paleoanthropological models. JHE 42, 167-187.

Ackermann, R.R. 2003. Using extant morphological variation to understand fossil relationships: A cautionary tale. SAJS 99, 255-258.

Ackermann, R.R. 2005. Ontogenetic integration of the hominoid face. JHE 48, 175-197.

Ackermann, R.R. & Krovitz, G.E. 2002. Common patterns of facial ontogeny in the hominid lineage. The Anatomical Record (New Anat.) 260, 142-147.

Ackermann, R.R. & Cheverud, J.M. 2004a. Detecting genetic drift versus selection in human evolution. PNAS 101, 17946-17951.

Ackermann, R.R. & Cheverud, J.M. 2004b. Morphological integration in primate evolution. In (Pigliucci, M. & Preston, K., Eds.) Phenotypic Integration: studying the ecology and evolution of complex phenotypes, pp. 302-319. New York: Oxford Uni. Press.

Ackermann, R.R., Roger, J. & Cheverud, J.M. 2006. Identifying the morphological signatures of hybridization in primate and human evolution. JHE 51, 632-645.

Ahern, J.C.M. 1998. Understanding intraspecific variation: The problem with excluding Sts 19 from Australopithecus africanus. AJPA 105, 461-480.

Ahern, J.C.M. 2005. Foramen magnum position in Pan troglodytes, Plio-Pleistocene Hominids, and recent Homo sapiens: Implications for recognizing the earliest hominids. AJPA 127, 267-276.

Aiello, L. 1981a. Locomotion in the Miocene Hominoidea. In, (Stringer, C. Ed.) Aspects of Human Evolution. Symp. Soc. For the study of human biology Vol. XXI, pp. 63-97. London: Taylor & Francis Ltd.

Aiello, L. 1981b. The allometry of primate body proportions. Symp. Zool. Soc. Lond. 48, 331-358.

Aiello, L. & Dean, M. 1982. The evolution of the locomotion in the early hominidae. In, Progress in Anatomy: Anatomical Society of Great Britain and Ireland, pp. 81-97. UK: Cambridge University Press.

Aiello, L. & Dean, C. 1990. An Introduction to Human Evolutionary Anatomy. London: Academic Press.

Aiello, L. & Dunbar, R.I.M. 1993. Neocortex size, group size and the evolution of language. Current Anthropology 34(2), 184-193.

Aiello, L. & Wood, B. 1994. Cranial variables as predictors of hominine body mass. AJPA 95, 409-426.

Aiello, L. & Wheeler, P. 1995. The expressive-tissue hypothesis: The brain and the digestive system in human and primate evolution. Current Anthropology 36(2), 199-221.

Aiello, L.C. & Wells, J.C.K. 2000. Energetics and the evolution of the genus Homo. Annu. Rev. Anthropol. 31, 323-338.

Aiello, L.C. & Andrews, P. 2000. The Australopithecines in review. Human Evolution 15, 17-38.

421 Aiello, L.C., Collard, M., Thackeray, J.F. & Wood, B.A. 2000. Assessing exact randomization-based methods for determining the taxonomic significance of variability in the human fossil record. SAJS 96, 179- 183.

Aiello, L.C. & Collard, M.C. 2001. Our newest oldest ancestor. Nature 410, 526-527.

Aimi, M. 1981. Fossil Macaca nemestrina (Linnaeus, 1766) from Java, Indonesia. Primates 22, 409-413.

Aimi, M., & Nogami, Y. 1989. Macaca fuscata develops thin enamel on lingual sides of lower incisors. Primates 30, 261-264.

Aimi, M. & Bakar, A. 1992. Taxonomy and distribution of Presbytis melalophos group in Sumatra, Indonesia. Primates 33(2), 191-206.

Albrecht, G.H. 1978. The craniofacial morphology of the Sulawesi macaques: Multivariate approaches to biological problems. Contributions to Primatology 13, 1-123.

Albrecht, G.H. 1980. Multivariate analysis and the study of form, with special reference to canonical variates analysis. Amer. Zool. 20, 679-693.

Albrecht, G.H. 1992. Assessing the affinities of fossils using canonical variates and generalized distances. Human Evolution 7(4), 49-69.

Albrecht, G.H. & Miller, J.M.A. 1993. Geographic variation in primates: a review with implications for interpreting fossils. In, (W.H. Kimbel & B.M. Lawrence, Eds.) Species, Species Concepts, and Primate Evolution, pp. 123-161. New York: Plenum Press.

Albrecht, G.H., Gelvin, B.R. & Hartman, S.E. 1993. Ratios as a size adjustment in morphometrics. AJPA 91, 441-468.

Albrecht, G.H., Gelvin, B.R. & Miller, J.M.A. 2003. The hierarchy of intraspecific craniometric variation in gorillas: A population-thinking approach with implications for fossil species recognition studies. In, (Taylor, A. & Goldsmith, M.L., Eds.) Gorilla biology: a multidisciplinary perspective, pp. 62-103. UK: Cambridge Uni. Press.

Alemseged, Z., Coppens, Y. & Geraads, D. 2002. Hominid cranium from Omo: Description and taxonomy of Omo-323-1976-896. AJPA 117, 103-112.

Alemseged, Z., Spoor, F., Kimbel, W., Bobe, R., Geraads, D., Reed, D. & Wynn, J.G. 2006. A juvenile early hominin skeleton from Dikika, Ethiopia. Nature 443, 296-301.

Alexander, Jr., R.W. 1981. Sexual dimorphism in the cranium Cercopithecus aethiops. JHE 10, 311-317.

Alexander, Jr., R.W. 1982. Sexual dimorphism in the cranium of Cercocebus albigena. JHE 10, 143-150.

Alpagut, B., Andrews, P., Fortelius, M., Kappleman, J., Temizsoy, Celebi, H. & Lindsay, W. 1996. A new specimen of Ankarapithecus meteai from the Sinap formation of central Anatolia. Nature 382, 349-351.

Anapol, F. & Gray, J.P. 2003. Fiber architecture of the intrinsic muscles of the shoulder and arm in semiterrestrial and arboreal guenons. AJPA 122, 51-65.

Anapol, F., Shahnoor, N. & Gray, J.P. 2004. Fiber architecture, muscle function, and behavior: gluteal and hamstring muscles of semiterrestrial and arboreal guenons. In, (Anapol, F., German, R.Z. & Jablonski, N.G., Eds) Shaping primate evolution: form, function and behavior, pp. 99-133. Cambridge: Cambridge University Press.

422 Anapol, F., Turner, T.R., Mott, C.S. & Jolly, C.J. 2005. Comparative postcranial body shape and locomotion in Chlorocebus aethiops and Cercopithecus mitis. AJPA 127, 231-239.

Anderson, S. 1975. On the number of categories in biological classification. American Museum Novitates 2584, 1-9.

Anderson, S. & Jones, J.K., Eds. 1967. Recent mammals of the world: a synopsis of families. New York: Ronald Press Company.

Andrews, P. 1981. Species diversity and diet in the monkeys and apes during the Miocene. In, (Stringer, C. Ed.) Aspects of Human Evolution, pp. 25-61. Symp. Soc. For the study of human biology Vol. XXI. London: Taylor & Francis Ltd.

Andrews, P. 1981 & 1985. The origin of the Cercopithecoidea. In (Napier, P.H.) Catalogue of Primates in the British Museum (Natural History) & elsewhere in the British Isles, Parts I & II: Family Cercopithecidae, Subfamily Cercopithecinae & Colobinae, p.1-2. London: British Museum (Natural History).

Andrews, P. 1985. Family group systematics and evolution among catarrhine primates. In, (E. Delson, Ed.) Ancestors: the Hard Evidence, pp. 14-22. New York: Alan R. Liss, Inc.

Andrews, P. 1986. Molecular evidence for catarrhine evolution. In (Wood, B., Martin, L. & Andrews, P., Eds.) Major topics in primate and human evolution, pp. 107-129. Cambridge: Cambridge University Press.

Andrews, P. 1987. Aspects of hominoid phylogeny. In, (C. Patterson, Ed.) Molecules and Morphology in Evolution: Conflict or Compromise?, pp. 23-53. Cambridge: Cambridge University Press.

Andrews, P. 1992. Evolution and the environment in the Hominoidea. Nature 360, 641-646.

Andrews, P. 1995. Ecological apes and ancestors. Nature 376, 555-556.

Andrews, P. 1996. The nature of the evidence. Nature 379, 123-124.

Andrews, P. & Williams, D.B. 1973. The use of principal components analysis in physical anthropology. AJPA 39, 291-304.

Andrews, P. & Groves, C. 1976. Gibbons and Brachiation. Gibbon and Siamang, vol. 4, pp. 167-218. Basel: Karger.

Andrews, P. & Cronin, J.E. 1982. The relationships of Sivapithecus and Ramapithecus and the evolution of the orang-utan. Nature 297, 541-546.

Andrews, P. & Aiello, L. 1984. An evolutionary model for feeding and positional behavior. In, (Chivers, D.J., Wood, B.A. & Bilsborough, A., Eds.) Food acquisition and processing in primates, pp. 429-466. New York: Plenum Press.

Andrews, P. & Martin, L. 1987. Cladistic relationships of extant and fossil hominoids. JHE 16, 101-118.

Andrews, P. & Martin, L. 1991. Hominoid dietary evolution. Phil. Tran: Biol. Sci. 334, 199-209.

Andrews, P., Harrison, T., Delson, E., Bernor, R.L. & Martin, L. 1996. Distribution and biochronology of European and southwest Asian Miocene catarrhines. In (Bernor, R.L., Fahlbusch, V. & Mittmann, H-W., Eds.) The evolution of Western Eurasian Neogene Mammal Faunas, pp. 168-207.

Andrews, P., Begun, D.R. & Zylstra, M. 1997. Interrelationships between functional morphology and paleoenvironments in Miocene hominoids. In, (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 29-58. New York: Plenum Press.

423

Andrews, P. & Humphrey, L. 1999. African Miocene environments and the transition to early hominines. In, (Bromage, T.G. & Schrenk, F., Eds.) African biogeography, climate change, & human evolution, pp. 282- 300. New York: Oxford University Press.

Andrews, P. & Harrison, T. 2005. The last common ancestor of apes and humans. In (Lieberman, D.E., Smith, R.J., Kelley, J., Eds.) Interpreting the past: Essays on human, primate, and mammal evolution in honor of David Pilbeam, p. 103-121. Boston: Brill Academic Publishers, Inc.

Ankel-Simons, F. 1999. Primate Anatomy: an introduction, 2nd Ed. San Diego: Academic Press.

Anton, S.C. 2002. Evolutionary significance of cranial variation in Asian Homo erectus. AJPA 118, 301-323.

Anzai, T., Shiina, T., Kimura, N., Yanagiya, K., Kohara, S., Shigenari, A., Yamagata, T., Kulski, J.K., Naruse, T.K., Fujimori, Y., Fukuzumi, Y., Yamazaki, M., Tashiro, H., Iwamoto, C., Umehara, Y., Imanishi, T., Meyer, A., Ikeo, K., Gojobori, T., Bahram, S. & Inoko, H. 2003. Comparative sequencing of human and chimpanzee MHC class I regions unveils insertions/deletions as the major path to genomic divergence. PNAS 100, 7708-7713.

Ardito, G. & Mottura, A. 1987. An overview of the geographic and chronologic distribution of Western European Cercopithecids. Human Evolution 2(1), 29-45.

Arnason, U., Gullberg, A. & Xu, X. 1996a. A complete mitochondrial DNA molecule of the white-handed gibbon, Hylobates lar, and comparison among individual mitochondrial genes of all hominoid genera. Hereditas 124, 185-189.

Arnason, U., Gullberg, A., Janke, A & Xu, X. 1996b. Pattern and timing of evolutionary divergences among hominoids based on analyses of complete mtDNAs. J. Mol. Evol. 43, 650-661.

Arnason, U., Gullberg, A., Burguete, A.S. & Janke, A. 2000. Molecular estimates of primate divergences and new hypotheses for primate dispersal and the origin of modern humans. Hereditas 133, 217-228.

Arnold, M.L. & Meyer, A. Natural hybridization in primates: One evolutionary mechanism. Zoology 109, 261-276.

Arsuaga, J.L., Martínez, I., Gracia, A. & Lorenzo, C. 1997. The Sima de los Huesos crania (Sierra de Atapuerca, Spain). A comparative study. JHE 33, 219-281.

Asfaw, B., White, T., Lovejoy, O., Latimer, B., Simpson, S. & Suwa, G. 1999. Australopithecus garhi: A new species of early hominid from Ethiopia. Science 284, 629-635.

Asfaw, B., Gilbert, W.H., Beyene, Y., Hart, W.K., Renne, P.R., WoldeGabriel, G., Vrba, E.S. & White, T.D. 2002. Remains of Homo erectus from Bouri, Middle Awash, Ethiopia. Nature 416, 317-320

Ashton, E.H. & Oxnard, C.E. 1964a. Locomotor patterns in primates. Pro. Zool. Soc. Lond. 142, 1-28.

Ashton, E.H. & Oxnard, C.E. 1964b. Functional adaptations in the primate shoulder girdle. Pro. Zool. Soc. Lond. 142, 49-66.

Ashton, E.H., Oxnard, C.E. & Spence, T.F. 1965. Scapular shape and primate classification. Pro. Zool. Soc. Lond. 145, 125-142.

Ashton, E.H., Flinn, R.M. & Oxnard, C.E. 1975. The taxonomic and functional significance of overall body proportions in primates. J. Zool., Lond. 175, 73-105.

424 Atchley, W.R., Kunkel, J.G., Cherry, L.M., Case, S.M. & Wilson, A.C. 1980. M-statistics and morphometric divergence. Science 208, 1059-1061.

Avise, J.C. & Aquadro, C.F. 1982. A comparative summary of the genetic distances in the vertebrates. Evol. Biol. 15, 151-184.

Avise, J.C., Walker, D. & Johns, G.C. 1998. Speciation durations and Pleistocene effects on vertebrate phylogeography. Proc. R. Soc. Lond. B 265, 1707-1712.

Avise, J.C. & Johns, G.C. 1999. Proposal for a standardized temporal scheme of biological classification for extant species. PNAS 96, 7358-7363.

Ayala, F.J. & Coluzzi, M. 2005. Chromosome speciation: Humans, Drosophila, and mosquitoes. PNAS 102, 6535-6542.

Backwell, L. & d’Errico, F. 2001. Evidence of termite foraging by Swartkrans early hominids. PNAS 98, 1358-1363.

Bailey, W.J., Fitch, D.H.A., Tagle, D.A., Czelusniak, J., Slightom, J.L. & Goodman, M. 1991. Mol. Biol. Evol. 8(2), 155-184.

Bakewell, M.A., Shi, P. & Zhang, J. 2007. More genes underwent positive selection in chimpanzee evolution than in human evolution. PNAS 104, 7489-7494.

Barry, J.C. 1987. The history and chronology of Siwalik Cercopithecids. Human Evolution 2(1), 47-58.

Bartlett, T.Q. 2007. The Hylobatidae: Small apes of Asia. In, (Campbell, C.J., Fuentes, A., MacKinnon, K.C., Panger, M. & Bearder, S.K., Eds.) Primates in Perspective, pp. 274-289. UK: Oxford University Press.

Barton, R. 1999. The evolutionary ecology of the primate brain. In, (P.C. Lee, Ed.) Comparative primate socioecology, pp. 167-203. UK: Cambridge University Press.

Bass, W.M. 1995. Human Osteology: a laboratory and field manual. USA: Missouri Archaeological Society.

Begun, D.R. 1992a. Miocene fossil hominids and the chimp-human clade. Science 257, 1929-1933.

Begun, D.R. 1992b. Phyletic diversity and locomotion in primitive European hominids. AJPA 87, 311-340.

Begun, D.R. 1994. Relations among the great apes and humans: New interpretations based on the fossil great ape Dryopithecus. Yearbook of Physical Anthropology 37, 11-63.

Begun, D.R. 1995. Late Miocene European orang-utans, gorillas, humans, or none of the above? JHE 29, 169-180.

Begun, D.R. & Kordos, L. 1997. Phyletic affinities and functional convergence in Dryopithecus and other Miocene and living hominids. In, (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 291-316. New York: Plenum Press.

Begun, D.R., Ward, C.V. & Rose, M.D. 1997. Events in hominoid evolution. In, (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 389-415. New York: Plenum Press

Begun, D.R. 2001. African and Eurasian Miocene hominoids and the origins of the Hominidae. In (De Bonis, L., Koufos, G.D. & Andrews, P., Eds.) Hominoid evolution and climatic change in Europe volume 2 Phylogeny of the Neogene hominoid Primates of Eurasia, pp. 231-253. UK: Cambridge University Press.

425 Begun, D.R. 2002a. The Pliopithecoidea. In, (W.C. Hartwig, Ed.) The Primate Fossil Record, pp. 221-240. UK: Cambridge University Press.

Begun, D.R. 2002b. European Hominoids. In, (W.C. Hartwig, Ed.) The Primate Fossil Record, pp. 339-368. UK: Cambridge University Press.

Begun, D.R. 2004a. The earliest hominins – is less more? Science 303, 1478-1480.

Begun, D.R. 2004b. Enhanced cognitive capacity as a contingent fact of hominid phylogeny. In (Russon, A.E. & Begun, D.R., Eds.) The Evolution of Thought: Evolutionary origins of Great Ape Intelligence, pp. 15-28. UK: Cambridge Uni. Press.

Begun, D.R. & Kordos, L. 2004. Cranial evidence of the evolution of intelligence in fossil apes. In (Russon, A.E. & Begun, D.R., Eds.) The Evolution of Thought: Evolutionary origins of Great Ape Intelligence, pp. 260-279. UK: Cambridge Uni. Press.

Benefit, B.R. 1999. Biogeography, dietary specialization, and the diversification of African Plio-Pleistocene monkeys. In, (Bromage, T.G. & Schrenk, F., Eds.) African biogeography, climate change, & human evolution, pp. 172-188. New York: Oxford University Press.

Benefit, B. 2000. Old World monkey origins and diversification: an evolutionary study of diet and dentition. In, (Whitehead, P.F. & Jolly, C.J., Eds.) Old World Monkeys, pp. 133-179. UK: Cambridge Uni. Press.

Benefit, B.R. & McCrossin, M.L. 1993. Facial anatomy of Victoriapithecus and its relevance to the ancestral cranial morphology of old world monkeys and apes. AJPA 92, 329-370.

Benefit, B. & McCrossin, M. 1997. Earliest known Old World monkey skull. Nature 388, 368-371.

Benefit, B.R. & McCrossin, M.L. 2002. The Victoriapithecidae, Cercopithecoidea. In, (W.C. Hartwig, Ed.) The Primate Fossil Record, pp. 241-254. UK: Cambridge University Press.

Bennett, E.L. & Davies, A.G. 1994. The ecology of Asian colobines. In, (Davies, A.G. & Oates, J.F., Eds.) Colobine monkeys: their ecology, behavior and evolution, pp. 129-172. Great Britain: Cambridge Uni. Press.

Benton, M. 2000. Stems, nodes, crown clades, and rank-free lists: is Linnaeus dead? Biological Reviews 75: 633-648.

Benton, M.J. & Pearson, P.N. 2001. Speciation in the fossil record. TREE 16, 405-411.

Berger, L.R. & Tobias, P.V. 1996. A chimpanzee-like tibia from Sterkfontein, South Africa and its implications for the interpretation of bipedalism in Australopithecus africanus. JHE 30, 343-348.

Bernstein, I.S. & Gordon, T.P. 1980. Mixed taxa introductions, hybrids and macaque systematics. In, (D.G. Lindburg, Ed.) The Macaques: Studies in Ecology, Behavior, and Evolution, pp. 10-30. New York: Van Nostrand Reinhold Company.

Bigoni, F., Stanyon, R., Wimmer, R. & Schempp, W. 2003. Chromosome painting shows that the Proboscis monkey (Nasalis larvatus) has a derived karyotypes and is phylogenetically nested within Asian colobines. AJP 60, 85-93.

Bigoni, F., Houck, M.L., Ryder, O.A., Weinberg, J. & Stanyon, R. 2004. Chromosome painting shows that Pygathrix nemaeus has the most basal karyotypes among Asian Colobinae. IJP 25, 679-688.

Bingham, P.M. 1999. Human uniqueness: A general theory. The Quarterly Review of Biology 74(2), 133- 169.

426 Bingham, P.M. 2000. Human evolution and human history: A complete theory. Evolutionary Anthropology 9, 248-257.

Bleisch, W.V. & Jiahua, X. 1998. Ecology and behavior of the Guizhou snub-nosed langur (Rhinopithecus [Rhinopithecus] brelichi), with a discussion of socioecology in the genus. In, (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub-Nosed Monkeys, pp.217-240. Singapore: World Scientific.

Bleisch, W.V., Zhenmei, L., Dierenfield, E.S. & Jiahua, X. 1998. Selected nutrient analysis of plants in the diet of the Guizhou snub-nosed monkey (Rhinopithecus [Rhinopithecus] brelichi). In, (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub-Nosed Monkeys, pp. 241-254. Singapore: World Scientific.

Blue, K.T., McCrossin, M.L. & Benefit, B. 2006. Terrestriality in Middle Miocene context: Victoriapithecus from Maboko Island, Kenya. In, (Ishida, H., Tuttle, R., Pickford, M., Ogihara, N. & Nakatsukasa, M., Eds.) Human Origins and Environmental Background, pp. 45-58. USA: Springer.

Blumenberg, B. & Lloyd, A. 1983. Australopithecus and the origin of genus Homo: Aspects of biometry and systematics with accompanying catalog of tooth metric data. Biosystems 16, 127-167.

Blumenberg, B. 1985. Biometrical studies upon hominoid teeth: The coefficient of variation, sexual dimorphism and questions of phylogenetic relationship. Biosystems 18, 149-184.

Boaz, N.T. 1983. Morphological trends and phylogenetic relationships from Middle Miocene hominoids to Late Pliocene hominids. In, (R.L. Ciochon & R.S. Corruccini, Eds.) New Interpretations of Ape and Human Ancestry, pp. 705-720. New York: Plenum Press.

Bobe, R. & Behrensmeyer, A.K. 2004. The expansion of grassland ecosystems in Africa in relation to mammalian evolution and the origin of the genus Homo. Paleogeography, Paleoclimatology, Paleoecology 207, 399-420.

Bock, W.J. 1981. Functional-adaptive analysis in evolutionary classification. Amer. Zool. 21, 5-20.

Boesch, C. & Boesch, H. 1989. Hunting behavior of wild chimpanzees in the Tai National Park. AJPA 78, 547-573.

Boesch, C. & Tomasello, M. 1998. Chimpanzee and human cultures. Current Anthropology 39, 591-614.

Boesch, C. 2003. Is culture a golden barrier between human and chimpanzee? Evolutionary Anthropology 12, 82-91.

Bogin, B. & Rios, L. 2003. Rapid morphological change in living humans: implications for modern human origins. Comparative Biochemistry and Physiology Part A 136, 71-84.

Bolter, D.R. & Zihlman, A.L. 2003. Morphometric analysis of growth and development in wild-collected vervet monkeys (Cercopithecus aethiops), with implications for growth patterns in Old World monkeys, apes and humans. Journal of Zoology 260(1), 99-110.

Bolter, D. & Zhilman, A. 2007. Primate growth and development: A functional and evolutionary approach. In, (Campbell, C.J., Fuentes, A., MacKinnon, K.C., Panger, M. & Bearder, S.K., Eds.) Primates in Perspective, pp. 408-422. UK: Oxford University Press.

Boonratana, R. & Canh, L.X. 1998. Preliminary observations of the ecology and behavior of the Tonkin snub- nosed monkey (Rhinopithecus [Presbyticus] avunculus) in Northern Vietnam. In, (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub-Nosed Monkeys, pp. 207-216. Singapore: World Scientific.

Brace, C.L. 2005. “Neutral theory” and the dynamics of the evolution of “Modern” human morphology. Human Evolution 20, 19-38.

427

Bramble, D. & Lieberman, D. 2004. Endurance running and the evolution of Homo. Nature 432, 345-352.

Brandon-Jones, D. 1996. The Asian Colobinae (Mammalia: Cercopithecidae) as indicators of Quaternary climatic change. BJLS 59, 327-350.

Brandon-Jones, D. 2004. A taxonomic revision of the langurs and leaf-monkeys (Primates: Colobinae) South Asia. Zoos’ Print Journal 19(8), 1552-1594.

Brandon-Jones, D., Eudey, A.A., Geissmann, T., Groves, C.P., Melnick, D.J., Morales, J.C., Shekelle, M. & Stewart, C.-B. 2004. Asian primate classification. IJP 25, 97-164.

Britten, R.J. 2002. Divergence between samples of chimpanzee and human DNA sequences is 5% counting indels. PNAS 90, 13633-13635.

Bromage, T.G. 1999. Habitat specificity and early hominid craniodental ecomorphology. In, (Bromage, T.G. & Schrenk, F., Eds.) African biogeography, climate change, & human evolution, pp. 364-368. New York: Oxford University Press.

Broom, R., 1940. The South African Pleistocene cercopithecid apes. Annals of the Transvaal Museum 20, 89-100.

Broom, R. 1950. The genera and species of the South African fossil ape-men. AJPA 8(1), 1-14.

Brown, F., Harris, J., Leakey, R. & Walker, A. 1985. Early Homo erectus skeleton from west Lake Turkana, Kenya. Nature 316, 788-792.

Brown, B. 1997. Miocene hominoid mandibles: Functional and Phylogenetic perspectives. In, (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 153-171. New York: Plenum Press.

Brown, P., Sutikna, T., Morwood, M., Soejono, R.P., Jatmiko, Saptomo, E.W. & Due, R.A. 2004. A new small-bodied hominin from the Late Pleistocene of Flores, Indonesia. Nature 431, 1055-1061.

Brown, B., Kappelman, J. & Ward, S. 2005. Lots of faces from different places: What craniofacial morphology does(n’t) tell us about hominoid phylogenetics. In (Lieberman, D.E., Smith, R.J., Kelley, J., Eds.) Interpreting the past: Essays on human, primate, and mammal evolution in honor of David Pilbeam, p. 167- 187. Boston: Brill Academic Publishers, Inc.

Bruce, E.J. & Ayala, F.J. 1978. Humans and apes are genetically very similar. Nature6 27, 264-265.

Brunet, M., Beauvilan, A., Coppens, Y., Heintz, E., Moutaye, A.H.E. & Pilbeam, D. 1995. The first australopithecine 2,500 kilometers west of the Rift Valley (Chad). Nature 378, 273-240.

Brunet, M., Guy, F., Pilbeam, D., Mackaye, H.T., Likius, A., Ahounta, D., Beauvilain, A., Blondel, C., Bocherens, H., Boisserie, J., De Bonis, L., Coppens, Y., Dejax, J., Denys, C., Duringer, P., Eisenmann, V., Fanone, G., Fronty, P., Geraads, D., Lehmann, T., Lihoreau, F., Louchart, A., Mahamat, A., Merceron, G., Mouchelin, G., Otero, O., Campomanes, P., Ponce De Leon, M., Rage, J., Sapanet, M., Schuster, M., Sudre, J., Tassy, P., Valentin, X., Vignaud, P., Viriot, L., Zazzo, A. & Zollikofer, C. 2002. A new hominid from the Upper Miocene of Chad, Central Africa. Nature 418, 145-151.

Burney, D.A. 1995. Historical perspectives on human-assisted biological invasions. Evolutionary Anthropology 5, 216-221.

Buettner-Janusch, J. 1963. A problem in evolutionary systematics: Nomenclature and classification of baboons. Folia Primatol. 4: 288–308.

428

Buikstra, J.E. & Ubelaker, D.H., Eds. 1994. Standards for data collection from human skeletal remains. USA: Arkansas Archaeological Survey Research Series No. 44.

Butynski, T.M. 2002. The Guenons: an overview of diversity and taxonomy. In, (Glenn, M.E. & Cords, M., Eds.) The Guenons: Diversity and adaptation in African monkeys, pp. 3-14. New York: Kluwer Academic / Plenum Publishers.

Buzzard, P.J. 2006a. Ecological partitioning of Cercopithecus campbelli, C. petaurista, and C. diana in the Tai Forest. IJP 27, 529-558.

Buzzard, P.J. 2006b. Cheek pouch use in relation to interspecific competition and predator risk for three guenon monkeys (Cercopithecus spp.). Primates 47, 336-341.

Byron, C.D. & Covert, H.H. 2004. Unexpected locomotor behaviour: Brachiation by an Old World monkey (Pygathrix nemaeus) from Vietnam. J. Zool., Lond. 263, 101-106.

Caccone, A. & Powell, J.R. 1989. DNA divergence among hominoids. Evolution 43, 925-942.

Caceres, M., Lachuer, J., Zapala, M.A., Redmond, J.C., Kudo, L., Geschwind, D.H., Lockhart, D.J., Preuss, T.M. & Barlow, C. 2003. Elevated gene expression levels distinguish human from non-human primate brains. PNAS 100, 13030-13035.

Caldecott, J.O., Feistner, T.C. & Gadsby, E.L. 1996. A comparison of ecological strategies of pig-tailed macaques, mandrills and drills. In (Fa, J.E. & Lindburg, D.G., Eds.) Evolution and Ecology of Macaque Societies, pp. 73-94. UK: Cambridge Uni. Press.

Cameron, D.W. 2003. Early hominin speciation at the Plio/Pleistocene transition. Homo 54, 1-28.

Campbell, B. 1962. The systematics of man. Nature 194, 225-232.

Campbell, B. 1963. Quantitative taxonomy and human evolution. In, (S. Washburn, Ed.) Classification and Human Evolution, pp. 50-74. Chicago: Aldine Publishing Co.

Campbell, B. 1965. The nomenclature of the hominidae. Royal Anthropological Institute of Great Britain and Ireland 22, 1-33.

Campbell, N.A. 1978. Multivariate analysis in biological anthropology: Some further considerations. JHE 7, 197-203.

Cantalupo, C. & Hopkins, W.D. 2001. Asymmetric Broca’s area in great apes. Nature 414, 505.

Cantino, P.D., Bryant, H.N., de Queiroz, K., Donoghue, M.J., Eriksson, T., Hillis, D.M. & Lee, M.S. 1999. Species names in phylogenetic nomenclature. Systematic Biology 48, 790-807.

Cardini, A., Jansson, A-U., Elton, S. 2007. A geometric morphometric approach to the study of ecogeographical and clinal variation in vervet monkeys. Journal of Biogeography

Carrier, D.R. 2007. The short legs of great apes: evidence for aggressive behavior in australopiths. Evolution, 596-605.

Cartmill, M. 1990. Human uniqueness and theoretical content in paleoanthropology. IJP 11, 173-192.

Cartmill, M. 2001. Taxonomic revolution and the animal-human boundary. In, (Corbey & Roebroeks, Eds.) Studying Human Origins: Disciplinary History and Epistemology, pp. 97-106. Amsterdam University Press.

429 Cartmill, M. 2002. Paleoanthropology: science or mythological charter? Journal of Anthropological Research 58, 183-201.

Carroll, S.B. 2003. Genetics and the making of Homo sapiens. Nature 422, 849-857.

Castresana, J. 2001. Cytochrome b phylogeny and the taxonomy of the great apes and mammals. Mol. Biol. Evol. 18, 465-471.

Caton, J.M. 1999. Digestive strategy of the Asian colobine genus Trachypithecus. Primates 40(2), 311-325.

Cela-Conde, C. 1996. Bipedal/Savanna/Cladogeny Model. Can it still be held? Hist. Phil. Life Sci. 18, 213- 224.

Cela-Conde, C. 1998. The problem of hominoid systematics, and some suggestions for solving it. SAJS 94, 255-262.

Cela-Conde, C.J. 2001. Hominid taxon and systematics of the Hominoidea. In, (P.V. Tobias, M.A. Raath, J. Moggi-Cecchi & G.A. Doyle, Eds.) Humanity from African Naissance to Coming Millenia, pp. 271-278, Firenze University Press and Witwatersrand University Press.

Cela-Conde, C.J. & Altaba, C.R. 2002. Multiplying genera versus moving species: a new taxonomic proposal for the family Hominidae. SAJS 98, 229-232.

Cela-Conde, C.J. & Ayala, F.A. 2003. Genera of the human lineage. PNAS 100, 7684-7689.

Chaimanee, Y., Jolly, D., Benammi, M., Tafforeau, P., Duzer, D., Moussa, I. & Jaeger, J. 2003. A Middle Miocene hominoid from Thailand and orangutan origins. Nature 422, 61-65.

Chaimanee, Y., Suteethorn, V., Jintasakul, P., Vidthayanon, C., Marandat, B. & Jaeger, J.-J. 2004. A new orang-utan relative from the Late Miocene. Nature 427, 439-441.

Chamberlain, A.T. & Wood, B. 1987. Early hominid phylogeny. JHE 16, 119-133.

Chaplin, G. & Jablonski, N.G. 1998. The integument of the “odd-nosed” colobines. In, (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub-Nosed Monkeys, pp. 79-104. Singapore: World Scientific.

Chapman, C.A., Chapman, L.J., Cords, M., Gathua, M.J., Gauthier-Horn, A., Lambert, J.E., Rode, K., Tutin, C.E.G. & White, L.J.T. 2002. Variation in the diets of Cercopithecus species: differences within forest, among forests, and across species. In, (Glenn, M.E. & Cords, M., Eds.) The Guenons: Diversity and adaptation in African monkeys, pp. 325-350. New York: Kluwer Academic / Plenum Publishers.

Chatterjee, H.J. 2006. Phylogeny and biogeography of gibbons: a dispersal-vicariance analysis. IJP 27, 699- 712.

Chazan, M. 1995. The meaning of Homo sapiens. In, (R. Corbey & B. Theunissen, Eds.) Ape, Man, Apeman: Changing Views since 1600, pp. 229-240. Leiden University: Department of Prehistory.

Chen, F.-C., Vallender, E.J., Wang, H., Tzeng, C.-S. & Li, W.-H. 2001. Genomic divergence between human and chimpanzee estimated from large-scale alignments of genomic sequences. Journal of Heredity 92, 481-489.

Chen, F. & Li, W. 2001. Genomic divergences between humans and other hominoids and the effective population size of the common ancestor of humans and chimpanzees. Am. J. Hum. Genet. 68, 444-456.

Cherry, L.M., Case, S.M. & Wilson, A.C. 1978. Frog perspectives on the morphological differences between humans and chimps. Science 200, 209-211.

430

Cherry, L.M., Case, S.M., Kunkel, J.G., Wyles, J.S. & Wilson, A.C. 1982. Body shape metrics and organismal evolution. Evolution 36(5), 914-933.

Cheverud, J.M. 1982. Phenotypic, genetic, and environmental morphological integration in the cranium. Evolution 36, 499-516.

Cheverud, J.M. 1989. A comparative analysis of morphological variation patterns in the Papionins. Evolution 43, 1737-1747.

Cheyne, S.M. & Chivers, D.J. 2006. Sexual swellings of female gibbons. Folia primat. 77(5), 345-352.

Chiarelli, B. 1985. Chromosomes and the origin of man. In, (P.V. Tobias, Ed.) Hominid Evolution: Past, Present and Future, pp. 397-400. New York: Alan R. Liss, Inc

[The] Chimpanzee Sequencing and Analysis Consortium. 2005. Initial sequence of the chimpanzee genome and comparison with the human genome. Nature 437, 69-87.

Chism, J. & Rowel, T.E. 1988. The natural history of the patas monkey. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 412-438. UK: Cambridge Uni. Press.

Chivers, D.J. 1974. The siamang in Malaya: A field study of a primate in tropical rain forest. Contrib. to Primatol. 4, 1-335.

Chopra, S.R.K., Kaul, S. & Pathak, R.K. 1982. Morphometric affinities in innominate bones of the Old World Primates including man. JHE 11, 105-108.

Chou, H., Takematsu, H., Diaz, S., Iber, J., Nickerson, E., Wright, K.L., Muchmore, E.A., Nelson, D.L., Warren, S.T. & Varki, A. 1998. A mutation in human CMP-sialic acid hydroxylase occurred after the Homo- Pan divergence. PNAS 95, 11751-11756.

Chou, H., Hayakawa, T., Diaz, S., Krings, M., Indriati, E., Leakey, M., Paabo, S., Satta, Y., Takahata, N. & Varki, A. 2002. Inactivation of CMP-N-acetylneuraminic acid hydroxylase occurred prior to brain expansion during human evolution. PNAS 99, 11736-11741.

Ciochon, R.L. & Chiarelli, A.B. 1980. Paleobiogeographic perspectives on the origin of the Platyrrhini. In, (Ciochon, R.L. & Chiarelli, A.B., Eds.) Evolutionary biology of the New World monkeys and continental drift, pp. 459-493. New York: Plenum Press.

Ciochon, R.L. 1983. Hominoid cladistics and the ancestry of modern apes and humans: A summary statement. In, (R.L. Ciochon & R.S. Corruccini, Eds.) New Interpretations of Ape and Human Ancestry, pp. 783-843. New York: Plenum Press.

Ciochon, R.L. 1993. Evolution of the cercopithecoid forelimb: Phylogenetic and functional implications from morphometric analyses. Berkley: University of California Press.

Clark, A.G., Glanowski, S., Nielsen, R., Thomas, P.D., Kejariwal, A., Todd, M.A., Tanenbaum, D.M., Civello, D., Lu, F., Murphy, B., Ferriera, S., Wang, G., Zheng, X., White, T.J., Sninsky, J.J., Adams, M.D., & Cargill, M. 2003. Inferring nonneutral evolution from Human-Chimp-Mouse orthologous gene trios. Science 302, 1960-1963.

Clarke, R.J. 1985. Australopithecus and early Homo in Southern Africa. In, (E. Delson, Ed.) Ancestors: the Hard Evidence, pp. 171-177. New York: Alan R. Liss, Inc.

431 Clifford, S.L., Abernethy, K.A., White, L.J.T., Tutin, C.E.G., Bruford, M.W. & Wickings, E.J. 2003. Genetic studies of western gorillas. In, (Taylor, A. & Goldsmith, M.L., Eds.) Gorilla biology: a multidisciplinary perspective, pp. 269-292. UK: Cambridge Uni. Press.

Clutton-Block, T.H. 1974. Primate social organization and ecology. Nature 250, 539-542.

Coe, M.J. 1984. Primates: their niche structure and habitats. In, (Chivers, D.J., Wood, B.A. & Bilsborough, A., Eds.) Food acquisition and processing in primates, pp. 1-32. New York: Plenum Press.

Collard, M. 2002. Grades and transitions in human evolution. Proceedings of the British Academy 106, 61- 100.

Collard, M. & Wood, B. 1999. Grades among the African early hominids. In, (Bromage, T.G. & Schrenk, F., Eds.) African biogeography, climate change, & human evolution, pp. 316-327. New York: Oxford University Press.

Collard, M. & Wood, B. 2000. How reliable are human phylogenetic hypotheses? PNAS 97, 5003-5006.

Collard, M. & Wood, B.A. 2001a. Homoplasy and the early hominid masticatory system: Inferences from analyses of extant hominoids and papionins. JHE 41, 167-194.

Collard, M. & Wood, B.A. 2001b. How reliable are current estimates of fossil catarrhine phylogeny? An assessment using great apes and Old World monkeys. In (De Bonis, L., Koufos, G.D. & Andrews, P., Eds.) Hominoid evolution and climatic change in Europe volume 2 Phylogeny of the Neogene hominoid Primates of Eurasia, pp. 118-150. UK: Cambridge University Press.

Collins, D.A. & McGrew, W.C. 1988. Habitat of three groups of chimpanzees (Pan troglodytes) in western Tanzania compared. JHE 17, 553-574.

Colyn, M. & Deleporte, P. 2002. Biogeographic analysis of central African forest guenons. In, (Glenn, M.E. & Cords, M., Eds.) The Guenons: Diversity and adaptation in African monkeys, pp. 61-78. New York: Kluwer Academic / Plenum Publishers.

Conroy, G., Pickford, M., Senut, B., van Couvering, J. & Mein, P. 1992. Otavipithecus namibiensis, first Miocene hominoid from southern Africa. Nature 356, 144-147.

Conroy, G. 1990. Primate evolution. New York: W-W Norton & Company.

Conroy, G. 2002. Speciosity in the early Homo lineage: too many, too few, or just about right? JHE 43, 759- 766.

Conroy, G. 2003. The inverse relationship between species diversity and body mass: do primates play by the ‘rules’? JHE 45, 43-55.

Conroy, G. 2005. Reconstructing Human Origins, 2nd edition. New York: W.W. Norton.

Constantino, P. & Wood, B. 2007. The evolution of Zinjanthropus boisei. Evolutionary Anthropology 16, 49- 62.

Coolidge, H.J. & Shea, B. 1982. External body dimensions of Pan paniscus and Pan troglodytes chimpanzees. Primates 23(2), 245-251.

Cope, D.A. 1993. Measures of dental variation as indicators of multiple taxa in samples of sympatric Cercopithecus species. In, (W.H. Kimbel & B.M. Lawrence, Eds.) Species, Species Concepts, and Primate Evolution, pp. 211-237. New York: Plenum Press.

432 Cope, D.A. & Lacy, M.G. 1995. Comparative application of the coefficient of variation and range-based statistics for assessing the taxonomic composition of fossil samples. JHE 29, 549-576.

Coqueugniot, H., Hublin, J.-J., Veillon, F., Houët, F. & Jacob, T. 2004. Early brain growth in Homo erectus and implications for cognitive ability. Nature 431, 299-302.

Cords, M. 1988. Mating systems of forest guenons: a preliminary review. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 323-339. UK: Cambridge Uni. Press.

Corruccini, R.S. 1975. Multivariate analysis in biological anthropology: some considerations. JHE 4, 1-19.

Corruccini, R.S. 1994. How certain are hominoid phylogenies? The role of confidence intervals in cladistics. In, (Corruccini, R.S. & Ciochon, R.L., Eds.) Integrative paths to the past: Paleoanthropological advances in honor of F. Clark Howell, pp. 167-183.

Corruccini, R.S. & Ciochon, R. 1979. Primate facial allometry and interpretations of australopithecine variation. Nature 281, 62-64.

Cote, S.M. 2004. Origins of the African hominoids: An assessment of the paleobiogeographical evidence. C.R. Palevol 3, 321-338.

Coyne, J.A. 1992. Genetics and speciation. Nature 355, 511-515.

Cracraft, J. 1983. Species concepts and speciation analysis. In, (Johnston, R.F., Ed.) Current Ornithology Vol. 1, pp. 159-187. New York: Plenum Press.

Cramer, D.L. 1977. Craniofacial morphology of Pan paniscus. Contributions to Primatology 10, 1-64.

Creel, N. & Preuschoft, H. 1971. Hominoid taxonomy: A canonical analysis of cranial dimensions. Proc. 3rd int. Congr. Primat., Zurich 1970, vol. 1, pp. 36-43.

Creel, N. & Preuschoft, H. 1976. Cranial morphology of the lesser apes. Gibbon and Siamang, vol. 4, pp. 219- 303. Basel: Karger.

Cronin, J.E., Boaz, N.T., Stringer, C.B. & Rak, Y. 1981. Tempo and mode in hominid evolution. Nature 292, 113-122.

Cronin, J.E. & Sarich, V.M. 1976. Molecular evidence for dual origin of mangabeys among Old World monkeys. Nature 260, 700-702.

Cronin, J.E. & Meikle, W.E. 1979. The phyletic position of Theropithecus: Congruence among molecular, morphological, and paleontological evidence. Systematic Zoology 28, 259-269.

Cronin, J.E., Cann, R. & Sarich, V.M. 1980. Molecular evolution and systematics of the genus Macaca. In, (D.G. Lindburg, Ed.) The Macaques: Studies in Ecology, Behavior, and Evolution, pp. 31-51. New York: Van Nostrand Reinhold Company.

Cronin, J.E. & Meikle, W.E. 1982. Hominid and gelada baboon evolution: Agreement between molecular and fossil time scales. IJP 3(4), 469-482.

Cronin, J.E. 1983. Apes, humans, and molecular clocks: A reappraisal. In, (R.L. Ciochon & R.S. Corruccini, Eds.) New Interpretations of Ape and Human Ancestry, pp. 115-150. New York: Plenum Press.

Curnoe, D. 2003. Problems with the use of cladistic analysis in palaeoanthropology. Homo 53(3), 225-234.

433 Curnoe, D. & Thorne, A. 2003. Number of ancestral human species: a molecular perspective. Homo 53(3), 201-224.

Curnoe, D., Thorne, A. & Coate, J.A. 2006. Timing and tempo of primate speciation. Journal of Evolutionary Biology 19, 59-65.

Curnoe, D. & Tobias, P.V. 2006. Description, new reconstruction, comparative anatomy, and classification of the Sterkfontein Stw 53 cranium, with discussions about the taxonomy of other southern African early Homo remains. JHE 50(1), 36-77.

Curten, S.H. 1976. Niche separation in sympatric Malaysian leaf-monkeys (Presbytis obscura and Presbytis melalophos). Yearbook of Physical Anthropology 20, 421-439.

Czarnetzki, A., Jakob, T. & Pusch, C.M. 2003. Palaeopathological and variant conditions of the Homo heidelbergensis type specimen (Mauer, Germany). JHE 44, 479-295.

Dainton, M. & Macho, G.A. 1999a. Did knuckle-walking evolve twice? JHE 36, 171-194.

Dainton, M. & Macho, G.A. 1999b. Heterochrony: somatic, skeletal and dental development in Gorilla, Homo and Pan. In (Hoppa & Fitzgerald, Eds.) Human growth in the past, pp. 32-64. UK: Cambridge Uni. Press.

Darroch, J.N. & Mosimann, J.E. 1985. Canonical and principal components of shape. Biometrika 72, 241- 252.

Dart, R.A. 1925. Australopithecus africanus: the man-ape of South Africa. Nature 115, 235-236.

Davenport, T.R.B., Stanley, W.T., Sargis, E.J., De Luca, D.W., Mpunga, N.E., Machaga, S.J. & Olson, L.E. 2006. A new genus of African monkey, Rungwecebus: morphology, ecology, and molecular phylogenetics. Science 312, 1378-1381.

Davies, A.G. & Oates, J.F., Eds. 1994. Colobine monkeys: their ecology, behavior and evolution. Great Britain: Cambridge Uni. Press.

Davies, A.G., Oates, J.F. & Dasilva, G.L. 1999. Patterns of frugivory in three West African colobine monkeys. IJP 20(3), 327-357.

Davies, A.G. 1994. Colobine populations. In, (Davies, A.G. & Oates, J.F., Eds.) Colobine monkeys: their ecology, behavior and evolution, pp. 285-310. Great Britain: Cambridge Uni. Press.

Davison, G.W.H. 1982. Convergence with terrestrial cercopithecines by the monkey Rhinopithecus roxellanae. Folia Primatol. 37, 209-215.

Day, M.H. 1967. Olduvai hominid 10: a multivariate analysis. Nature, 323-324.

Day, M.H. 1969. Femoral fragment of a Robust Australopithecine from Olduvai Gorge, Tanzania. Nature 221, 230-233.

Day, M.H. 1971. Postcranial remains of Homo erectus from Bed IV, Olduvai Gorge, Tanzania. Nature 232, 383-387.

Day, M.H. & Leakey, R.E.F. 1973. New evidence of the genus Homo from East Rudolf, Kenya, I. AJPA 39, 341-354.

Day, M.H., Leakey, R.E.F., Walker, A.C. & Wood, B.A. 1975. New hominids from East Rudolf, Kenya, I. AJPA 42, 461-476.

434

Day, M.H., Leakey, R.E.F., Walker, A.C. & Wood, B.A. 1976. New hominids from East Turkana, Kenya. AJPA 45, 369-436.

Day, M.H., Leakey, M.D., Olson, T.R., Leakey, R.E.F., Walker, A., McHenry, H.M., Corruccini, R.S., Johanson, D.C., White, T.D. 1980. On the Status of Australopithecus afarensis. Science 207, 1102-1105.

De Bonis, L. 1983. Phyletic relationships of Miocene hominoids and higher primate classification. In, (R.L. Ciochon & R.S. Corruccini, Eds.) New Interpretations of Ape and Human Ancestry, pp. 625-649. New York: Plenum Press.

De Bonis, L., Bouvrian, G., Geraads, D. & Koufos, G. 1990. New hominoid skull material from the late Miocene of Macedonia in Northern Greece. Nature 345, 712-714.

De Bonis, L. & Kuofos, G. 1993. The face and the mandible of Ouranopithecus macedoniensis: description of new specimens and comparisons. JHE 24, 469-491.

De Bonis, L. & Kuofos, G. 1997. The phylogenetic and functional implications of Ouranopithecus macedoniensis. In, (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 317-328. New York: Plenum Press

De Heinzelin, J., Clark, J.D., White, T., Hart, W., Renne, P., WoldeGabriel, G., Beyene, Y. & Vrba, E. 1999. Environment and behavior of 2.5-million-year-old Bouri Hominids. Science 284, 625-629.

De Lumley, M.-A., Gabounia, L., Vekua, A. & Lordkipanidze, D. 2006. Human remains from the Upper Pliocene-Early Pleistocene Dmanisi site, Georgia (1991-2000). Part I: the fossil skulls (D 2280, D 2282 and d 2700). L’anthropologie 110, 1-110.

De Queiroz, K. & Gautheir, J. 1992. Phylogenetic taxonomy. Ann. Rev. Ecol. Syst. 23, 449-480.

De Queiroz, K. & Good, D.A. 1997. Phenetic clustering in biology: A critique. The Quarterly Review of Biology 72 (1), 3-30.

Dean, D. & Delson, E. 1992. Second gorilla of third chimp? Nature 359, 676-677.

Dean, M.C. 1988a. Growth of teeth and development of the dentition in Paranthropus. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.43-53. New York: Aldine de Gruyter.

Dean, M.C. 1988b. Growth processes in the cranial base of hominoids and their bearing on morphological similarities that exist in the cranial base of Homo and Paranthropus. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.107-112. New York: Aldine de Gruyter.

Dean, C., Leakey, M.G., Reid, D., Schrenk, F., Schwartz, G.T., Stringer, C. & Walker, A. 2001. Growth processes in teeth distinguish modern humans from Homo erectus and earlier hominins Nature 414, 628-631.

Deinard, A. & Kidd, K. 1999. Evolution of a HOXB6 intergenic region within the great apes and humans. JHE 36, 687-703.

Deinard, A. & Smith, D.G. 2001. Phylogenetic relationships among the macaques: Evidence from the nuclear locus NRAMP1 JHE 41, 54-59.

Deloison, Y. 1985. Comparative study of calcanei of primates and Pan-Australopithecus-Homo relationship. In, (P.V. Tobias, Ed.) Hominid Evolution: Past, Present and Future, pp. 143-147. New York: Alan R. Liss, Inc.

435 DelPero, M., Masters, J.C., Zuccon, D., Cervella, P., Crovella, S. & Ardito, G. 2000. Mitochondrial sequences as indicators of generic classification in Bush Babies. IJP 21, 889-904.

Delson, E. 1975. The evolutionary history of the Cercopithecidae. Contrib. Primat. Vol. 5, pp.167-217.

Delson, E. & Andrews, P. 1975. Evolution and interrelationships of the Catarrhine Primates. In (Luckett & Szalay, Eds.) Phylogeny of the Primates: a multidisciplinary approach, pp.405-446. New York: Plenum Press.

Delson, E. 1977. Catarrhine phylogeny and classification: principles, methods and comments. JHE 6, 433- 459.

Delson, E., Eldredge, N. & Tattersall, I. 1977. Reconstruction of hominid phylogeny: A testable framework based on cladistic analysis. JHE 6, 263-278.

Delson, E. 1980. Fossil macaques, phyletic relationships and a scenario of development. In, (D.G. Lindburg, Ed.) The Macaques: Studies in Ecology, Behavior, and Evolution, pp. 10-30. New York: Van Nostrand Reinhold Company.

Delson, E. 1985. Catarrhine evolution. In, (Delson, E., Ed.) Ancestors: The hard evidence pp. 9-13. NY: Alan R. Liss, Inc.

Delson, E. 1993. Theropithecus fossils from Africa and India and the taxonomy of the genus. In, (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 157-189. UK: Cambridge University Press.

Delson, E. & Hoffstetter, R. 1993. Theropithecus from Ternifine, Algeria. In, (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 191-208. UK: Cambridge University Press.

Delson, E. 1994. Evolutionary history of the colobine monkeys in paleoenvironmental perspective. In (Davies, A.G. & Oates, J.F., Eds.) Colobine monkeys: their ecology, behavior and evolution, pp. 11-44. Great Britain: Cambridge Uni. Press.

Delson, E. & Andrews, P. 1975. Evolution ad interrelationships of the catarrhine primates. In, (Luckett, W.P. & Szalay, F.S., Eds.) Phylogeny of the Primates: A multidisciplinary approach, pp. 405-446.

Delson, E., Eldredge, N. & Tattersall, I. 1977. Reconstruction of Hominid phylogeny: A testable framework based on cladistic analysis. JHE 6, 263-278.

Delson, E. & Rosenberger, A.L. 1980. Are there any anthropoid primate living fossils. In, (Ciochon, R.L. & Chiarelli, A.B., Eds.) Evolutionary biology of the New World monkeys and continental drift, pp. 50-61. New York: Plenum Press.

Delson, E. & Dean, D. 1993. Are Papio baringensis R. Leakey, 1969, and P. quadratirostris Iwamoto, 1982, species of Papio or Theropithecus? In, (N. Jablonski, Ed.) Theropithecus: The Rise and Fall of a Primate Genus, pp. 245-270. Cambridge: Cambridge University Press.

Delson, E., Terranova, C.J., Jungers, W.L., Sargis, E., Jablonski, N. & Dechow, P.C. 2000. Body mass in Cercopithecidae (Primates, Mammalia): Estimation and scaling in extinct and extant taxa. American Museum of Natural History, Anthropological Papers, Number 83, pp. 1-159.

DeSilva, J., Shoreman, E. & MacLatchy, L. 2006. A fossil hominoid proximal femur from Kikorongo Crater, southerwestern Uganda. JHE 50, 687-695.

Detwiler, K.M. 2002. Hybridization between Red-tailed monkeys (Cercopithecus ascanius) and Blue monkeys (C. mitis) in East African forests. In, (Glenn, M.E. & Cords, M., Eds.) The Guenons: Diversity and adaptation in African monkeys, pp. 79-98. New York: Kluwer Academic / Plenum Publishers.

436

Di Fore, A. & Gagneux, P. 2007. Molecular primatology. In, (Campbell, C.J., Fuentes, A., MacKinnon, K.C., Panger, M. & Bearder, S.K., Eds.) Primates in Perspective, pp. 369-393. UK: Oxford University Press.

Diamond, J. 1988. DNA-based phylogenies of the three chimpanzees. Nature 332, 685-686.

Diamond, J. 1992. The rise and fall of the third chimpanzee. London: Vintage.

Disotell, T., Honeycutt, R.L. & Ruvolo, M. 1992. Mitochondrial DNA phylogeny of the Old-World Monkey tribe Papionini. Mol. Biol. Evol. 9(1), 1-13.

Disotell, T. 1994. Generic level relationships of the Papionini (Cercopithecoidea). AJPA 94, 47-57.

Disotell, T. 2000. The molecular systematics of the Cercopithecidae. In, (Whitehead, P.F. & Jolly, C.J., Eds.) Old World Monkeys, pp. 29-56. UK: Cambridge Uni. Press.

Disotell, T.R. & Raaum, R.L. 2002. Molecular timescale and gene tree incongruence in the guenons. In, (Glenn, M.E. & Cords, M., Eds.) The Guenons: Diversity and adaptation in African monkeys, pp. 27-36. New York: Kluwer Academic / Plenum Publishers.

Dobzhansky, T. 1963. Genetic entities in hominid evolution. In (S. Washburn, Ed.) Classification and Human Evolution, pp. 347-362. Chicago: Aldine Publishing Co.

Dorus, S., Vallender, E.J., Evans, P.D., Anderson, J.R., Gilbert, S.L., Mahowald, M., Wyckoff, G.J. Malcom, C.M., Lahn, B.T. 2004. Accelerated evolution of nervous system genes in the origin of Homo sapiens. Cell 119, 1027–1040.

Dover, G. 1999. Human evolution: our turbulent genes and why we are not chimps. In (Sykes, B., Ed.) The Human Inheritance, pp. 75-92.

Duarte, C., Maurício, J., Pettitt, P.B., Souto, P., Trinkaus, E., van der Plicht, H. & Zilhão, J. 1999. The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia. PNAS 96, 7604-7609.

Duchin, L.E. 1990. The evolution of articulate speech: comparative anatomy of the oral cavity in Pan and Homo. JHE 19, 687-697.

Dunbar, R.I.M. & Dunbar, E.P. 1974. Ecological relations and niche separation between sympatric primates in Ethiopia. Folia primat. 21, 36-60.

Dunbar, R.I.M. 1983. Theropithecines and hominids: contrasting solutions to the same ecological problem. JHE 12, 647-658.

Dunbar, R.I.M. 1992a. Neocortex size as a constraint on group size in primates. JHE 20, 469-493.

Dunbar, R.I.M. 1992b. Behavioural ecology of the extinct papionins. JHE 22, 407-421.

Dunbar, R.I.M. 1993. Social organization of the gelada. In, (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 425-439. UK: Cambridge University Press.

Dunham, A., Matthews, L.H., Burton, J., Ashurst, J.L., Howe, K.L., Ashcroft, K.J., Beare, D.M., Burford, D.C., Hunt, S.E., Griffiths-Jones, S., Jones, M.C., Keenan, S.J., Oliver, K., Scott, C. E., Ainscough, R., Almeida, J.P., Ambrose, K.D., et al. 2004. The DNA sequence and analysis of human chromosome 13. Nature 428, 522 – 528.

437 Dutrillaux, B., Mulersis, M. & Couturier, J. 1988. Chromosomal evolution of the Cercopithecinae. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 150-159. UK: Cambridge Uni. Press.

Dunsworth, H. & Walker, A. 2002. Early genus Homo. In, (W.C. Hartwig, Ed.) The Primate Fossil Record, pp. 419-436. UK: Cambridge University Press.

Easteal, S., Collet, C. & Betty, D. 1995. The mammalian molecular clock. New York: Springer-Verlag.

Eck, G. & Howell, F.C. 1972. New fossil Cercopithecus material from the Lower Omo Basin, Ethiopia. Folia primat. 18, 325-355.

Eck, G.G. 1976. Cercopithecoidea from Omo Group Deposits. In (Y. Coppens, F.C. Howell, G.Ll. Isaac & R.E.F. Leakey, eds.), Earliest Man and Environments in the Lake Rudolf Basin: Stratigraphy, Paleoecology, and Evolution, pp. 332-344. Chicago: University of Chicago Press.

Eck, G.G. 1977. Diversity and Frequency Distribution of Omo Group Cercopithecoidea. JHE 6, 55-63.

Eck, G.G. 1993. Theropithecus darti from the Hadar Formation, Ethiopia. In, (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 15-83. UK: Cambridge University Press.

Eckhardt, R. 2000. Human Paleobiology. UK: Cambridge University Press.

Elango, N., Thomas, J.W., NISC Comparative sequencing Program & Yi, S.V. 2006. Variable molecular clocks in hominoids. PNAS 103, 1370-1375.

Eldredge, N. & Tattersall, I. 1975. Evolutionary models, phylogenetic reconstruction, and another look at the hominid phylogeny. Contrib. Primat. Vol. 5, pp. 218-243.

Elmore, K.L. & Richman, M.B. 2001. Euclidean distance as a similarity metric for principal component analysis. Monthly Weather Review 129, 540-549.

Elton, S. 2001. Locomotor and habitat classifications of cercopithecoid postcranial material from Sterkfontein member 4, Bolt’s Farm and Swartkrans members 1 and 2, South Africa. Palaeont. afr. 37, 115-126.

Elton, S., Bishop, L.C. & Wood, B. 2001. Comparative context of Plio-Pleistocene hominin brain evolution. JHE 41, 1-27.

Elton, S. & Morgan, B.J. 2006. Muzzle size, paranasal swelling size and body in Mandrillus leucophaeus. Primates 47, 151-157.

Elton, S. 2006. Forty years on and still going strong: the use of hominin-cercopithecid comparisons in paleoanthropology. J. Roy. Anthrop. Inst. (N.S.) 12, 19-38.

Enard, W., Khaitovich, P., Klose, J., Zöllner, S., Heissig, F., Giavalisco, P., Nieselt-Struwe, K., Muchmore, E., Varki, A., Ravid, R., Doxiadis, G.M., Bontrop, R.E. & Pääbo, S. 2002. Intra- and interspecific variation in primate gene expression patterns. Science 296, 340-343.

Enard, W., Przeworski, M., Fisher, S.E., Lai, C.S.L., Wiebe, V., Kitano, T., Monaco, A.P. & Pääbo, S. 2002. Molecular evolution of FOXP2, a gene involved in speech and language. Nature 418, 869-872.

Enstam, K.L. & Isbell, L.A. 2007. The guenons (Genus Cercopithecus) and their allies: Behavioral ecology of polyspecific associations. In, (Campbell, C.J., Fuentes, A., MacKinnon, K.C., Panger, M. & Bearder, S.K., Eds.) Primates in Perspective, pp. 252-273. UK: Oxford University Press.

438 Ereshefsky, M. 1992. Species, higher taxa, and the units of evolution. In, (Ereshefsky, M., Ed.) The units of evolution: Essays on the nature of species, pp.381-398. Cambridge, MA: MIT press.

Ereshefsky, M. 1997. The evolution of the Linnean hierarchy. Biology and Philosophy 12, 493-519.

Ereshefsky, M. 2001. The poverty of the Linnaean Hierarchy: a philosophical study of biological taxonomy. USA: Cambridge Uni. Press.

Eudey, A.A. 1980. Pleistocene glacial phenomena and the evolution of Asian macaques. In, (D.G. Lindburg, Ed.) The Macaques: Studies in Ecology, Behavior, and Evolution, pp. 52-83. New York: Van Nostrand Reinhold Company.

Ewing, B. & Green, P. 2000. Analysis of expressed sequence tags indicates 35,000 human genes. Nature Genetics 25, 232-234.

Fa, J.E. 1989. The genus Macaca: a review of taxonomy and evolution. Mammal Review 19, 45-81.

Falk, D. 1985. Hadar AL 162-28 endocast as evidence that brain enlargement preceded cortical reorganization in hominid evolution. Nature 313, 45-47.

Falk, D., Hildebolt, C., Smith, K., Morwood, M.J., Sutikna, T., Brown, P., Jatmiko, Saptomo, E.W., Brunsden, B. & Prior, F. 2005a. The brain of LB 1, Homo floresiensis. Science 308: 242-245.

Falk, D., Hildebolt, C., Smith, K. Morwood, M.J., Sutikna, T., Jatmiko, Saptomo, E.W., Brunsden, B. & Prior, F. 2005b. Response to Comment on "The Brain of LB1, Homo floresiensis". Science 310: 236.

Falk, D., Hildebolt, C., Smith, K., Morwood, M.J., Sutikna, T., Jatmiko, Saptomo, E.W., Brunsden, B. & Prior, F. 2006. Response to Comment on "The Brain of LB1, Homo floresiensis". Science 312: 999.

Fashing, P.J. 2007. African colobines monkeys: Patterns of between-group interaction. In, (Campbell, C.J., Fuentes, A., MacKinnon, K.C., Panger, M. & Bearder, S.K., Eds.) Primates in Perspective, pp. 201-223. UK: Oxford University Press.

Fedigan, L. & Fedigan, L.M. 1988. Cercopithecus aethiops: a review of field studies. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 389-411. UK: Cambridge Uni. Press.

Felsenstein, J. 1978. The Number of Evolutionary Trees. Systematic Zoology, 27, 27-33.

Fenart, R. & Deblock, R. 1973. Pan paniscus et Pan troglodytes craniometrie: Etude comparative et ontogenique selon les methodes classiques et vestibulaire. Musee Royal de L’Afrique Centrale – Tervuren, Belgique Annales – Serie IN-8 – Sciences Zoologiques – no 204.

Ferguson, J.M.H. 2002. On the use of genetic divergence for identifying species. Biol. J. Linn. Soc. 75, 509- 516.

Finarelli, J.A. & Clyde, W.C. 2004. Reassessing hominoid phylogeny: evaluating congruence in the morphological and temporal data. Paleobiology 30(4), 614-651.

Fleagle, J.G. 1977. Locomotor behavior and skeletal anatomy of sympatric Malaysian leaf-monkeys (Presbytis obscura and Presbytis melalophos). Yearbook of Physical Anthropology 20, 440-453.

Fleagle, J.G. 1984. Are there any fossil gibbons? In, (Preuschoft, H., Chiver, D.J., Brockelman, W.Y. & Creel, N., Eds.) The Lesser Apes: Evolutionary and behavioural biology, pp. 431-447. UK: Edinburgh University Press.

439 Fleagle, J.G. 1985. Size and adaptation in primates. In (Jungers, W.L., Ed.), Size and scaling in primate biology, pp. 1-19. Plenum Press: New York.

Fleagle, J.G. 1986. The fossil record of early catarrhine evolution. In, (Wood, B., Martin, L. & Andrews, P., Eds.) Major topics in primate and human evolution, pp. 130-149. Cambridge: Cambridge University Press.

Fleagle, J.G. 1998. Primate Adaptation and Evolution. 2nd ed. San Diego: Academic Press.

Fleagle, J.G. & Simons, E.L. 1978a. Micropithecus clarki, a small ape from the Miocene of Uganda. AJPA 49, 427-440.

Fleagle, J.G. & Simons, E.L. 1978b. Humeral morphology of the earliest apes. Nature 276, 705-707.

Fleagle, J.G., Stern Jr., J.T., Jungers, W., Susman, R., Vangor, A.K. & Wells, J.P. 1981. Climbing: A biomechanical link with brachiation and with bipedalism. Symp. Zool. Soc. Lond. 48, 359-375.

Fleagle, J.G. & Kay, R.F. 1985. The paleobiology of the catarrhines. In, (E. Delson, Ed.) Ancestors: the Hard Evidence, pp. 23-36. New York: Alan R. Liss, Inc.

Fleagle, J.G. & Meldrum, D.J. 1988. Locomotor behavior and skeletal morphology of two sympatric Pitheciine monkeys, Pithecia pithecia and Chiropotes satanas. AJP 16, 227-249.

Fleagle, J.G. &Reed, K.E. 1996. Comparing primate communities: a multivariate approach. JHE 30, 489-510.

Fleagle, J.G. & McGraw, W.S. 1999. Skeletal and dental morphology supports diphyletic origin of baboons and mandrills. PNAS 96, 1157-1161.

Fleagle, J.G. & McGraw, W.S. 2002. Skeletal and dental morphology of African papionins: unmasking a cryptic clade. JHE 42, 267-292.

Fleagle, J.G. & Reed, K.E. 2004. The evolution of primate ecology: patterns of geography and phylogeny. In, (Anapol, F., German, R.Z. & Jablonski, N.G., Eds) Shaping primate evolution: form, function and behavior, pp. 353-367. Cambridge: Cambridge University Press.

Foley, R. 1987. Another Unique Species: Patterns in human evolutionary ecology. UK: Longman Scientific & Technical.

Foley, R. 1991. How many species of hominid should there be? JHE 20, 413-427.

Foley, R. 1993. African terrestrial primates: The comparative evolutionary biology of Theropithecus and the Hominidae. In, (N. Jablonski, Ed.) Theropithecus: The Rise and Fall of a Primate Genus, pp. 245-270. Cambridge: Cambridge University Press.

Foley, R.A. 1994. Speciation, extinction and climatic change in hominid evolution. JHE 26, 275-289.

Foley, R.A. & Elton, S. 1998. Time and energy: The ecological context for the evolution of bipedalism. In, (Strasser, E., Fleagle, J., Rosenberger, A. & McHenry, H., Eds.) Primate locomotion: Recent advances, pp. 419-432. New York: Plenum Press.

Foley, R. 1999a. Evolutionary geography of Pliocene African hominids. In, (Bromage, T.G. & Schrenk, F., Eds.) African biogeography, climate change, & human evolution, pp. 328-348. New York: Oxford University Press.

Foley, R. 1999b. Hominid behavioral evolution: missing links in comparative primate socioecology. In, (Lee, P.C., Ed.) Comparative primate socioecology, pp. 363-388. UK: Cambridge University Press.

440 Foley, R. 2001. In the Shadow of the Modern Synthesis? Alternative Perspectives on the Last Fifty Years of Paleoanthropology. Evolutionary Anthropology 10, 5-14.

Fooden, J. 1976. Provisional classification and key to living species of macaques (Primates: Macaca). Folia primatol. 25, 225-236.

Fooden, J. 1982. Ecogeographic segregation of macaque species. Primates 23(4), 574-579.

Fooden, J. 1990. The bear macaque, Macaca arctoides: a systematic review. JHE 19, 607-686.

Fooden, J. & Albrecht, G.H. 1993. Latitudinal and insular variation of skull size in crab-eating macaques (Primates, Cercopithecidae: Macaca fascicularis). AJPA 92, 521-538.

Foote, M. 1997. Sampling, taxonomic description, and our evolving knowledge of morphological diversity. Paleobiology 23, 187-206.

Fox, E.A., van Schaik, C.P., Sitompul, A. & Wright, D. 2004. Intra-and Interpopulational differences in orangutan (Pongo pygmaeus) activity and diet: Implications for the invention of tool use. AJPA 125, pp. 162- 174.

Fragaszy, D.M. & Bard, K. 1997. Comparison of development and life history in Pan and Cebus. IJP 18, 683-701.

Franklin, D., Freedman, L. & Milne, N. 2005. Three-dimensional technology for linear morphological studies: a re-examination of cranial variation in four southern Africa indigenous populations. Homo 56, 17-34.

Freedman, L. 1957. The fossil Cercopithecoidea of South Africa. Ann. Of the Transvaal Museum 23, pp. 121- 262.

Frisch, J.E. 1973. The hylobatid dentition. Gibbon and Siamang, vol. 2, pp. 55-95. Basel: Karger.

Froehlich, J.W. & Supriatna, J. 1996. Secondary intergradation between Macaca maurus and M. tonkeana in South Sulawesi, and the species status of M. togeanus. In (Fa, J.E. & Lindburg, D.G., Eds.) Evolution and Ecology of Macaque Societies, pp. 43-70. UK: Cambridge Uni. Press.

Frost, S.R. 2001. New Early Pliocene Cercopithecidae (Mammalia: Primates) from Aramis, Middle Awash Valley, Ethiopia. American Museum Novitates 3350, 1-36.

Frost, S.R. & Delson, E. 2002. Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia. JHE 43, 687-748

Frost, S.R., Marcus, L.F., Bookstein, F.L., Reddy, D.P. & Delson, E. 2003. Cranial allometry, phylogeography, and systematics of large-bodied papionins (Primates: Cercopithecidae) inferred from geometric morphometric analysis of landmark data. The Anatomical Record Part A 275A, 1048-1072.

Frost, S.R. & Alemseged, Z. 2007. Middle Pleistocene fossil Cercopithecidae from Asbole, Afar Region, Ethiopia. JHE 53, 227-259.

Fujiyama, A., Watanabe, H., Toyoda, A., Taylor, T.D., Itoh, T., Tsai, S., Park, H., Yaspo, M., Lehrach, H., Chen, Z., Fu, G., Saitou, N., Osoegawa, K., de Jong, P.J., Suto, Y., Hattori, M. & Sakaki, Y. 2002. Construction and Analysis of a Human-Chimpanzee Comparative Clone Map. Science 295, 131-134.

Gabunia, L., Vekua, A., Lordkipanidze, D., Swisher, C.C.III, Ferring, R., Justus, A., Nioradze, M., Tvalchrelidze, M., Antón, S.C., Bosinski, G., Jöris, O., de Lumley, M-A. Majsuradze, G. & Mouskhelishvili, A. 2000. Earliest Pleistocene hominid cranial remains from Dmanisi, Republic of Georgia: taxonomy, geological setting, and age. Science 288, 1019-1025.

441

Gagneux, P., Wills, C., Gerloff, U., Tautz, D., Morin, P.A., Boesch, C., Fruth, B., Hohmann, G., Ryder, O.A. & Woodruff, D.S. 1999. Mitochondrial sequences show diverse evolutionary histories of African hominoids. PNAS 96, 5077-5082.

Gagneux, P. & Varki, A. 2001. Genetic differences between humans and great apes. MPE 18, 2-13.

Gagneux, P., Gonder, M.K., Goldberg, T.L. & Morin, P.A. 2001. Gene flow in wild chimpanzee populations: What genetic data tell us about chimpanzee movement over space and time. Proc. Trans. R. Soc. Lond. B 356, 889-897.

Galik, K., Senut, B., Pickford,, M., Gommery, D., Treil, J., Kuperavage, A.J. and Eckhardt, R.B. 2004. External and internal morphology of the BAR 1002’00 Orrorin tugenensis femur. Science 305: 1450-1453.

Galik, K., Eckhardt, R.B. & Kuperavage, A.J. 2005. Internal functional morphology of BAR 1002’00 documents ape-human divergence circa 6 Ma: That’s impossible, but anyway, everyone knew it all the time. AJPA 126 (Suppl. 40), 105.

Gannon, P.J., Holloway, R.L., Broadfield, D.C. & Braun, A.R. 1998. Asymmetry of chimpanzee planum temporale: Humanlike pattern of Wernicke’s brain language area homolog. Science 279, 220-222.

Garner, K.J. & Ryder, O.A. 1996. Mitochondrial DNA diversity in gorillas. MPE 6, 39-48.

Gautheir-Hion, A. 1973. Social and ecological features of talapoin monkey – comparisons with sympatric cercopithecines. In (Michael, R.P. & Crook, J.H., Eds.) Comparative ecology and behaviour in primates, pp. 147-170. London: Academic Press.

Gautheir-Hion, A.1988a. Polyspecific associations among forest guenons: ecological, behavioral and evolutionary aspects. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 452-476. UK: Cambridge Uni. Press.

Gautheir-Hion, A. 1988b. The diet and dietary habits of forest guenons. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 257-283. UK: Cambridge Uni. Press.

Gauthier, J-P., Drubbel, R.V. & Deleporte, P. 2002. Phylogeny of the Cercopithecus lhoesti group revisited: combining multiple character sets. In, (Glenn, M.E. & Cords, M., Eds.) The Guenons: Diversity and adaptation in African monkeys, pp. 37-48. New York: Kluwer Academic / Plenum Publishers.

Gebo, D.L. 1992. Plantigrady and foot adaptation in African Apes: Implications for hominid origins. AJPA 89, 29-58.

Gebo, D.L. 1996. Climbing, brachiation, and terrestrial quadrupedalism: historical precursors of hominid bipedalism. AJPA 101, 55-92.

Gebo, D.L. & Sargis, E.J. 1994. Terrestrial adaptations in the postcranial skeleton of guenon. AJPA 93, 341- 371.

Gebo, D.L. & Chapman, C.A. 1995. Positional behavior in five sympatric old world monkeys. AJPA 97, 49- 76.

Gebo, D.L. 2004. Palaeontology, terrestriality, and the intelligence of great apes. In (Russon, A.E. & Begun, D.R., Eds.) The Evolution of Thought: Evolutionary origins of Great Ape Intelligence, pp. 320-334. UK: Cambridge Uni. Press.

442 Gebo, D.L., MacLatchy, L., Kityo, R., Deino, A., Kingston, J. & Pilbeam, D. 1997. A hominoid genus from the Early Miocene of Uganda. Science 276, 401-404.

Gebo, D.L. & Chapman, C.A. 2000. Locomotor behavior in Ugandan monkeys. In, (Whitehead, P.F. & Jolly, C.J., Eds.) Old World Monkeys, pp. 480-495. UK: Cambridge Uni. Press.

Gee, H. 2000. Deep Time: Cladistics, The Revolution in Evolution. London: Fourth Estate.

Geissmann, T. 2002. Taxonomy and evolution of gibbons. EA 11(S1), 28-31.

Geissmann, T., Groves, C.P. & Roos, C. 2004. The Tenasserim Lutung, Trachypithecus barbei (Blyth, 1847) (Primates: Cercopithecidae): Description of a live specimen, and a reassessment of phylogenetic affinities, taxonomic history, and distribution. Contributions to Zoology 73(4). http://dpc.uba.uva.nl/ctz/vol73/nr04/art03

Geraads, D. 1987. Dating the Northern African cercopithecid fossil record. Human Evolution 2(1), 19-27.

Ghiglieri, M.P. 1987. Sociobiology of the great apes and the hominid ancestor. JHE 16, 319-357.

Gibbs, S., Collard, M. & Wood, B. 2002. Soft-tissue anatomy of the extant hominoids: a review and phylogenetic analysis. Journal of Anatomy 200, 3-49.

Gibert, J., Ribot, F., Gibert, L., Leakey, M., Arribas, A. & Martínez-Navarro, B. 1995. Presence of the cercopithecid genus Theropithecus in Cueva Victoria (Murcia, Spain). JHE 28, 487–493.

Gilbert, C.C. 2007. Craniomandibular morphology supporting the diphyletic origin of mangabeys and a new genus of the Cercocebus/Mandrillus clade, Procercocebus. JHE 53, 69-102.

Gilbert, C.C. & Rossie, J.B. 2007. Congruence of molecules and morphology using a narrow allometric approach. PNAS 104, 11910-11914.

Gingerich, P.D. 1979. Paleontology, phylogeny, and classification: An example from the mammalian fossil record. Systematic Zoology 28, 451-464.

Gittleman, J.L. & Luh, H-K. 1992. On comparing comparative methods. Annual Review of Ecology and Systematics 23, 383-404.

Glazko, G., Veeramachaneni, V., Nei, M. & Makaowski, W. 2005. Eighty percent of proteins are different between humans and chimpanzees. Gene 346, 215-219.

Glenn, M.E. & Cords, M., Eds. 2002. The Guenons: Diversity and adaptation in African monkeys. New York: Kluwer Academic/Plenum Publishers.

Godfrey, L. & Marks, J. 1991. The nature and origin of primate species. Yearbook of Physical Anthropology 34, 39-68.

Gomberg, D.N. 1985. Functional differences of three ligaments of the transverse tarsal joint in hominoids. JHE 14, 553-562.

Gonder, M.K., Oates, J.F., Disotell, T.R., Forstner, M.R.J., Morales, J.C. & Melnick, D.J. 1997. A new west African chimpanzee subspecies? Nature 388, 337-337.

Goodman, M. 1963a. Serological analysis of the phyletic relationships of recent hominoids. Human Biology 35(3), 377-406.

Goodman, M. 1963b. Man’s place in the phylogeny of the primates as reflected in serum proteins. In, (S. Washburn, Ed.) Classification and Human Evolution, pp. 204-224. Chicago: Aldine Publishing Co.

443

Goodman, M., Koop, B.F., Czelusiak, J., Fitch, D.H.A., Tagle, D.A. & Slightom, J.L. 1989. Molecular phylogeny of the family of apes and humans. Genome 31, 316-335.

Goodman, M., Tagle, D.A., Fitch, D.H.A., Bailey, W., Czelusniak, J., Koop, B.F., Benson, P. & Slightom, J.L. 1990. Primate evolution at the DNA level and a classification of hominoids. J. Mol. Evol. 30, 260-266.

Goodman, M., Porter, C.A., Czelusniak, J., Page, S.L., Schneider, H., Shoshani, J., Gunnell, G. & Groves, C.P. 1998. Toward a phylogenetic classification of primates based on DNA evidence complemented by fossil evidence. MPE 9, 585-598.

Goodman, M., Czelusniak, J., Page, S. & Meireles, C.M. 2001. Where DNA sequences place Homo sapiens in a phylogenetic classification of primates. In, (P.V. Tobias, M.A. Raath, J. Moggi-Cecchi & G.A. Doyle, Eds.) Humanity from African Naissance to Coming Millenia, pp. 279-289. Firenze University Press and Witwatersrand University Press.

Gould, S.J. 1975. Allometry in Primates, with emphasis on scaling and the evolution of the brain. Contrib. Primat. Vol. 5, pp. 244-292.

Gould, S.J. 1976. Ladders, bushes and human evolution. Natural History 85, 24-31.

Gould, S.J. 1985. A clock of evolution. Nat. Hist. 4, 12-25.

Goswani, A. 2006. The evolution of morphological integration in the mammalian skull. PhD thesis Uni. Of Chicago. USA: ProQuest information & Learning Company.

Gray, J.E. 1825. An outline of an attempt at the disposition of Mammalia into tribes and families, with a list of the genera apparently appertaining to each tribe. Ann. Of Phil. 10, 337-344.

Green, D.J., Gordon, A.D. & Richmond, B.G. 2007. Limb-size proportions in Australopithecus afarensis and Australopithecus africanus. JHE 52, 187-200.

Green, R.E., Krause, J., Ptak, S.E., Briggs, A.W., Ronan, M.T., Simons, J.F., Du, L., Egholm, M., Rothberg, J.M., Paunovic, M. & Pääbo, S. 2006. Analysis of one million base pairs of Neanderthal DNA. Nature 444, 330-336.

Greenwood, L.O. 1983. Toward the resolution of discrepancies between phenetic and paleontological data bearing on the question of human origins. In, (R.L. Ciochon & R.S. Corruccini, Eds.) New Interpretations of Ape and Human Ancestry, pp. 695-703. New York: Plenum Press.

Grehan, J.R. 2006. Mona Lisa Smile: the morphological enigma of human and great ape evolution. The Anatomical Record (Part B) 2898B, 139-157.

Grimwood, J., Gordon, L.A., Olsen, A., Terry, A., Schmutz, J., Lamerdin, J., Hellsten, U., Goodstein, D., Couronne, O., Tran-Gyamfi, M., et al. 2004. The DNA sequence and biology of human chromosome 19. Nature 428, 529 – 535.

Grine, F.E. 1981. Tropic differences between ‘Gracile’ and ‘Robust’ Australopithecines: A scanning electron microscope analysis of occlusal events. SAJS 77, 203-230.

Grine, F.E. 1986. Dental evidence for dietary differences in Australopithecus and Paranthropus: a quantitative analysis of permanent molar microwear. JHE 15, 783-822.

Grine, F.E. & Kay, R.F. 1988. Early hominid diets from quantitative image analysis of dental microwear. Nature 333, 765-768.

444 Grine, F.E. & Martin, L.B. 1988. Enamel thickness and development in Australopithecus and Paranthropus. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.3-42. New York: Aldine de Gruyter.

Grine, F.E., Jungers, W.L., & Schultz, J. 1996. Phenetic affinities among early Homo crania from East and South Africa. JHE 30, 189-225.

Grine, F.E., Ungar, P.S., Teaford, M.F. & El-Zaatari, S. 2006. Molar microwear in Praeanthropus afarensis: Evidence for dietary stasis through time and under diverse paleoecological conditions. JHE 51, 297-319.

Groves, C.P. & Mazak, V. 1975. An approach to the taxonomy of the Hominidae: Gracile Villafrachian hominids of Africa. Casopis pro mineralogii a geologii 20, 225-246.

Groves, C.P. 1970. The forgotten leaf-eaters and the phylogeny of the Colobinae. In, (Napier, J.R. & Napier, P.H., Eds.) Old World monkeys: evolution, systematics, and behavior, pp. 555-588. New York: Academic Press.

Groves, C. 1972. Systematics and phylogeny of the gibbons. Gibbon and Siamang, vol. 1, pp. 1-89. Basel: Karger.

Groves, C. 1978. Phylogenetic and population systematics of the mangabeys (Primates: Cercopithecoidea). Primates 19(1):1-34.

Groves, C. 1986. Systematics of the great apes. In, (J. Erwin & D.R. Swindler, Eds.) Comparative Primate Biology, Volume 1: Systematics, Evolution and Anatomy, pp. 187-217. New York: Alan R. Liss, Inc.

Groves, C.P. 1989. A Theory of Human and Primate Evolution. Oxford: Clarendon Press.

Groves, C.P. 1993. Speciation in living hominoid primates. In, (W.H. Kimbel & B.M. Lawrence, Eds.) Species, Species Concepts, and Primate Evolution, pp. 109-121. New York: Plenum Press.

Groves, C.P. 1999. Nomenclature of African Plio-Pleistocene hominins. JHE 37, 869-872.

Groves, C. 2000. The phylogeny of the Cercopithecoidea. In, (Whitehead, P.F. & Jolly, C.J., Eds.) Old World Monkeys, pp. 77-98. UK: Cambridge Uni. Press.

Groves, C. 2001a. Primate Taxonomy. Washington D.C.: Smithsonian Institute.

Groves, C.P. 2001b. Towards taxonomy of the Hominidae. In, (P.V. Tobias, M.A. Raath, J. Moggi-Cecchi & G.A. Doyle, Eds.) Humanity from African Naissance to Coming Millenia, pp. 307-318. Firenze University Press and Witwatersrand University Press.

Groves, C. 2001c. Time and taxonomy. Ludas Vitalis IX(15), 91-96.

Groves, C. 2003. A history of gorilla taxonomy. In, (Taylor, A. & Goldsmith, M.L., Eds.) Gorilla biology: a multidisciplinary perspective, pp. 15-34. UK: Cambridge Uni. Press.

Groves, C. 2004. The what, why and how of primate taxonomy. IJP 25(5), 1105-1126.

Grubb, P. 1990. Primate geography in the Afro-tropical biome. In (Peters, G., Ed.) Vertebrates in the Tropics, pp.187-214.

Grubb, P. 1999. Evolutionary processes implicit in distribution patterns on modern African mammals. In, (Bromage, T.G. & Schrenk, F., Eds.) African biogeography, climate change, & human evolution, pp. 150- 166. New York: Oxford University Press.

445 Grubb, P., Sandrock, O., Kullmer, O., Kaiser, T.M. & Schrenk, F. 1999. Relationships between Eastern and Southern African mammalian faunas. In, (Bromage, T.G. & Schrenk, F., Eds.) African biogeography, climate change, & human evolution, pp. 253-270. New York: Oxford University Press.

Grubb, P. Butynski, T.M., Oates, J.F., Bearder, S.K., Disotell, T.R., Groves, C.P. & Struhsaker, T.T. 2003. Assessment of the diversity of African primates. IJP 24, 1301-1357.

Gundling, T. Hill, A. 2000. Geological context of fossil Cercopithecoidea from eastern Africa. In, (Whitehead, P.F. & Jolly, C.J., Eds.) Old World Monkeys, pp. 180-213. UK: Cambridge Uni. Press.

Guy, F., Brunet, M., Schmittbuhl, M. & Viriot, L. 2003. New approaches in hominoid taxonomy: morphometrics. AJPA 121, 198-218.

Guy, F., Lieberman, D., Pilbeam, D., Ponce de León, M., Likius, A., Mackaye, H.T., Vignaud, P., Zollikofer, C. & Brunet, M. 2005. Morphological affinities of the Sahelanthropus tchadensis (Late Miocene hominid from Chad) cranium. PNAS 102, 18836-18841.

Haeusler, M. & McHenry, H.M. 2004. Body proportions of Homo habilis reviewed. JHE 46, 433-465.

Haeusler, M. & McHenry, H.M. 2007. Evolutionary reversals of limb proportions in early hominids? Evidence from KNM-ER 3735 (Homo habilis). JHE 1-23. doi:10.1016/j.jhevol.2007.06.001.

Haile-Selassie, Y., Suwa, G. & White, T.D. 2004. Late Miocene teeth from Middle Awash, Ethiopia, and early hominid dental evolution. Science 303, 1503-1505.

Haile-Selassie, Y. 2001. Late Miocene hominids from the Middle Awash, Ethiopia. Nature 412, 178-181.

Hall, K.R.L. 1968. Behavior and ecology of the wild patas monkey, Erythrocebus patas, in Uganda. In (Jay, P.C., Ed.) Primates: Studies in adaptation and variability, pp. 32-119.

Hall, L.M., Jones, D.S. & Wood, B.A. 1998. Evolution of the gibbon subgenera inferred from Cytochrome b DNA sequence data. MPE 10, 281-286.

Hamilton, A.C. 1988. Guenon evolution and forest history. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 13-34. UK: Cambridge Uni. Press.

Hamada, Y., Watanabe, T., Chatani, K. Hayakawa, S. & Iwamoto, M. 2005. Morphometrical comparison between Indian- and Chinese-derived rhesus macaques (Macaca mulatta). Anthro. Sci. 113, 183-188.

Hammer, O., Harper, D.A.T. & Ryan, P.D. Sept. 28, 2006. PAST – PAlaeontological STatistics, ver. 1.53; http://folk.uio.no/ohammer/past.

Hanihara, T. 1996. Comparison of craniofacial features of major human groups. AJPA 99, 389-412.

Hardt, T. & Henke, W. 2006. Facial patterns in Cercopithecoidea and Hominoidea: a geometric approach. Anthrop. Anz. 64, 129-146.

Harris, E.E., & Disotell, T.R., 1998. Nuclear gene trees and the phylogenetic relationships of the mangabeys (Primates: Papionini). Mol. Biol. Evol. 15(7), 892–900.

Harris, E.E. 2000. Molecular systematics of the Old World monkey tribe Papionini: analysis of the total available genetic sequences. JHE 38, 235-256.

Harrison, G.A. & Weiner, J.S. 1963. Some considerations in the formulation of theories of human evolution. In (S. Washburn, Ed.) Classification and Human Evolution, pp. 75-84. Chicago: Aldine Publishing Co.

446

Harrison, G.A., Tanner, J.M., Pilbeam, D. & Baker, P.T. 1995. Human Biology, 3rd ed. Oxford: Oxford University Press.

Harrison, T. 1987. The phylogenetic relationships of the early catarrhine primates: a review of the current evidence. JHE 16, 41-80.

Harrison, T. 1993. Cladistic concepts and the species problems in hominoid evolution. In, (W.H. Kimbel & B.M. Lawrence, Eds.) Species, Species Concepts, and Primate Evolution, pp. 345-371. New York: Plenum Press.

Harrison, T. 2005. The zoogeographic and phylogenetic relationships of early catarrhine primates in Asia. Anthro. Sci. 113, 43-51.

Harrison, T. & Harris, E.E. 1996. Plio-Pleistocene cercopithecids from Kanam East, western Kenya. JHE 30, 539-561.

Harrison, T. & Rook, L. 1997. Enigmatic anthropoid or misunderstood ape?: The phylogenetic status of Oreopithecus bambolii reconsidered. In, (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 327-362. New York: Plenum Press.

Hartman, S.E. 1988. A cladistic analysis of hominoid molars. JHE 17, 489-502.

Harvati, K. 2000. Dental eruption sequence among colobine primates. AJPA 112, 69-85.

Harvati, K. 2003. Quantitative analysis of Neanderthal temporal bone morphology using three-dimensional geometric morphometrics. AJPA 120, 323-338.

Harvati, K. & Weaver, T.D. 2006. Human cranial anatomy and the differential preservation of population history and climate signatures. The Anatomical Record Part A 288a, 1225-1233.

Harvati, K. & Frost, S.R. 2007. Dental eruption sequences in fossil colobines and the evolution of primate life histories. Int. J. Primatol.

Harvey, P.H. & Clutton-Bock, T.H. 1985. Life history variation in primates. Evolution 39, 559-581.

Hayashi, S., Hayasaka, K., Takenaka, O. & Horai, S. 1995. Molecular phylogeny of gibbons inferred from mtDNA sequences. J. Mol. Evol. 41, 359-365.

Heizmann, E.P.J. & Begun, D.R. 2001. The oldest Eurasian hominoid. JHE 41, 463-481.

Hellmann, I., Zoller, S., Enard, W., Ebersberger, I., Nickel, B. & Paabo, S. 2003. Selection on human genes as revealed by comparisons to chimpanzee DNA. Genome Research 13, 831-837.

Henneberg, M. & De Miquel, C. 2004. Hominins are a single lineage: brain and body size variability does not reflect postulated taxonomic diversity of hominins. Homo 55, 21-37.

Henneberg, M., Stephan, C.N. & Norris, R.M. 2005. Sources of biological variation. Is sex really important? AJPA 126 (Suppl.40), 114.

Hennig, W. 1966. Phylogenetic systematics. USA: University of Illinois Press.

Henzi, P. & Barrett, L. 2002. Evolutionary ecology, sexual conflict, and behavioral differentiation among baboon populations. EA 12, 217-230.

Henzi, P. & Barrett, L. 2005. The historical socioecology of savanna baboons. J. Zool. Lond. 265, 215-226.

447

Hershkovitz, I., Kornreich, L. & Laron, Z. 2007. Comparative skeletal features between Homo floresiensis and patients with primary growth hormone insensitivity (Laron syndrome). AJPA, DOI: 10.1002/ajpa.20655.

Hill, A. & Ward, S. 1988. Origin of the hominidae: The record of African large hominoid evolution between 14 and 4 My. Yearbook of Physical Anthropology 31, 49-83.

Hill, A., Ward, S., Deino, A., Curtis, G. & Drake, R. 1992. Earliest Homo. Nature 355, 719-722.

Hill, A. & Gundling, T. 2000. Geological context of the fossil Cercopithecoidea from eastern Africa. In, (Whitehead, P.F. & Jolly, C.J., Eds.) Old World Monkeys, pp. 180-213. UK: Cambridge Uni. Press.

Hill, A., Leakey,M., Kingston, J.D. & Ward, S. 2002. New cercopithecoids and a hominoid from 12·5 Ma in the Tugen Hills succession, Kenya. JHE 42, 75-93.

Hill, W.C.O. 1934. The monograph on the purple-faced leaf monkeys (Pithecus vetulus). Ceylon Journal of Science, Section B, Zoology 19, 23-88.

Hill, W.C.O. 1936. Supplementary observations on purple-faced leaf monkeys (Genus Kasi). Ceylon Journal of Science, Section B, Zoology 20, 115-133.

Hill, W.C.O. 1966. Primates: Comparative Anatomy and Taxonomy VI, Catarrhini, Cercopithecoidea, Cercopithecinae. Edinburgh: Edinburgh University Press.

Hill, W.C.O. 1968. The genera of old world apes and monkeys. In, (Chiarelli, B., Ed.) Taxonomy and Phylogeny of Old World Primates with References to the Origin of Man, pp. 7-15. Torino, Italy: Casa Editrice Libraria - Rosenberg & Sellier.

Hill, W.C.O. 1969a. The discovery of the chimpanzee. The Chimpanzee, vol. 1, pp. 1-21. Basel: Karger.

Hill, W.C.O. 1969b. The nomenclature, taxonomy and distribution of chimpanzees. The Chimpanzee, vol. 1, pp. 22-49. Basel: Karger.

Hill, W.C.O. 1970. Primates: Comparative Anatomy and Taxonomy, VIII Cynopithecinae, Papio, Mandrillus, Theropithecus. Edinburgh: Edinburgh University Press.

Hill, W.C.O. 1974. Primates: Comparative Anatomy and Taxonomy, 7 Cynopithecinae, Cercocebus, Macaca, Cynopithecus. Edinburgh: Edinburgh University Press.

Hillis, D.M. & Wiens, J.J. 2000. Molecules versus morphology in systematics: Conflicts, artifacts, and Misconceptions. In, (Weins, J.J., Ed.) Phylogenetic analysis of morphological data, pp. 1-19. Washington: Smithsonian Institution Press.

Hladik, C.M. 1977. Comparative studies of the strategies of two sympatric species of leaf-monkeys: Presbytis senex and Presbytis entellus. In (Clutton-Brock, T.H., Ed.) Primate Ecology: Studies of feeding and ranging behaviour in lemurs, monkeys and apes, pp. 324-354. London: Academic Press.

Hlusko, L.J. 2004. Integrating the genotype and phenotype in hominid paleontology. PNAS 101, 2653-2657.

Hlusko, L.J. 2006. A new large Pliocene colobine species (Mammalia: Primates) from Asa Issie, Ethiopia. Geobios 39, 57-69.

Hoelzer, G.A. & Melnick, D.J. 1996. Evolutionary relationships of the macaques. In (Fa, J.E. & Lindburg, D.G., Eds.) Evolution and Ecology of Macaque Societies, pp. 3-19. UK: Cambridge Uni. Press.

448 Hofreiter, M., Siedel, H., van Neer, W. & Vigilant, L. 2003. Mitochondrial DNA sequence from an enigmatic gorilla population (Gorilla gorilla uellensis). AJPA 121, 361-368.

Holliday, T.W. 2003. Species Concepts, reticulation, and human evolution. Current Anthropology 44, 653- 673.

Holloway, R.L. 1983. Cerebral brain endocast pattern of Australopithecus afarensis hominid. Nature 303, 420-422.

Holloway, R.L., Broadfield, D.C. & Yuan, M.S. 2003. Morphology and histology of chimpanzee primary visual striate cortex indicate that brain reorganization predated brain expansion in early hominid evolution. The Anatomical Record Part A 273, 594-602.

Hooijer, D.A. 1962. Quaternary langurs and macaques from the Malay archipelago. Zool. Verhandelingen 55, 1-64.

Horai, S., Satta, Y., Hayasaka, K., Kondo, R., Inoue, T., Ishida, T., Hayashi, S. & Takahata, N. 1992. Man’s place in Hominoidea revealed by mtDNA genealogy. J. Mol. Evol. 35, 32-43.

Horai, S., Hayasaka, K., Kondo, R., Tsugane, K. & Takahata, N. 1995. Recent African origin of modern humans revealed by complete sequences of hominoid mitochondrial DNAs. PNAS 92, 532-536.

Horn, A.D. 1987a. The socioecology of the Black mangabey (Cercocebus aterrimus) near Lake Tumba, Zaire. AJP 12, 165-180.

Horn, A.D. 1987b. Taxonomic assessment of the allopatric gray-cheeked mangabey (Cercocebus albigena) and Black mangabey (C. aterrimus): Comparative socioecological data and the species concept. AJP 12, 181- 187.

Horovitz, I., Zardoya & Meyer, A. 1998. Platyrrhine Systematics: A simultaneous analysis of molecular and morphological data. AJPA 106, 261-281.

Horovitz, I. 1999. A phylogenetic study of living and fossil Platyrrhines. Amer. Mus. Nov. 3269, 1-40.

Howells, W.W. 1950. Origin of the human stock: Concluding remarks of the chairman. Cold Springs Harbor Symposia on Quantitative Biology 15, 79-86.

Howells, W. 1973. Cranial Variation in Man: A study by multivariate analysis of patterns of difference among recent human populations. Cambridge, MA: Harvard University.

Howells, W. 1989. Skull Shapes and the Map: Craniometric analyses in the dispersion of modern Homo. Cambridge, MA: Harvard University.

Hughes, A.R. & Tobias, P.V. 1977. A fossil probably of the genus Homo from Sterkfontein, Transvaal. Nature 265, 310-312.

Hull, D. 1979. A craniometric study of the Black and White Colobus Illiger 1811 (Primates: Cercopithecidae). AJPA 51, 163-182.

Hunt, K.D. Ecological morphology of Australopithecus afarensis: Traveling terrestrially, eating arboreally. In, (Strasser, E., Fleagle, J., Rosenberger, A. & McHenry, H., Eds.) Primate locomotion: Recent advances, pp. 397-418. New York: Plenum Press.

Hunt, K.D. 2003. The single species hypothesis: Truly dead and pushing up bushes, or still twitching and ripe for resuscitation. Human Biology 75, 485-502.

449 Hutchins, M., Kleiman, D.G., Geist, V. & McDade, M.C. (Eds.) 2003. Primates. In Grzimek’s Animal Life Encyclopedia 2nd Ed. Vol. 14, Mammals III, pp. 1-253. MI: Gale Group.

Huxley, T.H. 1894. Man’s Place in Nature and other Anthropological Essays. London: MacMillan and Co.

Hylander, W.L. 1975. Incisor Size and Diet in Anthropoids with Special Reference to Cercopithecidae. Science 189, 1095-1098.

Hylander, W.L. 1988. Implications of In Vivo experiments for interpreting the functional significance of “Robust” Australopithecine jaws. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp. 55-83. New York: Aldine de Gruyter.

Hylander, W.L., Picq, P.G. & Johnson, K.R. 1991. Masticatory-stress hypotheses and the supraorbital region of primates. AJPA 86, 1-36.

Hylander, W.L. & Johnson, K.R. 1997. In Vivo bone strain patterns in the zygomatic arch of macaques and the significance of these patterns for functional interpretations of craniofacial form. AJPA 102, 203-232.

Inger, R.F. 1958. Comments on the definition of genera. Evolution 12, 370-384.

Ingold, T. 1995. ‘People like us’: The concept of anatomically modern humans. In, (R. Corbey & B. Theunissen, Eds.) Ape, Man, Apeman: Changing Views since 1600, pp. 241-262. Leiden University: Department of Prehistory.

International Commision on Zoological Nomenclature. 1999. International Commision on Zoological Nomenclature 4th Ed. UK: International Trust for Zoological Nomenclature.

International Human Genome Sequencing Consortium. 2001. Initial sequencing and analysis of the human genome. Nature 409, 860-921.

Isbell, L.A., Pruetz, J.D., Lewis, M. & Young, T.P. 1998. Locomotor activity differences between sympatric Patas monkeys (Erythrocebus patas) and Vervet monkeys (Cercopithecus aethiops): Implications for the evolution of long hindlimb length in Homo. AJPA 105, 199-207.

Isbell, L.A., Cheney, D.L. & Seyfarth, R.M. 2002. Why vervet monkeys (Cercopithecus aethiops) live in multimale groups. In, (Glenn, M.E. & Cords, M., Eds.) The Guenons: Diversity and adaptation in African monkeys, pp. 173-188. New York: Kluwer Academic / Plenum Publishers.

Ishida, H. & Pickford, M. 1997. A new Late Miocene hominoid from Kenya: Samburupithecus kiptalami gen. et sp. nov. Comptes Rendus de l'Académie des Sciences - Series IIA - Earth and Planetary Science, Volume 325, 823-829.

Iwamoto, T. 1993. The ecology of Theropithecus gelada. In (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 441-452. UK: Cambridge University Press.

Jablonski, N.G. & Yuerong, P. 1988. The evolution and paleobiogeography of monkeys in China. In (Whyte et al, Eds.) The palaeoenvironment of East Asia from the Mid-Tertiary vol. 2, pp. 849-867.

Jablonski, N.G. 1993a. The phylogeny of Theropithecus. In, (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 209-224. UK: Cambridge University Press.

Jablonski, N.G. 1993b. Evolution of the masticatory apparatus in Theropithecus. In, (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 299-329. UK: Cambridge University Press.

Jablonski, N.G. 1995. The Phyletic Position and Systematics of the Douc Langurs of Southeast Asia. AJP 35, 185-205.

450

Jablonski, N.G. 1998. The evolution of the Doucs and Snub-nosed monkeys and the question of the phyletic unity of the Odd-nosed colobines. In, (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub-Nosed Monkeys, pp.13-52. Singapore: World Scientific.

Jablonski, N.G. 2002. Fossil Old World monkeys: The Late Neogene radiation. In, (W.C. Hartwig, Ed.) The Primate Fossil Record, pp. 255-300. UK: Cambridge University Press.

Jablonski, N. & Yumin, G. 1991. Reassessment of Megamacaca lantianensis, a large monkey from the Pleistocene of north-central China. JHE 20(1), 51-66.

Jablonski, N.G. & Ruliang, P. 1995. Sexual dimorphism in the snub-nosed langurs (Colobinae: Rhinopithecus). AJPA 96, 251-272.

Jablonski, N.G. & Kelley, J. 1997. Did a major immunological event shape the evolutionary histories of apes and old world monkeys? JHE 33, 513-520.

Jablonski, N.G., Ruliang, P. & Chaplin, G. 1998. Mandibular morphology of the Doucs and Snub-Nosed Monkeys in relation to diet. In, (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub-Nosed Monkeys, pp.105-128. Singapore: World Scientific.

Jablonski, N. & Tyler, D.E. 1999. Trachypithecus auratus sangiranensis, A new fossil monkey from Sangiran, Central Java, Indonesia. IJP 20(3): 319-326.

Jablonski, N.G., Leakey, M.G., Kiarie, C. & Antón, M. 2002. A new skeleton of Theropithecus brumpti (Primates: Cercopithecidae) from Lomekwi, West Turkana, Kenya. JHE 43, 887-923

Jablonski, N.G. & Chaplin, G. 2004. Becoming bipedal: how do theories of bipedalism stand up to anatomical scrutiny? In, (Anapol, F., German, R.Z. & Jablonski, N.G., Eds) Shaping primate evolution: form, function and behavior, pp. 281-295. Cambridge: Cambridge University Press.

Jacob, T., Indriati, E., Soejono, R.P., Hsü, K., Frayer, D.W., Eckhardt, R.B., Kuperavage, A.J., Thorne, A. & Henneberg, M. 2006. Pygmoid Australomelanesian Homo sapiens skeletal remains from Liang Bua, Flores: Population affinities and pathological abnormalities. PNAS 103: 13421-13426.

James, W.W. 1960. The jaws and teeth of primates. London: Pitman Med. Pub. Co, Ltd.

Jenkins, P.D. 1990. Catalogue of the Primates in the British Museum (Natural History) and elsewhere in the British Isles Part V: The apes, Superfamily Hominoidea. London: Natural History Museum Publications.

Jensen-Seaman, M.I., Deinard, A.S. & Kidd, K.K. 2001. Modern African ape populations as genetic and demographic models of the last common ancestor of humans, chimpanzees and gorillas. Journal of Heredity 92, 475-480.

Jensen-Seaman, M.I., Deinard, A.S. & Kidd, K. 2003. Mitochondrial and nuclear DNA estimates of divergence between western and eastern gorillas. In, (Taylor, A. & Goldsmith, M.L., Eds.) Gorilla biology: a multidisciplinary perspective, pp. 247-268. UK: Cambridge Uni. Press.

Jerison, H.J. 1973. Evolution of the brain and intelligence. NY: Academic Press.

Jobling, M.A., Hurles, M.E. & Tyler-Smith, C. 2004. Human evolutionary genetics: Origins, peoples and diseases (chapter 7: Humans as apes). New York: Garland Science.

Johanson, D. & Edgar, B. 1996. From Lucy to Language. New York: Simon & Schuster Editions.

451 Johns, G.C. & Avise, J.C. 1998. A comparative summary of genetic distances in the vertebrates from mitochondrial Cytochrome b gene. Mol. Biol. Evol. 15(11), 1481-1490.

Johnson, S.C. 1981. Bonobos: Generalized hominid prototype or specialized insular dwarfs? Current Anthropology 22, 363-375.

Johnson, M.E., Viggiano, L., Bailey, J.A., Abdul-Rauf, M., Goodwin, G., Rocchi, M. & Eichler, E.E. 2001. Positive selection of a gene family during the emergence of humans and African apes. Nature 413, 514-519.

Jolicoeur, P. & Mosimann, J.E. 1960. Size and shape variation in the painted turtle, a principal component analysis. Growth 24, 339-354.

Jolly, C.J. 1966. Introduction to the Cercopithecoidea, with notes on their use as laboratory animals. Symp. Zool. Soc. Lond. 17, 427-457.

Jolly, C.J. 1970. The large African monkeys as an adaptive array. In, (Napier, J.R. & Napier, P.H., Eds.) Old World monkeys: evolution, systematics, and behavior, pp. 139-174. New York: Academic Press.

Jolly, C.J. 1993. Species, subspecies, and baboon systematics. In, (W.H. Kimbel & B.M. Lawrence, Eds.) Species, Species Concepts, and Primate Evolution, pp. 67-107. New York: Plenum Press.

Jolly, C.J. 2001. A proper study for mankind: Analogies from the Papionin monkeys and their implications for human evolution. Yearbook of Physical Anthropology 44, 177-204.

Jolly, C.J. 2003. Cranial anatomy and baboon diversity. The Anatomical Record Part A. 275A, 1043-1047.

Jolly, C.J. 2007. Baboons, mandrills, and mangabeys: Afro-papionin socioecology in a phylogenetic perspective. In, (Campbell, C.J., Fuentes, A., MacKinnon, K.C., Panger, M. & Bearder, S.K., Eds.) Primates in Perspective, pp. 240-251. UK: Oxford University Press.

Jolly, C.J., Woolley-Barker, T., Beyene, S., Distotell, T.R. & Phillips-Conroy, J.E. 1997. Intergeneric hybrid baboons. IJP 18, 597-627.

Jones, S., Martin, R. & Pilbeam, D., Eds. 1992. The Cambridge encyclopedia of human evolution. UK: Cambridge University Press.

Jones, N.B., Hawkes, K. & O’Connell, J.F. 1999. Some current ideas about the evolution of the human life history. In, (Lee, P.C., Ed.) Comparative primate socioecology, pp. 140-166. UK: Cambridge University Press.

Jones, T., Ehardt, C.L., Butynski, Davenport, T.R.B., Mpunga, N.E., Machaga, S.J. & De Luca, D.W. 2005. The highland mangabeys Lophocebus kipunji: A new species of African Monkey. Science 308, 1161-1164.

Jungers, W. 1988a. Relative joint size and hominoid locomotor adaptations with implications for the evolution of hominid bipedalism. JHE 17, 247-265.

Jungers, W. 1988b. New estimates of body size in Australopithecines. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.115-132. New York: Aldine de Gruyter.

Jungers, W.L., Falsetti, A.B. & Wall, C.E. 1995. Shape, relative size, and size-adjustments in morphometrics. Yearbook of Physical Anthropology 38, 137-161.

Jungers, W.L., Burr, D.B. & Cole, M.S. 1998. Body size and scaling of long bone geometry, bone strength, and positional behavior in Cercopithecoid primates. In, (Strasser, E., Fleagle, J., Rosenberger, A. & McHenry, H., Eds.) Primate locomotion: Recent advances, pp. 309-330. New York: Plenum Press.

452 Kaessmann, H., Wiebe, V. & Paabo, S. 1999. Extensive nuclear DNA sequence diversity among chimpanzees. Science 286, 1159-1162.

Kamilar, J.M. 2006. Geographic variation in savanna baboon (Papio) ecology and its taxonomic and evolutionary implications. In (Lehman, S.M. & Fleagle, J.G., Eds.) Primate biogeography: Progress and prospects, pp. 169-200. USA: Springer.

Kappleman, J. 1996. The evolution of body mass and relative brain size in fossil hominids. JHE 30, 243-276.

Kay, R.F. 1977. The evolution of molar occlusion in the cercopithecidae and early catarrhines. AJPA 46, 327- 352.

Kay, R.F. & Grine, F. 1988. Tooth morphology, wear and diet in Australopithecus and Paranthropus from South Africa. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.427-447. New York: Aldine de Gruyter.

Kay, R.F. & Ungar, P.S. 1997. Dental evidence for diet in some Miocene catarrhines with comments on the effects of phylogeny on the interpretation of adaptation. In, (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 131-151. New York: Plenum Press.

Kay, R.F., Ross, C. & Williams, B.A. 1997. Anthropoid origins. Science 275, 797-804.

Keller, M.P., Seifried, B.A. & Chance, P.E. 1999. Molecular evolution of the CMT1A-REP region: A human- and chimpanzee-specific repeat. Mol. Biol. Evol. 16(8), 1019-1026.

Kelley, J. 1997. Paleobiological and phylogenetic significance of life history in Miocene hominoids. In, (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 173-208. New York: Plenum Press.

Kelley, J. 2002. The hominoid radiation in Asia. In (W.C. Hartwig, Ed.) The Primate Fossil Record, pp. 369-384. UK: Cambridge University Press.

Kelly, J. 2004. Life history and cognitive evolution in the apes. In (Russon, A.E. & Begun, D.R., Eds.) The Evolution of Thought: Evolutionary origins of Great Ape Intelligence, pp. 280-297. UK: Cambridge Uni. Press.

Kennedy, G.E. 1992. The evolution of Homo sapiens as indicated by features of the postcranium. In, (Brauer, G. & Smith, F., Eds.) Continuity or Replacement: Controversies in Homo sapiens evolution, pp. 209-218. Rotterdam: A.A. Balkema.

Kennedy, G.E. 1999. Is “Homo rudolfensis” a valid species? JHE 36, 119-121.

Kidd, R.S. 2004. On the nature of morphology: selected canonical variates analyses of the hominoid hindtarsus and their interpretation. In, (Anapol, F., German, R.Z. & Jablonski, N.G., Eds) Shaping primate evolution: form, function and behavior, pp. 162-192. Cambridge: Cambridge University Press.

Kidder, J.H., Lantz, R.L. & Smith, F. 1992. Defining modern humans: a multivariate approach. In, (Brauer, G. & Smith, F., Eds.) Continuity or Replacement: Controversies in Homo sapiens evolution, pp. 157-177. Rotterdam: A.A. Balkema.

Kidder, J.M. & Durband, A.C. 2004. A re-evaluation of the metric diversity within Homo erectus. JHE 46, 299-315.

Kimbel, W.H., White, T.D. & Johanson, D. 1984. Cranial morphology of Australopithecus afarensis: A comparative study based on a composite reconstruction of the adult skull. AJPA 64, 337-388.

453

Kimbel, W.H., White, T.D. & Johanson, D.C. 1985. Craniodental morphology of the hominids from Hadar and Laetoli: Evidence of “Paranthropus” and Homo in the Mid Pliocene of Eastern Africa. In, (E. Delson, Ed.) Ancestors: the Hard Evidence, pp. 120-137. New York: Alan R. Liss, Inc.

Kimbel, W.H., White, T.D., Johanson, D.C., Yaroch, L.A., Skelton, R.R. & McHenry, H.M. 1986. On the Phylogenetic Analysis of Early Hominids. Current Anthropology 27, (4) 361-365.

Kimbel, W.H. & White, T.D. 1988. Variation, sexual dimorphism and the taxonomy of Australopithecus. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.175-192. New York: Aldine de Gruyter.

Kimbel, W.H., White, T.D. & Johanson, D.C. 1988. Implications of KNM-WT 17000 for the evolution of “Robust” Australopithecus. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.259-268. New York: Aldine de Gruyter.

Kimbel, W.H. & Martin, L.B. 1993. Species and speciation: Conceptual issues and their relevance for primate evolutionary biology. In, (W.H. Kimbel & B.M. Lawrence, Eds.) Species, Species Concepts, and Primate Evolution, pp. 539-553. New York: Plenum Press.

Kimbel, W.H. & Rak, Y. 1993. The importance of species taxa in paleoanthropology and an argument for the phylogenetic concept of the species category. In, (W.H. Kimbel & B.M. Lawrence, Eds.) Species, Species Concepts, and Primate Evolution, pp. 461-484. New York: Plenum Press.

Kimbel, W.H., Johanson, D. & Rak, Y. 1994. The first skull and other new discoveries of Australopithecus afarensis at Hadar, Ethiopia. Nature 368, 449-451.

Kimbel, W.H., Rak, Y. & Johanson, D.C. 2004. The skull of Australopithecus afarensis. UK: Oxford Uni. Press.

Kimbel, W.H., Walter, R.C., Johanson, D.C., Reed, K.E., Aronson, J.L., Assefa, Z., Marean, C.W., Eck, G.G., Bobe, R., Hovers, E. et al. 1996. Late Pliocene Homo and Oldowan Tools from the Hadar Formation (Kada Hadar Member), Ethiopia. JHE 31, 549-561.

Kimbel, W.H., Lockwood, C.A., Ward, C.V., Leakey, M.G., Rak, Y. & Johanson, D.C. 2006. Was Australopithecus anamensis ancestral to A. afarensis? A case of anagenesis in the hominin fossil record. JHE 51, 134-152.

Kimbel, W.H. 1995. Hominid speciation and Pliocene climatic change. In, (E.S. Vrba, G.H. Denton, T.C. Partridge & L.H. Burkle, Eds.) Paleoclimate and Evolution with Emphasis on Human Origins, pp. 425-437. New Haven: Yale University Press.

King, M. & Wilson, A.C. 1975. Evolution at two levels in humans and chimpanzees. Science 188, 107-116.

Kingdon, J. 1988. Comparative morphology of hands and feet in the genus Cercopithecus. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 184-193. UK: Cambridge Uni. Press.

Kirkpatrick, R.C. 1998. Ecology and Behavior in Snub-nosed and Douc Langurs. In, (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub-Nosed Monkeys, pp.155-190. Singapore: World Scientific.

Kirkpatrick, R.C. 2007. The Asian Colobines: Diversity among leaf-eating monkeys. In, (Campbell, C.J., Fuentes, A., MacKinnon, K.C., Panger, M. & Bearder, S.K., Eds.) Primates in Perspective, pp. 186-200. UK: Oxford University Press.

454 Klein, R.G. 1988. The causes of “Robust” australopithecine extinction. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp. 499-505. New York: Aldine de Gruyter.

Kluge, A.G. 1983. Cladistics and the classification of the great apes. In (R.L. Ciochon & R.S. Corruccini, Eds.) New Interpretations of Ape and Human Ancestry, pp. 151-177. New York: Plenum Press.

Knox, E.B. 1998. The use of hierarchies as organizational models in systematics. Biological Journal of the Linnean Society 63, 1-49.

Kohler, M., Moya-Sola, S. & Alba, D.M. 2000. Macaca (Primates, Cercopithecidae) from the Late Miocene of Spain. JHE 38, 447-452.

Kool, K.M. 1993. The diet and feeding behavior of he silver leaf monkey (Trachypithecus auratus sondaicus) in Indonesia. IJP 14, 667-700.

Koop, B.F., Goodman, M., Xu, P., Chan, K. & Slightom, J.L. 1986. Primate -globin DNA sequences and man’s place among the great apes. Nature 319, 234-238.

Kordos, L. & Begun, D. 2002. Rudabanya: A Late Miocene subtropical swamp deposit with evidence of the origin of the African Apes and Humans. Evolutionary Anthropology 11, 45-57.

Krentz, H.B. 1993. Postcranial anatomy of extant and extinct species of Theropithecus. In, (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 383-422. UK: Cambridge University Press.

Krings, M., Stone, A., Schmitz, R.W., Krainitzki, H., Stoneking, M. & Paabo, S. 1997. Neandertal DNA sequences and the origin of modern humans. Cell 90, 19-30.

Krogman, W.M. 1969. Growth changes in skull, face, jaws, and teeth of the chimpanzee. The Chimpanzee, vol. 1, 104-164. Basel: Karger.

Krovitz, G.E., Thompson, J.L. & Nelson, A.J. 2003. Hominid growth and development from australopithecine to Middle Pleistocene Homo. In, (J.L. Thompson, G.E. Krovitz & A.J. Nelson, Eds.) Patterns of Growth and Development in the Genus Homo, pp. 271-292. Cambridge: Cambridge University Press.

Kruskal, J.B. 1964. Multidimensional scaling by optimizing goodness of fit to a nonmetric hypothesis. Psychometrika 29, 1-27.

Kumar, S. 2005. Molecular clocks: four decades of evolution. Nature Reviews Genetics 6, 654-662.

Kumar, S., Filipski, A., Swarna, V., Walker, A. & Hedges, S.B. Placing confidence limits on the molecular age of the human-chimpanzee divergence. PNAS 102, 18842-18847.

Kuykendall, K. 2003. Reconstructing australopithecine growth and development: What do we think we know? In, (J.L. Thompson, G.E. Krovitz & A.J. Nelson, Eds.) Patterns of Growth and Development in the Genus Homo, pp. 191-218. Cambridge: Cambridge University Press.

Lahr, M.M. 1996. The evolution of modern human diversity: A study of cranial variation. UK: Cambridge University Press.

Lambert, J.E. 2002. Resource switching and species coexistence in guenons: a community analysis of dietary flexibility. In, (Glenn, M.E. & Cords, M., Eds.) The Guenons: Diversity and adaptation in African monkeys, pp. 309-324. New York: Kluwer Academic / Plenum Publishers.

Lambert, J.E. 2005. Competition, predation, and the evolutionary significance of the Cercopithecine cheek pouch: The case of Cercopithecus and Lophocebus. AJPA 126, 183-192.

455

Larson, S.G. 1998. Parallel evolution in the hominoid trunk and forelimb. Evolutionary Anthropology 6, 87- 99.

Le Floch-Prigent, P. & Deloison, Y. 1985. Anatomical comparison between entire feet of chimpanzee, gorilla and man. In, (P.V. Tobias, Ed.) Hominid Evolution: Past, Present and Future, pp. 135-141. New York: Alan R. Liss, Inc.

Le Gros Clark, W.E. 1947. Observations on the anatomy of the fossil Australopithecinae. Journal of Anatomy 81, 300-333.

Le Gros Clark, W.E. 1964. The Fossil Evidence for Human Evolution 2nd edition. Chicago: University of Chicago Press.

Leakey, L.S.B. 1966. Homo habilis, Homo erectus and the Australopithecines. Nature 209, 1279-1281.

Leakey, L.S.B. 1967. A early Miocene member of Hominidae. Nature 213, 155-163.

Leakey, L.S.B., Tobias, P.V. & Napier, J.R. 1964. A new species of the genus Homo from Olduvai Gorge. Nature 202, 7-9.

Leakey, M.D., Clarke, R.J. & Leakey, L.S.B. 1971. New hominid skull Bed I, Olduvai Gorge, Tanzania. Nature 232, 308-312.

Leakey, M.D., Hay, R.L., Curtis, G.H., Drake, R.E., Jackes, M.K. & White, T. 1976. Fossil hominids from the Laetolil Beds. Nature 262, 460-466.

Leakey, R.E.F. 1973a. Further evidence of Lower Pleistocene hominids from East Rudolf, North Kenya, 1972. Nature 242, 170-173.

Leakey, R.E.F. 1973b. Evidence for an advanced Plio-Pleistocene hominid from East Rudolf, Kenya. Nature 242, 447-450.

Leakey, R.E.F. 1974. Evidence for an advanced Plio-Pleistocene hominid from East Rudolf, Kenya. Nature 248, 653-656..

Leakey, R.E.F. 1976. New hominid fossils from the Koobi Fora formation in Northern Kenya. Nature 261, 574-576.

Leakey, R.E.F. & Wood, B.A. 1973. New evidence of the genus Homo from East Rudolf, Kenya. II. AJPA 39, 355-368.

Leakey, R.E.F. & Walker, A. 1976. Australopithecus and Homo erectus and the single species hypothesis. Nature 261, 572-574.

Leakey, R.E.F. & Walker, A. 1985. New higher primates from the early Miocene of Buluk, Kenya. Nature 318, 173-175.

Leakey, M.G. 1976. Cercopithecoidea of the East Rudolf Succession. In (Y. Coppens, F.C. Howell, G.Ll. Isaac & R.E.F. Leakey, eds.), Earliest Man and Environments in the Lake Rudolf Basin: Stratigraphy, Paleoecology, and Evolution, pp. 345-350. Chicago: University of Chicago Press.

Leakey, M.G. & Leakey, R.E.F. 1976. Further Cercopithecinae (Mammalia, Primates) from the Plio/Pleistocene of East Africa. In, (Savage, R.J.G., & Coryndon, S.C., Eds.) Fossil Vertebrates of Africa vol. 4, pp.121-146.

456 Leakey, M.G. & Leakey, R.E.F. 1986a. A new Miocene hominoid from Kenya. Nature 324, 143-146.

Leakey, M.G. & Leakey, R.E.F. 1986b. A second new Miocene hominoid from Kenya. Nature 324, 146-148.

Leakey, M.G. & Leakey, R.E.F. 1986c. A new Miocene small-bodied ape from Kenya. JHE 16, 369-387.

Leakey, M.G. 1982. Extinct large colobines from Plio-Pleistocene of Africa. AJPA 58, 153-172.

Leakey, M.G. 1988. Fossil evidence for the evolution of the guenons. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 7- 12. UK: Cambridge Uni. Press.

Leakey, M.G. 1993. Evolution of Theropithecus in the Turkana basin. In, (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 85-123. UK: Cambridge University Press.

Leakey, M.G., Fiebel, C.S., McDougall, I. & Walker, A. 1995. New four-million-year-old hominid species from Kanapoi and Allia Bay, Kenya. Nature 376, 565-571.

Leakey, M.G. & Walker, A. 1997. Afropithecus: function and phylogeny. In, (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 225-239. New York: Plenum Press.

Leakey, M.G., Feibel, C.S., McDougall, I., Ward, C. & Walker, A. 1998. New specimens and confirmation of an early age for Australopithecus anamensis. Nature 393, 62-66.

Leakey, M.G., Spoor, F., Brown, F.H., Gathogo, P.N., Kiarie, C., Leakey, L.N., McDougall, I. 2001. New hominin genus from eastern Africa shows diverse Middle Pliocene lineages. Nature 410, 433-440.

Leakey, M.G., Teaford, M.F. & Ward, C. 2003. Cercopithecidae from Lothagam. In, (Leakey, M.G. & Harris, J.M., Eds.) Lothagam: The Dawn of Humanity in Eastern Africa, pp. 201-257.

Lee, P.C. 1999. Comparative ecology of postnatal growth and weaning among haplorhine primates. In, (Lee, P.C., Ed.) Comparative primate socioecology, pp.111-139. UK: Cambridge University Press.

Lee-Thorp, J. & Sillen, A. 2001. Chemical signals in fossils offer new opportunities for assessing and comparing dietary niches of South African hominids. In (P.V. Tobias, M.A. Raath, J. Moggi-Cecchi & G.A. Doyle, Eds.) Humanity from African Naissance to Coming Millenia, pp. 321-325. Firenze University Press and Witwatersrand University Press.

Lee-Thorp, J., Sponheimer, M & van der Merwe, N.J. 2003. What do stable isotopes tell us about hominid dietary and ecological niches in the Pliocene? International Journal of Osteoarchaeology 13, 104-113.

Lee-Thorp, J. & Sponheimer, M. 2006. Contributions of biogeochemistry to understanding hominin dietary ecology. Yearbook of Physical Anthropology 49, 131-148.

Leigh, S.R., Relethford, J.H., Park, P.B. & Konigsberg, L.W. 2003. Morphological differentiation of Gorilla subspecies. In, (Taylor, A. & Goldsmith, M.L., Eds.) Gorilla biology: a multidisciplinary perspective, pp. 104-131. UK: Cambridge Uni. Press.

Leigh, S.R., Shah, N.F. & Buchanan, L.S. 2003. Ontogeny and phylogeny in papionin primates JHE, 45, 285-316

Lemos, B. Meiklejohn, C.D., Cáceres, M. & Hartl, D.L. 2005. Rates of divergence in gene expression profiles in gene expression profiles of primates, mice and flies: stabilizing selection and variability among functional categories. Evolution 59, 126–137.

457 Leonard, W.R. & Robertson, M.L. 1997. Comparative primate energetics and hominid evolution. AJPA 102, 265-281.

Leonard, W.R. Robertson, M.L., Snodgrass, J.J. & Kuzawa, C.W. 2003. Metabolic correlates of hominid brain evolution. Comparative Biochemistry and Physiology Part A 136, 5-15

Lernould, J-M. 1988. Classification and geographical distributions of guenons: a review. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 54-78. UK: Cambridge Uni. Press.

Lessertisseur, J., Nakache, J-P. & Petit-Maire, N. 1974. Application of principal component analysis using the Chi-square metric to the study of cranial characters in Primates. JHE 3, 19-35.

Li., M., Wei, F., Huang, C., Pan, R. & de Ruiter, J. 2004. Phylogeny of snub-nosed monkeys inferred from mitochondrial DNA, cytochrome B and 12S rRNA sequences. Int. J. Primatol. 25(4), 861-873.

Li, Qing-Qing & Zhang, Ya-Ping. 2005. Phylogenetic relationships of the macaques (Cercopithecidae: Macaca), inferred from mitochondrial DNA sequences. Biochemical Genetics 43 (7/8), 375-386.

Li, W. & Tanimura, M. 1987. The molecular clock runs more slowly in man than in apes and monkeys. Nature 326, 93-96.

Li, Y. 2007. Terrestriality and tree stratum use in a group of Sichuan snub-nosed monkeys. Primates 48, 197- 207.

Liang, Z., Ruliang, P. & Curnoe, D. 2006. Fossil baboon found in Yunnan, China. Zoological Research 27(6), 635-636.

Lieberman, D.E. 1995. Testing hypotheses about recent human evolution from skulls: Integrating morphology, function, development and phylogeny. Curr. Anthro. 36(2), 159-197.

Lieberman, D.E. Wood, B.A. & Pilbeam, D.R. 1996. Homoplasy and early Homo: an analysis of the evolutionary relationships of H. habilis sensu stricto and H. rudolfensis. JHE, 30, 1996, 97-120.

Lieberman, D.E. 2001. Another face in the family tree. Nature 410, 419-420.

Lieberman, D.E., McBratney, B.M. & Krovitz, G. 2002. The evolution and development of cranial form in Homo sapiens. PNAS 99, 1134-1139.

Lieberman, D.E., Krovitz, G. & McBratney, B.M. 2004. Testing hypotheses about tinkering in the fossil record: the case of the human skull. Journal of Experimental Zoology (Mol Dev Evol) 302B, 284-301.

Linneaus, C. 1758. Systema Naturae. Stockholm: Laurentii Salvii.

Lippold, L.K. 1998. Natural history of the Douc Langurs. In, (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub-Nosed Monkeys, pp.191-206. Singapore: World Scientific.

Lockwood, C.A. & Fleagle, J.G. 1999. The recognition and evaluation of homoplasy in primate and human evolution. Yearbook of Physical Anthropology 42, 189-232.

Lockwood, C.A. & Tobias, P.V. 1999. A large male hominin cranium from Sterkfontein, South Africa, and the status of Australopithecus africanus. JHE 36, 637-685.

Lockwood, C.A. & Tobias, C.A. 2002. Morphology and affinities of new hominin cranial remains from Member 4 of the Sterkfontein Formation, Gauteng Province, South Africa. JHE 42, 389-450.

458 Lockwood, C.A., Kimbel, W.H., & Lynch, J.M. 2004. Morphometrics and hominoid phylogeny: Support for a chimpanzee–human clade and differentiation among great ape subspecies. PNAS 101, 4356-4360.

Long, Y., Kirkpatrick, R.C., Xiao, L. & Zhong, T. 1998. Time budgets of the Yunnan snub-nosed monkey (Rhinopithecus [Rhinopithecus] bieti). In (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub- Nosed Monkeys, pp. 279-292. Singapore: World Scientific.

Lordkipanidze, D., Vekua, A., Ferring, R., Rightmire, G.P., Zollikofer, C.P.E., Ponce de León, M.S., Agusti, J., Kiladze, G., Mouskhelishvili, A., Nioradze, M. & Tappen, M. 2006. A fourth hominin skull from Dmanisi, Georgia. The Anatomical Record, Part A 288A, 1146-1157.

Lovejoy, C.O., Heiple, K.G. & Burstein, A.H. 1973. The gait of Australopithecus. AJPA 38, 757-780.

Lovejoy, C.O., Meindl, R.S., Ohman, J.C., Heiple, K.G. & White, T.D. 2002. The Maka femur and its bearing on the antiquity of human walking. AJPA 119, 97-133.

Lovejoy, C.O., McCollum, M.A., Reno, P.L. & Rosenman, B.A. 2003. Developmental biology and human evolution. Ann. Rev. Anthropology 32, 85-109.

Lucas, P.W., Corlett, R.T. & Luke, D.A. 1985. Plio-Pleistocene hominid diets: an approach combining masticatory and ecological analysis. JHE 14, 187-202.

Lucas, P.W. & Teaford, M.F. 1994. Functional morphology of colobine teeth. In, (Davies, A.G. & Oates, J.F., Eds.) Colobine monkeys: their ecology, behavior and evolution, pp. 173-204. Great Britain: Cambridge Uni. Press.

Macchiarelli, R., Bondioli, L., Debenath, A., Mazurier, A., Tournepiche, J.-F., Birch, W. & Dean, C. How Neanderthal molar teeth grew. 2006. Nature 444, 748-751.

MacDonald, D., Ed. 2001. The new encyclopedia of mammals. UK: Oxford University Press.

MacLaron, A. 1999. The comparative method: principles and illustrations from primate socioecology. In, (Lee, P.C., Ed.) Comparative primate socioecology, pp.5-22. UK: Cambridge University Press.

MacLatchy, L., Gebo, D., Kityo, R. & Pilbeam, D. 2000. Postcranial functional morphology of Morotopithecus bishopi, with implications for the evolution of modern ape locomotion. JHE 39, 159-183.

MacLatchy, L. 2004. The oldest ape. Evolutionary Anthropology 13, 90-103.

Mai, L.L. 1983. A model of chromosome evolution and its bearing on cladogenesis in the hominoidea. In, (R.L. Ciochon & R.S. Corruccini, Eds.) New Interpretations of Ape and Human Ancestry, pp. 87-114. New York: Plenum Press.

Maier, W. 1984. Tooth morphology and dietary specialization. In, (Chivers, D.J., Wood, B.A. & Bilsborough, A., Eds.) Food acquisition and processing in primates, pp. 303-330. New York: Plenum Press.

Mann, A. & Weiss, M. 1996. Hominoid phylogeny and taxonomy: A consideration of the molecular and fossil evidence in an historical perspective. MPE 5, 169-181.

Manaster, B.J. 1979. Locomotor adaptations within the Cercopithecus genus: a multivariate approach. AJPA 50, 169-182.

Manzi, G., Mallegni, F., & Ascenzi, A. 2001. A cranium for the earliest Europeans: Phylogenetic position of the hominid from Ceprano, Italy. PNAS 98, 10011-10016.

459 Manzi, G., Bruner, E. & Passarello, P. 2003. The one-million-year-old Homo cranium from Bouri (Ethiopia): a reconsideration of its H. erectus affinities. JHE 44, 731-736.

Marais, E. 1969. The soul of the ape. New York: Atheneum.

Marcus, L.F. 1969. Measurement of Selection Using Distance Statistics in the Prehistoric Orang-Utan Pongo pygmaeus palaeosumatrensis. Evolution 23(2), 301-307.

Marks, J. 2005. Phylogenetic trees and evolutionary trees. EA 14, 49-53.

Marroig, G. & Cheverud, J.M. 2001a. A comparison of phenotypic variation and covariation patterns and the role of phylogeny, ecology, and ontogeny during cranial evolution of new world monkeys. The American Naturalist 55(12), 2576-2600.

Marroig, G. & Cheverud, J.M. 2001b. A comparison of phenotypic variation and covariation patterns and the role of phylogeny, ecology and ontogeny during cranial evolution of new world monkeys. Evolution 55: 2576–2600.

Marshall, J. & Sugardjito, J. 1986. Gibbon systematics. In, (J. Erwin & D.R. Swindler, Eds.) Comparative Primate Biology, Volume 1: Systematics, Evolution and Anatomy, pp. 137-186. New York: Alan R. Liss, Inc.

Martin, L. 1986. Relationships among extant and extinct great apes and humans. In, (B. Wood, L. Martin & P. Andrews, Eds.) Major topics in primate and human evolution, pp. 161-187. Cambridge: Cambridge University Press.

Martin, R.D. 1981. Phylogenetic reconstruction versus classification: the case for clear demarcation. Biologist 28(3), 127-132.

Martin, R.D. 1984. Body size, brain size and feeding strategies. In, (Chivers, D.J., Wood, B.A. & Bilsborough, A., Eds.) Food acquisition and processing in primates, pp. 73-104. New York: Plenum Press.

Martin, R.D. 1990. Primate origins and evolution: A phylogenetic reconstruction. Princeton: Princeton University Press.

Martin, R.D. 1993. Allometric aspects of skull morphology in Theropithecus. In, (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 273-298. UK: Cambridge University Press.

Martin, R.D. & MacLarnon, M. 1988. Quantitative comparisons of the skull and teeth in guenons. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 160-183. UK: Cambridge Uni. Press.

Martin, R.D., MacLarnon, A.M., Phillips, J.L., Dussubieux, L., Williams, P.R. & Dobyns, W.B. 2006a. Comment on "The Brain of LB1, Homo floresiensis". Science 312: 999.

Martin, R.D., MacLarnon, A.M., Phillips, J.L., Dobyns, W.B. 2006b. Flores hominid: New species or microcephalic dwarf? The Anatomical Record Part A 288A (11), 1123-1145.

Mayr, E., Linsley, E.G. & Usinger, R.L. 1953. Methods and principles of systematic zoology. New York: McGraw Hill.

Mayr, E. 1950. Taxonomic categories in fossil hominids. Cold Springs Harbor Symposia on Quantitative Biology 15, 109-118.

Mayr, E. 1963. The taxonomic evaluation of fossil hominids. In, (S. Washburn, Ed.) Classification and Human Evolution, pp. 332-346. Chicago: Aldine Publishing Co.

460 Mayr, E. 1969. Principles of systematic zoology. New York: McGraw-Hill.

Mayr, E. 1974. Cladistic analysis or cladistic classification. Z. zool. Syst. Evolut.-forsch 12, 94-128.

Mayr, E. 1981. Biological classification: toward a synthesis of opposing methodologies. Science 214, 510- 516.

Mayr, E. 1982. The Growth of Biological Thought. Cambridge: Harvard University Press.

Mayr, E. 1992. Species concepts and their application. In, (Ereshefsky, M., Ed.) The units of evolution: Essays on the nature of species, pp.15-25. Cambridge, MA: MIT press.

Mayr, E. & Bock, W.J. 2002. Classifications and other ordering systems. J. Zool. Syst. Evol. Research 40, 169-194.

McBratney-Owen, B. & Lieberman, D.E. 2003. Postnatal ontogeny of facial position in Homo sapiens and Pan troglodytes. In, (J.L. Thompson, G.E. Krovitz & A.J. Nelson, Eds.) Patterns of Growth and Development in the Genus Homo, pp. 45-72. Cambridge: Cambridge University Press.

McBrearty, S. & Brooks, A.S. 2000. The revolution that wasn't: a new interpretation of the origin of modern human behavior. JHE 39, 453-563.

McBrearty, S. & Jablonski, N.G. 2005. First fossil chimpanzee. Nature 437, 105-108.

McCollum, M.A. 1999. The robust australopithecine face: A morphogenetic perspective. Science 284, 301- 305.

McCollum, M.A., Sherwood, C.C., Vinyard, C.J., Lovejoy, C.O. & Schachat, F. 2006. Of muscle-bound crania and human brain evolution: The story behind the MYH16 headlines. JHE 50, 232-236.

McConkey, E.H. 1997. The origin of human chromosome 18 from a human/ape ancestor. Cytogenet. Cell Genet 76, 189-191.

McCown, T.D. 1950. Classification of fossil men: The genus Palaeoanthropus and the problem of superspecific differentiation among the Hominidae. Cold Springs Harbor Symposia on Quantitative Biology 15, 87-96.

McCrossin, M.L. & Benefit, B.R. 1993. Recently recovered Kenyapithecus mandible and its implications for great ape and human origins. PNAS 90, 1962-1966.

McCrossin, M. & Benefit, B.R. 1997. On the relationship of Kenyapithecus, a large-bodied hominoid from the Middle Miocene of Eastern Africa. In, (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 241-267. New York: Plenum Press.

McCrossin, M.L., Benefit, B.R., Gitau, S.N., Palmer, A.K. & Blue, K.T. 1998. Fossil evidence for the origins of terrestriality among Old World higher primates. In, (Strasser, E., Fleagle, J., Rosenberger, A. & McHenry, H., Eds.) Primate locomotion: Recent advances, pp. 353-396. New York: Plenum Press.

McDougall, I., Brown, F.H. & Fleagle, J.G. 2005. Stratigraphic placement and age of modern humans from Kibish, Ethiopia. Nature 433, 733-736.

McGraw, W.S. 1994. Census, habitat preference, and polyspecific associations of the six monkeys in the Lomako Forest, Zaire. AJP 34, 295-307.

McGraw, W.S. 1998a. Comparative locomotion and habitat use of six monkeys in the Tai Forest, Ivory Coast. AJPA 105, 493-510.

461

McGraw, W.S. 1998b. Locomotion, support use, maintenance activities, and habitat structure: The case of the Tai forest Cercopithecids. In, (Strasser, E., Fleagle, J., Rosenberger, A. & McHenry, H., Eds.) Primate locomotion: Recent advances, pp. 79-94. New York: Plenum Press.

McGraw, W.S. 1998c. Posture and support use if Old World monkeys (Cercopithecidae): The influence of foraging strategies, activity patterns, and the spatial distribution of preferred food items. AJP 46, 229-250.

McGraw, W.S. 2002. Diversity of guenon positional behavior. In, (Glenn, M.E. & Cords, M., Eds.) The Guenons: Diversity and adaptation in African monkeys, pp. 113-132. New York: Kluwer Academic / Plenum Publishers.

McGraw, W.S. & Fleagle, J.G. 2006. Biogeography and evolution of the Cercocebus-Mandrillus clade: Evidence from the face. In (Lehman, S.M. & Fleagle, J.G., Eds.) Primate biogeography: Progress and prospects, pp. 201-224. USA: Springer.

McGrew, W.C., Baldwin, P.J. & Tutin, C.E.G. 1981. Chimpanzees in hot, dry and open habitat: Mt. Assirik, Senegal, West Africa. JHE 10, 227-244.

McHenry, H.M. 1978. Fore- and hindlimbs proportions in Plio-Pleistocene hominids. AJPA 49, 15-22.

McHenry, H.M. 1984a. Relative cheek-tooth size in Australopithecus. AJPA 64, 297-306.

McHenry, H.M. 1984b. The common ancestor: A study of the postcranium of Pan paniscus, Australopithecus, and other hominoids. In (Susman, R.L., Ed.) The pygmy chimpanzee: evolutionary biology and behavior, pp.201-230.

McHenry, H.M. 1985. Implications of postcanine megadontia for the origin of Homo. In, (E. Delson, Ed.) Ancestors: the Hard Evidence, pp. 178-183. New York: Alan R. Liss, Inc.

McHenry, H.M. 1988. New estimates of body weight in early hominids and their significance to encephalization and megadontia in “Robust” Australopithecines. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.133-148. New York: Aldine de Gruyter.

McHenry, H.M. 1994. Early hominid postcrania: Phylogeny and function. In (Ciochon, R.L. & Corruccini, R.S., Eds.) Integrative paths to the past: Paleoanthropological advances in honor of F. Clark Howell, pp. 251-268. USA: Prentice Hall.

McHenry, H.M. 2002. Introduction to the fossil record of human evolution. In, (W.C. Hartwig, Ed.) The Primate Fossil Record, pp. 401-406. UK: Cambridge University Press.

McHenry, H.M. & Corruccini, R.S. 1980. Late Tertiary hominoids and human origins. Nature 285, 397-398.

McHenry, H.M. & Corruccini, R.S. 1981. Pan paniscus and human evolution. AJPA 54, 355-367.

McHenry, H.M. & Skelton, R.R. 1985. Is Australopithecus africanus ancestral to Homo? In, (P.V. Tobias, Ed.) Hominid Evolution: Past, Present and Future, pp. 221-226. New York: Alan R. Liss, Inc.

McHenry, H.M. & Berger, L.R. 1998. Body proportions in Australopithecus afarensis and A. africanus and the origin of the genus Homo. JHE 51, 1-22.

McHenry, H.M. & Coffing, K. 2000. Australopithecus to Homo: Transformations in body and mind. Annu. Rev. Anthropol. 29, 125-146.

McHenry, H.M., Brown, C.C. & McHenry, L.J. 2007. Fossil hominin ulnae and the forelimb of Paranthropus. AJPA, DOI: 10.1002/ajpa.20656.

462

McKee, J.K. 1989. Australopithecine anterior pillars: Reassessment of the functional morphology and phylogenetic relevance. AJPA 80, 1-9.

McKee, J.K. 1991. Palaeo-ecology of the Sterkfontein Hominids: A Review and Synthesis. Palaeontologia Africana 28: 41-51.

McKee, J.K. 1993. Taxonomic and evolutionary affinities of Papio izodi fossils from Taung and Sterkfontein. Palaeontologia Africana 30, 43-49.

McKee, J.K. & A.W. Keyser. 1994. Cranio-dental Remains of Papio angusticeps from the Haasgat Cave Site, South Africa. IJP 15(6): 823-841.

McKee, J.K., Thackeray, J.F. & Berger, L. 1995. Faunal assemblage seriation of Southern African Pliocene and Pleistocene fossil deposits. AJPA 96, 235-250.

McKee, J.K. 1999. The autocatalytic nature of hominid evolution in the African Plio-Pleistocene Environments. In, (Bromage, T.G. & Schrenk, F., Eds.) African biogeography, climate change, & human evolution, pp. 57-67. New York: Oxford University Press.

McKenna, M.C. & Bell, S.K. 1997. Classification of Mammals above the Species Level. New York: Columbia University Press.

Medway, L. 1970. The monkeys of Sundaland: ecology and systematics of the cercopithecids of a humid equatorial environment. In, (Napier, J.R. & Napier, P.H., Eds.) Old World monkeys: evolution, systematics, and behavior, pp. 513-554. New York: Academic Press.

Meijaard, E. & Groves, C. 2004. The biogeographical evolution and phylogeny of the genus Presbytis. Primate Report 68, 71-90.

Mimmack, G.M., Mason, S.J. & Galpin, J.S. 2001. Choice of distance matrices in cluster analysis: defining regions. Journal of Climate 14, 2790-2797.

Ming, L., Fuwen, W., Chengming, H., Pan, R. & de Ruiter, J. 2004. Phylogeny of snub-nosed monkeys inferred from mitochondrial DNA, Cytochrome b and 12S rRNA sequences. IJP 25(4), 861-873.

Mitani, M. 1991. Niche overlap and polyspecific associations among sympatric cercopithecids in the Campo Animal Reserve, Southwestern Cameroon. Primates 32, 137-151.

Mitani, J.C. & Watts, D. P. 1999. Demographic influences on the hunting behavior of chimpanzees. AJPA 109, 439-454.

Miller, J.M.A., Albrecht, G.H. & Gelvin, B.R. 2004. Craniometric variation in early Homo compared to modern gorillas: a population-thinking approach. In, (Anapol, F., German, R.Z. & Jablonski, N.G., Eds) Shaping primate evolution: form, function and behavior, pp. 66-96. Cambridge: Cambridge University Press.

Mitteroecker, P. Gunz, P., Bernhard, M., Schaefer, K. & Bookstein, F.L. 2004. Comparison of cranial ontogenetic trajectories among great apes and humans. JHE 46, 679-698.

Monda, K., Simmons, R.E., Kressirer, P., Su, B. & Woodruff, D.S. 2007. Mitochondrial DNA hypervariable region-1 sequence variation and phylogeny of the concolor gibbons, Nomascus. AJP 69, 1-22.

Moore, C.M., Janish, C., Eddy, C.A., Hubbard, G.B., Leland, M.M. & Rogers, J. 1999. Cytogenetic and fertility studies of a Rheboon, Rhesus macaque (Macaca mulatta) x Baboon (Papio hamadryas) cross: Further support for a single karyotype nomenclature. AJPA 110, 119-127.

463 Mootnick, A. & Groves, C. 2005. A new generic name for the Hoolock gibbon (Hylobatidae). IJP 26(4), 971- 976.

Morales, J.C. & Melnick, D.J. 1998. Phylogenetic relationships of the macaques (Cercopithecidae: Macaca), as revealed by high resolution restriction site mapping of mitochondrial ribosomal genes. JHE 34, 1-23.

Morin, P.A., Moore, J.J., Chakraborty, R., Jin, L., Goodall, J. & Woodruff, D.S. 1994. Kin selection, social structure, gene flow, and the evolution of the chimpanzees. Science 265, 1193-1201.

Morwood, M.J., Brown, P., Jatmiko, Sutikna, T., Saptomo, E.W., Westaway, K.E., Due, R.A., Roberts, R.G., Maeda, T., Wasisto, S. & Djubiantono, T. 2005. Further evidence for small-bodied hominins from the Late Pleistocene of Flores, Indonesia. Nature 437, 1012-1017.

Mosimann, J.E. 1970. Size allometry: Size and shape variables with characterizations of the Lognormal and generalized gamma distributions. Journal of the American Statistical Association 65(330), 930-945.

Mosimann, J.E. & James, F.C. 1979. New statistical methods for allometry with application to Florida Red- Winged blackbirds. Evolution 33, 444-459.

Moya-Sola, S. & Kohler, M. 1993. Recent discoveries of Dryopithecus shed new light on evolution of great apes. Nature 365, 543-545.

Moya-Sola, S. & Kohler, M. 1996. A Dryopithecus skeleton and the origins of great-ape locomotion. Nature 379, 156-159.

Moyà-Solà, S., Köhler, M. & Alba, D.M. 2001. Egarapithecus narcisoi, a new genus of Pliopithecidae (primates, catarrhini) from the Late Miocene of Spain. AJPA, 114, 312-324.

Moya-Sola, S., Kohler, M., Alba, D.M., Casanovas-Vilar, I. & Galindo, J. 2004. Pierolapithecus catalaunicus, a new Middle Miocene Great Ape from Spain. Science 306, 1339-1344.

Mulder, M.B. 2001. Using phyogenetically based comparative methods in anthropology: More questions than answers. Evolutionary Anthropology 10, 99-111.

Myers, R.H. & Shafer, D.A. 1979. Hybrid ape offspring of a mating of gibbon and siamang. Science 205, 308-310.

Nadal-Roberts, M. & Collard, M. 2005. Impact of methodological choices on assessments of the reliability of fossil primate phylogenetic hypotheses. Folia Primatol. 76, 207-221.

Nadezhdin, E.V. & Sverdlov, E.D. 2005. How different is the human genome from the genomes of the Great Apes? In, (Sverdlov, E.D., Ed.) Retroviruses and Primate Genome Evolution, pp. 68-92.USA: Landes Bioscience.

Nakatsukasa, M. 1994. Morphology of the humerus and femur in African mangabeys and geunons: Functional adaptation and implications for the evolution of positional behavior. African Study Monographs, Suppl. 21, 1- 61.

Nakatsukasa, M. 1996. Locomotor differentiation and different skeletal morphologies in Mangabeys (Lophocebus and Cercocebus). Folia Primatol. 66, 15-24.

Nakatsukasa, M., Pickford, M., Egi, N. & Senut, B. 2007. Femur length, body mass, and stature estimates of Orrorin tugenensis, a 6 Ma hominid from Kenya. Primates 48, 171-178.

Napier, J.R. & Napier, P.H. 1967. A handbook of living primates: morphology, ecology and behaviour of nonhuman primates. London: Academic Press.

464

Napier, J.R. & Napier, P.H., Eds. 1970. Old World monkeys: evolution, systematics, and behavior. New York: Academic Press.

Napier, J.R. 1970. Paleoecology and catarrhine evolution. In, (Napier, J.R. & Napier, P.H., Eds.) Old World monkeys: evolution, systematics, and behavior, pp. 53-96. New York: Academic Press.

Napier, P.H. 1981. Catalogue of Primates in the British Museum (Natural History) and elsewhere in the British Isles, Part II: Family Cercopithecidae, Subfamily Cercopithecinae. London: Natural History Museum Publications.

Napier, P.H. 1985. Catalogue of Primates in the British Museum (Natural History) and elsewhere in the British Isles, Part III: Family Cercopithecidae, Subfamily Colobinae. London: Natural History Museum Publications.

Navarro, A. & Barton, N.H. 2003. Chromosomal speciation and molecular divergence-accelerated evolution in rearranged chromosomes. Science 300, 321-324.

Newman, T.K., Jolly, C.J. & Rogers, J. 2004. Mitochondrial phylogeny and systematics of baboons (Papio). AJPA 124, 17-27.

Newton, P.N. & Dunbar, R.I.M. 1994. Colobine monkey society. In, (Davies, A.G. & Oates, J.F., Eds.) Colobine monkeys: their ecology, behavior and evolution, pp. 311-346. Great Britain: Cambridge Uni. Press.

Nijman, V. 2000. Geographic distribution of ebony leaf monkey Trachypithecus auratus (E. Geoffroy Saint- Hilaire, 1812) (Mammalia: Primates: Cercopithecidae). Contributions to Zoology 69(3). http://dpc.uba.uva.nl/ctz/vol69/nr03/art02

Nimchinsky, E.A., Gilissen, E., Allman, J.M., Perl, D.P., Erwin, J.M. & Hof, P.R. 1999. A neuronal morphologic type unique to humans and great apes. PNAS 96, 5268-5273.

Noonan, J.P., Coop, G., Kudaravalli, S., Smith, D., Krause, J., Alessi, J., Chen, F., Platt, D., Paabo, S., Pritchard, J.K. & Rubin, E.M. 2006. Sequencing and analysis of Neanderthal genomic DNA. Science 314, 1113-1118.

Nowak, R.M. 1999. Walker’s Mammals of the world, 6th edition, vol. 1. Baltimore: Johns Hopkins University Press.

O’Higgins, P. & Pan, R.L. 2004. Advances in the analysis of form and pattern: facial growth in African colobines. In, (Anapol, F., German, R.Z. & Jablonski, N.G., Eds) Shaping primate evolution: form, function and behavior, pp. 24-44. Cambridge: Cambridge University Press.

Oates, J.F. 1977. The guereza and its food. In (Clutton-Brock, T.H., Ed.) Primate Ecology: Studies of feeding and ranging behaviour in lemurs, monkeys and apes, pp. 276-323. London: Academic Press.

Oates, J.F. 1988a. The distribution of Cercopithecus monkeys in West Africa forests. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 79-103. UK: Cambridge Uni. Press.

Oates, J.F. 1988b. The diet of the Olive colobus monkey, Procolobus verus, in Sierra Leone. IJP 9, 457-478.

Oates, J.F. & Davies, A.G. 1994. What are the colobines? In, (Davies, A.G. & Oates, J.F., Eds.) Colobine monkeys: their ecology, behavior and evolution, pp. 1-10. Great Britain: Cambridge Uni. Press.

465 Oates, J.F., Davies, G. & Delson, E. 1994. The diversity of the living colobines. In, (Davies, A.G. & Oates, J.F., Eds.) Colobine monkeys: their ecology, behavior and evolution, pp. 45-74. Great Britain: Cambridge Uni. Press.

Oates, J.F. 1994. The natural history of African colobines. In, (Davies, A.G. & Oates, J.F., Eds.) Colobine monkeys: their ecology, behavior and evolution, pp. 75-128. Great Britain: Cambridge Uni. Press.

Ohman, J.C., Lovejoy, C.O., White, T.D.; Eckhardt, R.B., Galik, K. & Kuperavage, A.J. 2005. Questions about Orrorin Femur. Science 307: 845.

Olson, E.C. & Miller, R.L. 1958. Morphological Integration. USA: Uni. of Chicago Press.

Olson, M.V. & Varki, A. 2003. Sequencing the chimpanzee genome: Insights into human evolution and disease. Nature Genetics 4, 20-28.

Olson, T.R. 1981. Basicranial morphology of the extant hominoids: the new material from the Hadar formation, Ethiopia, and its significance in early human evolution and taxonomy. In, (Stringer, C. Ed.) Aspects of Human Evolution. Symp. Soc. For the study of human biology Vol. XXI, pp. 99-128. London: Taylor & Francis Ltd.

Olson, T.R. 1985. Cranial morphology and systematics of the Hadar formation hominids and “Australopithecus” africanus. In, (Delson, E., Ed.) Ancestors: The hard evidence, pp. 102-119. NY: Alan R. Liss, Inc.

Omland, K.E. 1997. Correlated rates of molecular and morphological evolution. Evolution 51, 1381-1393.

Oxnard, C.E. 1975a. The place of the australopithecines in human evolution: grounds for doubt? Nature 258, 389-395.

Oxnard, C.E. 1975b. Uniqueness and diversity in human evolution: morphometric studies of Australopithecines. Chicago: The University of Chicago Press.

Oxnard, C.E. 1983. The Order of Man: A biomathematical anatomy of the primates. Hong Kong University Press.

Oxnard, C.E. 1998. The information content of morphometric data in primates: Function, development, and evolution. In, (Strasser, E., Fleagle, J., Rosenberger, A. & McHenry, H., Eds.) Primate locomotion: Recent advances, pp. 255-275. New York: Plenum Press.

Oxnard, C.E. 2004. Brain evolution: mammals, primates, chimpanzees, and humans. IJP 25(5), 1127-1158.

Page, S.L., Chiu, C. & Goodman, M., 1999. Molecular phylogeny of Old World monkeys (Cercopithecidae) as inferred from gamma-globin DNA sequences. Mol. Phylogenet. Evol. 13(2), 348–359.

Page, S.L. & Goodman, M. 2001. Catarrhine phylogeny: Noncoding DNA evidence for a diphyletic origin of the mangabeys and for a human-chimpanzee clade. MPE 18, 14-25.

Pan, Y.-R. & Jablonski, N.G. 1987. The age and geographical distribution of fossil Cercopithecids in China. Human Evolution 2(1), 59-69.

Pan, R. & P. Yanzhang. 1995. Body size of Macaca anderssoni and its offspring. Human Evolution 10(4), 283-287.

Pan, R., & Oxnard, C. 2001. Cranial morphology of the Golden monkey (Rhinopithecus) and Douc Langur (Pygathrix nemaeus). Human Evolution 16, 199-223.

466 Pan, R. & Wei, F. & Li, M. 2003. Craniofacial variation of the Chinese macaques explored with morphologika. Journal of Morphology 256, 342-348.

Pan, R.L. & Oxnard, C.E. 2003. Dental variation among Asian colobines (Nonhuman Primates): Phylogenetic similarities or functional correspondence? Zoological Studies 42, 93-105.

Pan, R.L. & Oxnard, C.E. 2004. Craniodental variation in the African macaque, with reference to various Asian species. Folia Primatologica 75(6), 355-375.

Pan, R.L. & Groves, C. 2004. Cranial variation among the Asian colobines. In, (Anapol, F., German, R.Z. & Jablonski, N.G., Eds.) Shaping primate evolution: form, function and behavior, pp. 45-65. Cambridge: Cambridge University Press.

Pan, R., Groves, C. & Oxnard, C. 2004. Relationships between the fossil colobine Mesopithecus pentelicus and extant cercopithecoids, based on dental metrics. AJP 62, 287-299.

Pan, R. 2006. Dental morphometric variation between African and Asian colobines, with special reference to the other Old World Monkeys. Journal of Morphology 267, 1087-1098.

Patterson, N., Richter, D.J. Gnerre, S., Lander, E.S. & Reich, D. 2006. Genetic evidence for complex speciation of humans and chimpanzees. Nature 441, 1103-1108.

Penin, X., Berge, C. & Baylac, M. 2002. Ontogenetic study of the skull in modern humans and the common chimpanzee: Neotenic hypothesis reconsidered with a tridimensional Procustes analysis. AJPA 118, 50-62.

Peters, C.R. & O’Brien, E.M. 1981. The early hominid plant-food niche: Insights from an analysis of plant exploitation by Homo, Pan, and Papio in Eastern and Southern Africa. Current Anthropology 22, 127-140.

Pickford, M. 1987. The chronology of the Cercopithecoidea of East Africa. Human Evolution 2(1), 1-17.

Pickford, M. 1993. Climatic change, biogeography, and Theropithecus. In, (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 227-243. UK: Cambridge University Press.

Pickford, M. & Senut, B. 1988. Habitat and locomotion in Miocene cercopithecoids. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 35-53. UK: Cambridge Uni. Press.

Pickford, M. & Ishida, H. 1998. Interpretation of Samburupithecus, an upper Miocene hominoid from Kenya. Comptes Rendus de l'Académie des Sciences - Series IIA - Earth and Planetary Science 326, 299-306.

Pickford, M. & Senut, B. 2001. The geological and faunal context of Late Miocene hominid remains from Lukeino, Kenya. Comptes Rendus de l’Académie des Sciences - Series IIA - Earth and Planetary Science, 332, 145-152.

Pickford, M., Senut, B., Gommery, D. & Treil, J. 2002. Bipedalism in Orrorin tugenensis revealed by its femora. C.R. Palevol. 1, 191-203.

Pickering, T.R. 2006. Subsistence behaviour of South African Pleistocene hominids. SAJS 102, 205-210.

Pilbeam, D. 1962. The earliest hominids. Nature 219, 1335-1338.

Pilbeam, D. 1982. New hominoid skull material from the Miocene of Pakistan. Nature 295, 232-234.

Pilbeam, D. 1996. Genetic and morphological records of the hominoidea and hominid origins: A synthesis. MPE 5, 155-168.

467 Pilbeam, D. 2002. Perspectives on the Miocene Hominoidea. In, (W.C. Hartwig, Ed.) The Primate Fossil Record, pp. 303-310. UK: Cambridge University Press.

Pilbeam, D., Rose, M.D., Barry, J.C. & Shah, S.M.I. 1990. New Sivapithecus humeri from Pakistan and the relationship of Sivapithecus and Pongo. Nature 348, 237-239.

Pilbeam, D. & Young, N. 2004. Hominoid evolution: Synthesizing disparate data. C.R. Palevol 3, 303-319.

Plavcan, J.M. 1993. Catarrhine dental variability and species recognition in the fossil record. In, (W.H. Kimbel & B.M. Lawrence, Eds.) Species, Species Concepts, and Primate Evolution, pp. 239-263. New York: Plenum Press.

Plavcan, J.M. 1999. Mating systems, intrasexual competition and sexual dimorphism in primates. In, (Lee, P.C., Ed.) Comparative primate socioecology, pp. 241-270. UK: Cambridge University Press.

Plavcan, J.M. 2002. Taxonomic variation in the patterns of craniofacial dimorphism in primates. JHE 42, 579-608.

Plavcan, J.M. 2003. Scaling Relationships between Craniofacial Sexual Dimorphism and Body Mass Dimorphism in Primates: Implications for the Fossil Record. AJPA 120, 38-60.

Plavcan, J.M. & Cope, D.A. 2002. Metric variation and species recognition in the fossil record. Evolutionary Anthropology 10, 204-222.

Plavcan, J.M., Lockwood, C.A., Kimbel, W.H., Lague, M.R. & Harmon, E.H. 2005. Sexual dimorphism in Australopithecus afarensis revisited: How strong is the case for a human-like pattern of dimorphism? JHE 48, 313-320.

Polanski, J.M. & Franciscus, R.G. Patterns of craniofacial integration in extant Homo, Pan, and Gorilla. AJPA 131, 38-49.

Poux, C. & Douzery, E.J.P. 2004. Primate phylogeny, evolutionary rate variations, and divergence times: A contribution from the nuclear gene IRBP. AJPA 124, 1-16.

Prat, S., Brugal, J.-P., Tiercelin, J.-J., Barrat, J.-A., Bohn, M., Delagnes, A., Harmand, S., Kimeu, K., Kibunjia, M., Texier, P.-J. 2005. First occurrence of early Homo in the Nachukui Formation (West Turkana, Kenya) at 2.3-2.4 Myr. JHE 49, 230-240.

Prueschoft, H. 1969. The mechanical basis of the morphological differences in the skeletons of apes and man. Proc. 2nd int. Congr. Primat., Atlanta, GA 1968 vol. 2, pp. 160-170. Basel: Karger.

Prueschoft, H. 1970. Functional anatomy of the lower extremity. The Chimpanzee vol. 3, pp. 221-294. Basel: Karger.

Prueschoft, H. 1973. Functional anatomy of the upper extremity. The Chimpanzee vol. 6, pp. 34-120. Basel: Karger.

Prueschoft, H. 2004. Functional structures of the skull in hominoidea. Folia Primatologica 75(4), 219-252.

Puech, P-F., Albertini, H. & Serratrice, C. 1983. Tooth microwear and dietary patterns in early hominids from Laetoli, Hadar and Olduvai. JHE 12, 721-729.

Puech, P.-F., Cianfarani, F. & Roth, H. 1986. Reconstruction of the maxillary dental arcade of Garusi Hominid 1. JHE 15, 325-332.

Purvis, A. 1995. A composite estimate of primate phylogeny. Phil. Trans. R. Soc. Lond. B. 348, 405-421.

468

Purvis, A., Nee, S. & Harvey, P.H. 1995. Macroevolutionary inferences from primate phylogeny. Phil. R. Soc. Lond. B. 260, 329-333.

Purvis, A. & Webster, A.J. 1999. Phylogenetically independent comparisons and primate phylogeny. In, (Lee, P.C., Ed.) Comparative primate socioecology, pp.44-70. UK: Cambridge University Press.

Raaum, R.L., Sterner, K.N., Noviello, C.M., Stewart, C-B. & Disotell, T. 2005. Catarrhine primate divergence dates estimated from complete mitochondrial genomes: concordance with fossil and nuclear DNA evidence. JHE 48, 237-257.

Rae, T.C. 1997. The early evolution of the hominoid face. In, (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 59-77. New York: Plenum Press.

Rae, T.C. 1999. Mosaic evolution in the origin of the hominoidea. Folia Primatologica 70, 126-135.

Rak, Y. 1983. The Australopithecine Face. New York: Academic Press.

Rak, Y. 1988. On variation in the masticatory system of Australopithecus boisei. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.193-198. New York: Aldine de Gruyter.

Rak, Y., Ginzburg, A. & Geffen, E. 2007 Gorilla-like anatomy on Australopithecus afarensis mandibles suggests Au. afarensis link to robust australopiths. PNAS 104, 6568-6572.

Rao, C.R. 1964. The use and interpretation of principal component analysis in applied research. Sankhya: The Indian Journal of Statistics, series A. 26, 329-358.

Raup, D.M. & Stanley, S.M. 1978. Principles of paleontology, 2nd ed. New York: W.H. Freeman.

Ravosa, M.J. & Profant, L.P. 2000. Evolutionary morphology of the skull in cercopithecoids. In, (Whitehead, P.F. & Jolly, C.J., Eds.) Old World Monkeys, pp. 237-268. UK: Cambridge Uni. Press.

Reed, C.A. 1967. The generic allocation of the hominid species habilis as a problem in systematics. SAJS 63, 3-5.

Reed, K.E. 1997. Early hominid evolution and ecological change through the African Plio-Pleistocene. JHE 32, 289-322.

Relethford, J.H. 1994. Craniometric variation among modern human populations. AJPA 95, 53-62.

Relethford, J.H. 2002. Apportionment of global human genetic diversity based on craniometrics and skin color. AJPA 118, 393-398.

Remis, M.J. 1998. The gorilla paradox: The effects of body size and habitat on the positional behavior of lowland and mountain gorillas. In, (Strasser, E., Fleagle, J., Rosenberger, A. & McHenry, H., Eds.) Primate locomotion: Recent advances, pp. 95-106. New York: Plenum Press.

Remis, M.J. 2003. Are gorillas vacuum cleaners of the forest floor? The roles of body size, habitat, and food preferences on dietary flexibility and nutrition. In, (Taylor, A. & Goldsmith, M.L., Eds.) Gorilla biology: a multidisciplinary perspective, pp. 385-404. UK: Cambridge Uni. Press.

Ren, R., Su, Y., Yan, K., Li, J., Yin, Z., Zhu, Z., Hu, Z. & Hu, Y. 1998a. Preliminary survey of the social organization of Rhinopithecus [Rhinopithecus] roxellana in Shennongjia National Natural Reserve, Hubei, China. In, (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub-Nosed Monkeys, pp. 269-278. Singapore: World Scientific.

469

Reno, P.L., DeGusta, D., Serrat, M.A., Meindl, R.S., White, T.D., Eckhardt, R.B., Kuperavage, A.J., Galik, K. & Lovejoy, C.O. 2005. Plio-Pleistocene hominid limb proportions. Current Anthropology 46(4), 575-588.

Reno, P.L., Meindl, R.S., McCollum, M.A. & Lovejoy, C.O. 2005. The case is unchanged and remains robust. Australopithecus afarensis exhibits only moderate skeletal dimorphism. JHE 49, 279-288.

Rhesus Macaque Genome Sequencing and Analysis Consortium. 2007. Evolutionary and biomedical insights from the Rhesus macaque genome. Science 316, 222-234.

Richmond, B.G. & Strait, D.S. 2000. Evidence that humans evolved from a knuckle-walking ancestor. Nature 404, 382-385.

Richmond, B.G., Begun, D.R. & Strait, D.S. 2001. Origin of human bipedalism: The knuckle-walking hypothesis revisited. Yearbook of Physical Anthropology 44, 70-106.

Richmond, B.G., Aiello, L.C. & Wood, B.A. 2002. Early hominin limb proportions. JHE 43, 529-548. + Erratum to 2002. ‘Early hominin limb proportions’. JHE 43, 529–548.

Rightmire, G.P. 1969. On the computation of Mahalanobis’ generalized distance (D2). AJPA 30, 157-160.

Rightmire, G.P. 1993. Variation among early Homo crania from Olduvai Gorge and the Koobi Fora Region. AJPA 90, 1-33.

Rightmire, G.P. 1995a. Geography, time and speciation in Pleistocene Homo. SAJS 91, 450-454.

Rightmire, G.P. 1995b. Diversity within the genus Homo. In (E.S. Vrba, G.H. Denton, T.C. Partridge & L.H. Burkle, Eds.) Paleoclimate and Evolution with Emphasis on Human Origins, pp. 483-492. New Haven: Yale University Press.

Rightmire, G.P. 1996. The human cranium from Bodo, Ethiopia: evidence for speciation in the Middle Pleistocene? JHE 31, 21-39.

Rightmire, G.P. 2004. Brain size and encephalization in early to Mid-Pleistocene Homo. AJPA 124, 109- 123.

Rightmire, G.P., Lordkipanidze, D. & Vekua, A. 2006. Anatomical descriptions, comparative studies and evolutionary significance of the hominin skulls from Dmanisi, Republic of Georgia. JHE 50(2), 115-141.

Robbins, M.M. 2007. Gorillas: Diversity in ecology and behavior. In, (Campbell, C.J., Fuentes, A., MacKinnon, K.C., Panger, M. & Bearder, S.K., Eds.) Primates in Perspective, pp. 305-320. UK: Oxford University Press.

Robinson, J.T. 1954. The genera and species of the Australopithecinae. AJPA 12, 181-200.

Robinson, J.T. 1956. The dentition of the Australopithecinae. Transvaal Museum Memoir No. 9. 1-179.

Robinson, J.T. 1965. Homo ‘habilis’ and the Australopithecines. Nature 205, 121-124.

Robinson, J.T. 1972. Early hominid posture and locomotion. Chicago: University of Chicago Press.

Rogers, J. 1993. The phylogenetic relationships among Homo, Pan and Gorilla: a population genetics perspective. JHE 25 201-215.

Rogers, J. 2000. Molecular genetic variation and population structure in Papio. In, (Whitehead, P.F. & Jolly, C.J., Eds.) Old World Monkeys, pp. 57-76. UK: Cambridge Uni. Press.

470

Rook, L., Martínez-Navarro, B. & Howell, F.C. 2004. Occurrence of Theropithecus sp. in the Late Villafranchian of Southern Italy and implication for Early Pleistocene “out of Africa” dispersals. JHE 47, 267-277.

Roos, C. & Geissman, T. 2001. Molecular phylogeny of major hylobatid divisions. MPE 19(3), 846-494.

Roos, C., Ziegler, T., Hodges, J.K., Zischler, H. & Abegg, C. 2003. Molecular phylogeny of Mentawai macaques: taxonomic and biogeographic implications. MPE 29, 139-150.

Rose, M.D. 1984. Food acquisition and the evolution of positional behavior: the case of bipedalism. In, (Chivers, D.J., Wood, B.A. & Bilsborough, A., Eds.) Food acquisition and processing in primates, pp. 509- 524. New York: Plenum Press.

Roseman, C.C. 2004. Detecting interregionally diversifying natural selection on modern human cranial form by using matched molecular and morphometric data. PNAS 101, 12824-21829.

Ross, C. 1992. Life history patterns and ecology of macaque species. Primates 33, 207-215.

Ross, C. & Jones, K.E. 1999. Socioecology and the evolution of primate reproductive patterns rates. In, (Lee, P.C., Ed.) Comparative primate socioecology, pp.73-110. UK: Cambridge University Press.

Rossie, J.B. & MacLatchy, L. 2006. A new pliopithecoid genus from the early Miocene of Uganda. JHE 50, 568-586.

Rowe, N. 1996. The Pictorial Guide to the Living Primates. New York: Pogonias Press.

Rowel, T.E. 1988. The social systems of guenons, compared with baboons, macaques and mangabeys. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 439-451. UK: Cambridge Uni. Press.

Ruff, C.B., Trinkaus, E. & Holliday, T.W. 1997. Body mass and encephalization in Pleistocene Homo. Nature 387, 173-176.

Ruff, C. 1998. Evolution of the hominid hip. In, (Strasser, E., Fleagle, J., Rosenberger, A. & McHenry, H., Eds.) Primate locomotion: Recent advances, pp. 449-469. New York: Plenum Press.

Ruvolo, M. 1988. Genetic evolution in the African guenons. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J- P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 127-139. UK: Cambridge Uni. Press.

Ruvolo, M. 1997a. Molecular Phylogeny of the Hominoids: Inferences from Multiple Independent DNA Sequence Data Sets. Mol. Biol. Evolution 14, 248-265.

Ruvolo, M. 1997b. Genetic diversity in hominoid primates. Ann. Rev. Anthrop. 26, 515-540.

Ruvolo, M., Pan, D., Zehr, S., Goldberg, T., Disotell, T.R. & von Dornum, M. 1994. Gene trees and hominoid phylogeny. PNAS 91, 8900-8904.

Ryan, A.S. 1981. Anterior dental microwear and its relationship to diet and feeding behavior in three African Primates (Pan troglodytes troglodytes, Gorilla gorilla gorilla, Papio hamadryas). Primates 22(4), 533-550.

Saitou, N. Evolution of hominoids and the search for a genetic basis for creating humanness. Cytogenet. Genome Res. 108, 16-21.

471 Sakka, M. 1984. Cranial morphology and masticatory adaptations. In, (Chivers, D.J., Wood, B.A. & Bilsborough, A., Eds.) Food acquisition and processing in primates, pp. 415-428. New York: Plenum Press.

Santa Luca, A.P. 1980. The Ngandong fossil hominids: A comparative study of a Far Eastern Homo erectus group. Yale University Publications in Anthropology number 78. New Haven: Dept. of Anthropology Yale University.

Sarich, V.M. & Wilson, A.C. 1967. Immunological time scale for hominid evolution. Science 158, 1200- 1203.

Sarmiento, E.E. 1994. Terrestrial traits in the hands and feet of gorillas. American Museum Novitates 3091, 1- 56.

Sarmiento, E.E. 1998. Generalized quadrupeds, committed bipeds and the shift to open habitats: an evolutionary model of hominoid divergence. American Museum Novitates 3250, 1-77.

Sarmiento, E.E. & Marcus, L.F. 2000. The os navicular of humans, great apes, OH 8, Hadar, and Oreopithecus: Function, Phylogeny, and Multivariate Analyses. American Museum Novitates 3288, 1-38.

Sarmiento, E.E. & Oates, J.F. 2000. The Cross River gorillas: A distinct subspecies, Gorilla gorilla diehli Matschie 1904. American Museum Novitates 3304, 1-56.

Sarmiento, E.E., Stiner, E. & Mowbray, K. 2002. Morphology-based systematics (MBS) and problems with fossil hominoid and hominid systematics. The Anatomical Record (New Anat.) 269, 50-66.

Satta, Y., Klein, J. & Takahata, N. 2000. DNA archives and our nearest relative: The trichotomy problem revisited. MPE 14, 259-275.

Sawada, Y., Pickford, M., Senut, B., Itaya, T., Hyodo, M., Miura, T., Kashine, C., Chujo, T. & Fujii, H. 2002. The age of Orrorin tugenensis, an early hominid from the Tugen Hills, Kenya. C.R. Palevol. 1, 293- 303.

Scheffrahn, W., de Ruiter, J.R. & van Hooff, J.A.R.A.M. 1996. Genetic relatedness within and between populations of Macaca fascicularis on Sumatra and off-shore islands. In (Fa, J.E. & Lindburg, D.G., Eds.) Evolution and Ecology of Macaque Societies, pp. 20-42. UK: Cambridge Uni. Press.

Schmitt, D. 1998. Forelimb mechanics during arboreal and terrestrial quadrupedalism in Old World monkeys. In, (Strasser, E., Fleagle, J., Rosenberger, A. & McHenry, H., Eds.) Primate locomotion: Recent advances, pp. 175-200. New York: Plenum Press.

Schneider, H., Schneider, M.P.C., Sampaio, M.I.C., Harada, M.L., Stanhope, M., Czelusniak, J. & Goodman, M. 1993. Molecular phylogeny of the New World Monkeys (Platyrrhini, Primates). Mol. Phylo. Evol. 2, 225- 242.

Schillaci, M.A. & Froehlich, J.W. 2001. Nonhuman primate hybridization and the taxonomic status of Neanderthals. AJPA 115, 157-166.

Schrenk, F., Bromage, T.G., Betzler, C.G., Ring, U. & Juwayeyi, Y.M. 1993. Oldest Homo and Pliocene biogeography of the Malawi Rift. Nature 365, 833-836.

Schultz, A.H. 1930. The skeleton of the trunk and limbs of higher primates. Human Biology 2, 303-438.

Schultz, A.H. 1933. Observations on the growth, classification and evolutionary specializations of gibbons and siamangs. Human Biology 5, 212-255 & 385-428.

472 Schultz, A.H. 1936a. Characters common to higher primates and characters specific for man. Quarterly Review of Biology 11, 259-283.

Schultz, A.H. 1936b. Characters common to higher primates and characters specific for man (continued). Quarterly Review of Biology 11, 425-455.

Schultz, A.H. 1940. Growth and development of the chimpanzee. Contrib. Embry. 170, 1-63.

Schultz, A.H. 1950. Origin of the human stock: The specializations of man and his place among the catarrhine primates. Cold Springs Harbor Symposia on Quantitative Biology 15, 37-53.

Schultz, A.H. 1958. Cranial and Dental variability in Colobus monkeys. PZSL 130, 79-105.

Schultz, A.H. 1963a. Age changes, sex differences, and variability as factors in the classification of primates. In (S.L. Washburn, ed.) Human Evolution and Classification, 85-115. London: Methuen & CO. Limited.

Schultz, A.H. 1963b. The relative lengths of the foot skeleton and its main parts in primates. Symp. Zool. Soc. Lond. 10, 199-206.

Schultz, A.H. 1969. The skeleton of the chimpanzee. The Chimpanzee vol. 1, pp. 50-103. Basel: Karger.

Schultz, A.H. 1970. The comparative uniformity of the Cercopithecoidea. In, (Napier, J.R. & Napier, P.H., Eds.) Old World monkeys: evolution, systematics, and behavior, pp. 39-52. New York: Academic Press.

Schultz, A.H. 1973. The skeleton of the Hylobatidae and other observations on their morphology. Gibbon and Siamang, vol. 2, pp. 1-54. Basel: Karger.

Schultz, M. 1986. The forelimb of the Colobinae. In, (J. Erwin & D.R. Swindler, Eds.) Comparative Primate Biology, Volume 1: Systematics, Evolution and Anatomy, pp. 559-670. New York: Alan R. Liss, Inc.

Schwartz, J.H. 1986. Primate systematics and a classification of the order. In, (J. Erwin & D.R. Swindler, Eds.) Comparative Primate Biology, Volume 1: Systematics, Evolution and Anatomy, pp. 1-42. New York: Alan R. Liss, Inc.

Schwartz, J.H. 1997. Lufengpithecus and hominoid phylogeny: problems in delineating and evaluating phylogenetically relevant characters. In (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 363-388. New York: Plenum Press

Schwartz, J.H., Gabunia, L., Vekua, A. & Lordkipanidze, D. 2000. Taxonomy of the Dmanisi crania. Science 289, 55-56.

Schwartz, J.H., Collard, M. & Cela-Conde, C.J. 2001. Systematics of “Humankind”. Evolutionary Anthropology 10, 1-3.

Schwartz, J.H. & Tattersall, I. 2002. The human fossil record: genus Homo (Europe), vol. 1. USA: Wiley- Liss.

Schwartz, J.H. & Tattersall, I. 2003. The human fossil record: genus Homo (Africa and Asia), vol. 2. USA: Wiley-Liss.

Schwartz, J.H. & Tattersall, I. 2005. The human fossil record: early hominids and overview, vol. 4. USA: Wiley-Liss.

Semaw, S., Renne, P., Harris, J.W.K., Feibel, C.S., Bernor, R.L., Fesseha, N. & Mowbray, K. 1997. 2.5- million-year-old stone tools from Gona, Ethiopia. Nature 385, 333-336.

473 Semendeferi, K., Lu, A., Schenker, N. & Damasio, H. 2002. Humans and great apes share a large frontal cortex. Nature Neuroscience 5, 272-276.

Senut, B. & Tardieu, C. 1985. Functional aspects of Plio-Pleistocene hominid limb bones: Implications for taxonomy and phylogeny. In, (E. Delson, Ed.) Ancestors: the Hard Evidence, pp. 193-201. New York: Alan R. Liss, Inc.

Senut, B. 2002. From apes to humans: locomotion as a key feature for phylogeny. Z. Morph. Anthrop. 83, 351-360.

Senut, B., Pickford, M., Gommery, D., Meint, P. & Cheboi, K. 2001. First hominid from the Miocene (Lukeino Formation, Kenya). C.R. Acad. Sci. Paris. 332, 137-144.

Setchell, J.M., Lee, P.C., Wickings, E.J. & Dixson, A.F. 2001. Growth and ontogeny of sexual size dimorphism in the Mandrill (Mandrillus sphinx). AJPA 115, 349-360.

Shea, B.T. 1985. On aspects of skull form in African apes and Orangutans, with implications for hominoid evolution. AJPA 68, 329-342.

Shea, B.T. 1987. Reproductive strategies, body size, and encephalization in primate evolution. IJP 8, 139- 156.

Shea, B.T. 1992. Ontogenetic scaling of skeletal proportions in the talapoin monkey. JHE 23, 283-307.

Shea, B.T. & Coolidge, H.J. 1988. Craniometric differentiation and systematics in the genus Pan. JHE 17, 671-685.

Shea, B.T., Leigh, S.R. & Groves, C.P. 1993. Multivariate craniometric variation in chimpanzees: Implications for species identification in paleoanthropology. In, (W.H. Kimbel & B.M. Lawrence, Eds.) Species, Species Concepts, and Primate Evolution, pp. 265-296. New York: Plenum Press.

Shelmidine, N., Borries, C. & Koenig, A. 2007. Genital swellings in silvered langurs: What do they indicate? Am. J. Primatol. 69(5), 519-532.

Shi, J., Xi, H., Wang, Y., Zhang, C., Jiang, Z., Zhang, K., Shen, Y., Jin, L., Zhang, K., Yuan, W., Wang, Y., Lin, J., Hua, Q., Wang, F., Xu, S., Ren, S., Xu, S., Zhao, G., Chen, Z., Li Jin, L. & Huang, W. 2003. Divergence of the genes on human chromosome 21 between human and other hominoids and variation of substitution rates among transcription units. PNAS 100, 8331-8336.

Shoshani, J., Groves, C.P., Simons, E.L. & Gunnell, G.F. 1996. Primate phylogeny: Morphological vs. molecular results. MPE 5, 102-154.

Sigmon, B.A. 1986. Evolution in the hominid pelvis. Palaeont. Africana 26(3), 25-32.

Simons, E.L. 1963. Some fallacies in the study of hominid phylogeny. Science 141, 879-889.

Simons, E.L. 1970. The deployment and history of Old World monkeys (Cercopithecidae, Primates). In, (Napier, J.R. & Napier, P.H., Eds.) Old World monkeys: evolution, systematics, and behavior, pp. 97-138. New York: Academic Press.

Simons, E.L. 1978. Diversity among the early hominids: A vertebrate paleontologist’s viewpoint. In, (C. Jolly, Ed.) Early Hominids of Africa, pp. 543-566. New York: St. Martin’s Press.

Simons, E.L. 1985. Origins and characteristics of the first hominoids. In, (E. Delson, Ed.) Ancestors: the Hard Evidence, pp. 37-41. New York: Alan R. Liss, Inc.

474 Simons, E.L. 2003. The fossil record of Tarsier evolution. In, (Wright, P.C., Simons, E.L. & Gursky, S., Eds.) Tarsiers: past, present, and future, pp.9-34. New Brunswick: Rutgers University Press.

Simons, E.L. & Fleagle, J.G. 1973. The history of extinct gibbon-like primates. Gibbon and Siamang, vol. 2, pp. 121-148. Basel: Karger.

Simons, E.L. & Rasmussen, D.T. 1996. Skull of Catopithecus browni, an early Tertiary Catarrhine. AJPA 100, 261-292.

Simpson, G.G. 1943. Criteria for genera, species, and subspecies in zoology and paleozoology. Ann. New York Acad. Sciences XLIV, 145-178.

Simpson, G.G. 1945. The principles of classification and a classification of mammals. Bulletin of the American Museum of Natural History 85.

Simpson, G.G. 1950. Origin of the human stock: Some principles of historical biology bearing on human origins. Cold Springs Harbor Symposia on Quantitative Biology 15, 55-66.

Simpson, G.G. 1959. The nature and origin of supraspecific taxa. Cold Springs Harbor Symposia on Quantitative Biology 24, 255-271.

Simpson, G.G. 1961. Principles of Animal Taxonomy. New York: Columbia University Press.

Simpson, G.G. 1963. The meaning of taxonomic statements. In (S.L. Washburn, Ed.) Classification and Human Evolution, pp. 1-31. Chicago: Aldine Publishing Co.

Simpson, G.G. 1966. The biological nature of man. Science 152, 472-478.

Simpson, G.G., Roe, A. & Lewontin, R. 1960. Quantitative Zoology. New York: Harcourt, Brace & World.

Singleton, M. 2002. Patterns of cranial shape variation in the Papionini (Primates: Cercopithecidae). JHE 42, 547-578.

Singleton, M. 2004a. Geometric morphometric analysis of functional divergence in mangabey facial form. JAS 82, 27-44.

Singleton, M. 2004b. Fossil hominoid diets, extractive foraging, and the origins of great ape intelligence. In (Russon, A.E. & Begun, D.R., Eds.) The Evolution of Thought: Evolutionary origins of Great Ape Intelligence, pp. 298-319. UK: Cambridge Uni. Press.

Skelton, R.R., McHenry, H.M., & Drawhorn, G.M. 1986. Phylogenetic Analysis of Early Hominids. Current Anthropology 27(1), 21-43.

Smith, A.B. & Littlewood, D.T.J. 1994. Paleontological data and molecular phylogenetic analysis. Paleobiology 20, 259-273.

Smith, B.H. 1986. Dental development in Australopithecus and early Homo. Nature 323, 327-330.

Smith, B.H. 1991. Dental development and the evolution of life history in hominidae. AJPA 86, 157-174.

Smith, B.H. 1994a. Sequence of emergence of the permanent teeth in Macaca. Pan, Homo, and Australopithecus: its evolutionary significance. AJHB 6, 61-76.

Smith, B.H. 1994b. Patterns of dental development in Homo, Australopithecus, Pan, and Gorilla. AJPA 94, 307-325.

475 Smith, F.A. Brown, J.H., Haskell, J.P., Lyons, S.K., Alroy, J., Charnov, E.L., Dayan, T., Enquist, B.J. Ernest, S.K.M., Hadly,E.A., Jones, K.E., Kaufman, D.M., Marquet, P.A., Maurer, B.A., Niklas, K.J., Porter, W.P., Tiffney, B. & Willig, M.R.. 2004. Similarity of Mammalian Body Size across the Taxonomic Hierarchy and across space and time. The American Naturalist 163(5), 672-691.

Smith, F.H. 2002. Migrations, radiations and continuity: Patterns in the evolution of Middle and Late Pleistocene humans. In (W.C. Hartwig, Ed.) The Primate Fossil Record, pp. 437-456. UK: Cambridge University Press.

Smith, R.J. 2005. Species recognition in paleoanthropology: Implications of small sample sizes. In (Lieberman, D.E., Smith, R.J., Kelley, J., Eds.) Interpreting the past: Essays on human, primate, and mammal evolution in honor of David Pilbeam, p. 207-219. Boston: Brill Academic Publishers, Inc.

Smith, R.J. & Jungers, W.L. 1997. Body mass in comparative primatology. JHE 32, 523-559.

Smith, K.K. 1997. Comparative patterns of craniofacial development in Eutherian and Metatherian mammals. Evolution 51: 1663–1678.

Sneath, P.H.A. & Sokal, R.R. 1973. Numerical Taxonomy: The principles and practice of numerical taxonomy. San Francisco: W.H. Freeman.

Sokal, R.R. 1974. Classification: Purposes, Principles, Progress, Prospects. Science 185, 1115-1123.

Spencer, F. 1995. Pithekos to Pithecanthropus: An abbreviated review of changing scientific views on the relationships of the anthropoid apes to Homo. In (R. Corbey & B. Theunissen, Eds.) Ape, Man, Apeman: Changing Views since 1600, pp. 13-27. Leiden University: Department of Prehistory.

Spocter, M. A. & Manger, P.R. 2007. The use of cranial variables for the estimation of body mass in fossil hominins. AJPA.

Sponheimer, M. & Lee-Thorp, J. 1999. Isotopic evidence for the diet of an early hominid, Australopithecus africanus. Science 283, 368-370.

Sponheimer, M., Passey, B.H., de Ruiter, D., Guatelli-Steinberg, D. Cerling, T.E. & Lee-Thorp, J. 2006. Isotopic evidence for dietary variability in the early hominin Paranthropus robustus. Science 314, 980-982.

Spoor, F., Wood, B. & Zonneveld, F. 1994. Implications of early hominid labyrinthine morphology for evolution of human bipedal locomotion. Nature 369, 645-648.

Spoor, F. 1997. Basicranial architecture and relative brain size of Sts 5 (Australopithecus africanus) and other Plio-Pleistocene hominids. SAJS 93, 182-186.

Spoor, F., Leakey, M.G., Gathogo, P.N., Brown, F.H., Anton, S.C., McDougall, I., Kiare, C., Manthi, F.K. & Leakey, L.N. 2007. Implications of new early Homo fossils from Ileret, east of Lake Turkana, Kenya. Nature 448, 688-691.

Stanford, C.B. 2006. The behavioral ecology of sympatric African apes: implications for understanding fossil hominoid ecology. Primates 47, 91-101.

Stanley, S.M. 1978. Chronospecies’ longevities, the origin of genera, and the punctuational model of evolution. Paleobiology 4, 26-40.

Stanley, S.M. 1992. An ecological theory for the origin of Homo. Paleobiology 18, 237-257.

Stanyon, R., Patton, W.H., Kurland, J.A. & Taub, D.M. 1980. G-Banding in the Barbary Macaque (Macaca sylvanus, Linnaeus, 1758) of Morocco. JHE 9, 147-152.

476

Stanyon, R., Bruening, R., Stone, G., Shearin, A. & Bigoni, F. 2005. Reciprocal painting between humans, De Brazza’s and patas monkeys reveals a major bifurcation in the Cercopithecini phylogenetic tree. CGR 108, 175-182.

StatSoft, Inc. 2007. Electronic Statistics Textbook. Tulsa, OK: StatSoft. WEB: http://www.statsoft.com/textbook/stathome.html.

Stauffer, R.L., Walker, A., Ryder, O.A., Lyons-Weiler, M. & Hedges, S.B. 2001. Human and ape molecular clocks and constraints on paleontological hypotheses. Journal of Heredity 92(6), 470-474.

Stearn, W.T. 1959. The background of Linnaeus’s contributions to the nomenclature and methods of systematic biology. Systematic Zoology 8, 4-22.

Stedman, H.H., Kozyak, B.W., Nelson, A., Theiser, D.M., Su, L.T., Low, D.W., Bridges, C.R., Shrager, J.B., Minugh-Purvis, N. & Mitchell, M.A. 2004. Myosin gene mutation correlates with anatomical changes in the human lineage. Nature 428, 415-418.

Steiper, M.E. 2006. Population history, biogeography, and taxonomy of orangutans (Genus: Pongo) based a population genetic meta-analysis of multiple loci. JHE 50, 509-522.

Steiper, M.E., Young, N.M. 2006. Primate molecular divergence dates. MPE 41, 384-394.

Steiper, M.E., Young, N.M. & Sukarna, T.Y. 2004. Genomic data support the hominoid slowdown and an Early Oligocene estimate for the hominoid-cercopithecoid divergence. PNAS 101, 17021-17026.

Sterner, K.N., Raaum, R.L., Zhang, Y-P., Stewart, C-B. & Disotell, T.R. 2006. Mitochondrial data support an odd-nosed colobine clade. MPE 40, 1-7.

Stiles, C.W. & Orleman, M.B. 1926. Retention of Cercopithecus, type Diana, for the guenons. Journal of Mammalogy 7(1), 48-53.

Stiner, M.C. 2002. Carnivory, coevolution, and the geographic spread of the genus Homo. Journal of Archaeological Science 10, 1-63.

Stone, A.C., Griffiths, R.C., Zegura, S.L. & Hammer, M.F. 2002. High levels of Y-chromosome nucleotide diversity in the genus Pan. PNAS 99, 43-48.

Strait, D.S., Grine, F.E. & Moniz, M.A. 1997. A reappraisal of early hominid phylogeny. JHE 32, 17-82.

Strait, D.S. & Ross, C.F. 1999. Kinematic data on primate head and neck posture: Implications for the evolution of basicranial flexion and an evaluation of registration planes used in paleoanthropology. AJPA 108, 205-222.

Strait, D. & Grine, F.E.; and Asfaw, B., White, T., Lovejoy, L., Latimer, B. & McCollum, M. 1999. Cladistics and Early Hominid Phylogeny; and response. Science 285, p. 1209-1210.

Strait, D.S. & Grine, F.E. 2001. The systematics of Australopithecus garhi. Ludas Vitalis 9, 109-135.

Strait, D.S. & Grine, F.E. 2004. Inferring hominoid and early hominid phylogeny using craniodental characters: the role of fossil taxa. JHE 47, 399-452.

Strasser, E. & Delson, E. 1987. Cladistic analysis of cercopithecid relationships. JHE 16: 81-99.

Strasser, E. 1988. Pedal evidence for the origin and diversification of the cercopithecid clades. JHE 17, 225- 245.

477

Strasser, E. 1992. Hindlimb proportions, allometry, and biomechanics in old world monkeys (Primates, Cercopithecidae). AJPA 87, 187-213.

Strasser, E. 1994. Relative development of the hallux and pedal digit formulae in Cercopithecidae. JHE 26, 413-440.

Stringer, C.B. 1974. A multivariate study of the Petralona skull. JHE 3, 397-404.

Stringer, C.B. 1986. The credibility of Homo habilis. In (B. Wood, L. Martin & P. Andrews, Eds.) Major topics in primate and human evolution, pp. 266-294. Cambridge: Cambridge University Press.

Stringer, C.B. 1987. A numerical cladistic analysis for the genus Homo. JHE 16, 135-146.

Struhsaker, T.T., Butynski, T.M. & Lwanga, J.S. 1988. Hybridization between redtail (Cercopithecus ascanius schmidti) and blue (Cercopithecus mitis stuhlmanni) monkeys in the Kibale Forest, Uganda. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 452-476. UK: Cambridge Uni. Press.

Stumpf, R.M., Polk, J.D., Oates, J.F., Jungers, W.L., Hesey, C.P., Groves, C. & Fleagle, J.G. 2003. Patterns of diversity in gorilla cranial morphology. In, (Taylor, A. & Goldsmith, M.L., Eds.) Gorilla biology: a multidisciplinary perspective, pp. 35-61. UK: Cambridge Uni. Press.

Stumpf, R. 2007. Chimpanzees and bonobos: Diversity within and between species. In, (Campbell, C.J., Fuentes, A., MacKinnon, K.C., Panger, M. & Bearder, S.K., Eds.) Primates in Perspective, pp. 321-344. UK: Oxford University Press.

Su, Y., Ren, R., Yan, K., Li, J., Yin, Z., Zhu, Z., Hu, Z. & Hu, Y. 1998. Preliminary survey of the home range and ranging behavior of Golden monkeys (Rhinopithecus [Rhinopithecus] roxellana) in Shennongjia National Natural Reserve, Hubei, China. In (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub-Nosed Monkeys, pp. 255-268. Singapore: World Scientific.

Susman, R.L. & Stern, J.T. 1982. Functional Morphology of Homo habilis. Science 217: 931-934.

Susman, R.L., Stern, Jr., J.T., Jungers, W.L. 1984. Arboreality and bipedality in the Hadar hominids. Folia. Primato. 43, 113-156.

Susman, R.L., Stern, Jr., J.T. & Jungers, W.L. 1985. Locomotor adaptations in the Hadar hominids. In, (E. Delson, Ed.) Ancestors: the Hard Evidence, pp. 184-192. New York: Alan R. Liss, Inc.

Susman, R. 1988a. New postcranial remains from Swartkrans and their bearing on the functional morphology and behavior of Paranthropus robustus. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.149-172. New York: Aldine de Gruyter.

Susman, R.L. 1988b. Hand of Paranthropus robustus from Member 1, Swartkrans: fossil evidence for tool behavior. Science 240: 781-784.

Susman, R.L. 1994. Fossil evidence of early hominid tool use. Science 265: 1570-1573.

Suwa, G. 1988. Evolution of the “Robust” Australopithecine in the Omo succession: Evidence from mandibular premolar morphology. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.199-222. New York: Aldine de Gruyter.

Suwa, G., Asfaw, B., Beyene, Y., White, T.D., Katoh, S., Nagaoka, S., Nakaya, H., Uzawa, K., Renne, P. & WoldeGabriel, G. 1997. The first skull of Australopithecus boisei. Nature 389, 489 – 492.

478 Suwa, G., Kono, R.T., Shighiro, K., Asfaw, B. & Beyene, Y. 2007. A new species of great ape from the late Miocene epoch in Ethiopia. Nature 448, 921-924.

Swindler, D.R. & Wood, C.D. 1973. An Atlas of Primate Gross Anatomy: Baboon, Chimpanzee, and Man. Seattle: University of Washington Press.

Swisher, C.C., Curtis, G.H., Jacob, T., A. G. Getty, A.G., Suprijo, A. & Widiasmoro. 1994. Age of the Earliest Known Hominids in Java, Indonesia. Science 263, 1118-1121.

Swisher III, C.C., Rink, W.J., Antón, S.C., Schwarcz, H.P., Curtis, G.H., A. & Widiasmoro. 1996. Latest Homo erectus of Java: Potential Contemporaneity with Homo sapiens in Southeast Asia. Science 274, 1870- 1874.

Szalay, F. & Delson, E. 1979. Evolutionary history of the primates. San Diego: Academic Press.

Szalay, F.S. 1999. Paleontology and Macroevolution: on the theoretical conflict between and expanded synthesis and hierarchic punctuationism. In, (Bromage, T.G. & Schrenk, F., Eds.) African biogeography, climate change, & human evolution, pp. 35-56. New York: Oxford University Press.

Szamalek, J.M., Cooper, D.N., Hoegel, J., Hameister, H. & Kehrer-Sawatzki, H. 2007. Chromosomal speciation of humans and chimpanzees revisited: studies of DNA divergence within inverted regions. Cytogenet. Genome Res. 116, 53-60.

Tabachnick, B.G. & Fidell, L.S. 2007. Using multivariate statistics,5th ed. USA: Pearson Education, Inc.

Takahata, N. & Satta, Y. 1997. Evolution of the primate lineage leading to modern humans: Phylogenetic and demographic inferences from DNA sequences. PNAS 94, 4811-4815.

Tamura, K., & Nei, M. 1993. Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Mol. Biol. Evol. 10: 512–526.

Tappen, N. 1960. Problems of distribution and adaptation of the African monkeys. Current Anthropology 1, 91-120.

Tappen, N. 1968. Problems of the distribution and adaptation of the African monkeys. In, (Chiarelli, B., Ed.) Taxonomy and Phylogeny of Old World Primates with References to the Origin of Man, pp. 197-270. Torino, Italy: Casa Editrice Libraria - Rosenberg & Sellier.

Tarzami, S.T., Kringston, A.M., Conte, R.A. & Verma, R.S. 1997. Unique genomic sequences in human chromosome 16p are conserved in the great apes. Mol. Gen. Genet. 253, 512-514.

Tattersall, I. 1986. Species recognition in human paleontology. JHE 15, 165-175.

Tattersall, I. 1992. Species concepts and species identification in human evolution. JHE 22, 341-349.

Tattersall, I. 2000. Paleoanthropology: The last half-century. Evolutionary Anthropology 9, 2-16.

Tattersall, I., Delson, E. & Van Couvering, J. 1988. Encyclopedia of human evolution and prehistory. New York: Garland Publishing.

Teaford, M.F. 1993. Dental microwear and diet in extant and extinct Theropithecus: preliminary analyses. In (Jablonski, N.G., Ed.) Theropithecus: the rise and fall of a primate genus, pp. 331-349. UK: Cambridge University Press.

Teaford, M.F. & Ungar, P. 2000. Diet and the evolution of the earliest human ancestors. PNAS 97, 13506- 13511.

479

Telfer, P.T., Souquière, S., Clifford, S.L. Abernethy, K.A., Bruford, M.W. Disotell, T.R., Sterner, K.N. Roques,P., Marx, P.A. & Wickings, E.J. 2003. Molecular evidence for deep phylogenetic divergence in Mandrillus sphinx. Molecular Ecology 12, 2019-2024.

Teumer, J. & Green, H. 1989. Divergent evolution of part of the involucrin gene in the hominoids: Unique intragenic duplications in the gorilla and human. PNAS 86, 1283-1286.

Thackeray, J.F., Helbig, J. & Moss, S. 1995. Quantifying morphological variability within extant mammalian species. Palaeont. Afr. 32, 23-25.

Thackeray, J.F. 2003. Comparison of hominid fossils attributed to Australopithecus and Homo. SAJS 99, 241- 242.

Thackeray, J.F., Bellamy, C.L., Bellars, d., Bronner, G., Bronner, L., et al. 1997. Probabilities of conspecificity: application of a morphometric technique to modern taxa and fossil specimens attributed to Australopithecus and Homo. SAJS 93, 195-196.

Thackeray, J.F., Mdaka, S., Navsa, N., Moshau, R. & Singo, S. 2000. Morphometric analyses of conspecific males and females: an exploratory study of extant primate and extinct hominid taxa. SAJS 96, 534-536.

Thackeray, J.F., Maureille, B., Vandermeersch, Braga, J. & Chaix, R. 2005. Morphometric comparisons between Neanderthals and ‘anatomically modern’ Homo sapiens from Europe and the Near East. Annals of the Transvaal Museum 42, 47-51.

Thierry, B. 2007. The macaques: A double-layered social organization. In, (Campbell, C.J., Fuentes, A., MacKinnon, K.C., Panger, M. & Bearder, S.K., Eds.) Primates in Perspective, pp. 224-239. UK: Oxford University Press.

Theirry, B., Iwaniuk, A.N. & Pellis, S.M. 2000. The influence of phylogeny on the social behaviour of Macaques (Primates: Cercopithecidae, genus Macaca). Ethology 106, 713-728.

Thorpe, S.K.S., Holder, R.L. & Crompton, R.H. 2007. Origin of Human Bipedalism as an adaptation for locomotion on flexible branches. Science 316, 1328-1331.

Tobias, P.V. 1966. The distinctiveness of Homo habilis. Nature 209, 953-957.

Tobias, P.V. 1967. Olduvai Gorge: Volume 2, The Cranium and maxillary dentition of Australopithecus (Zinjanthropus) boisei. Cambridge: Cambridge University Press.

Tobias, P.V. 1968. The taxonomy and phylogeny of the australopithecines. In, (Chiarelli, B., Ed.) Taxonomy and Phylogeny of Old World Primates with References to the Origin of Man, pp. 277-315. Torino, Italy: Casa Editrice Libraria - Rosenberg & Sellier.

Tobias, P.V. 1978. The earliest Transvaal members of the genus Homo with another look at some problems of hominid taxonomy and systematics. Z. Morph. Anthrop. 69, 225-265.

Tobias, P.V. 1980. “Australopithecus afarensis” and A. africanus: Critique and alternative hypothesis. Palaeont. Afr., 23, 1-17.

Tobias, P.V. 1985. Single characteristics and the total morphological pattern redefined: The sorting effected by a selection of morphological features of the early hominids. In, (E. Delson, Ed.) Ancestors: the Hard Evidence, pp. 94-101. New York: Alan R. Liss, Inc.

Tobias, P.V. 1987. The brain of Homo habilis: a new level of organization in cerebral evolution. JHE 16, 741- 761.

480

Tobias, P.V. 1988. Numerous apparently synapomorphic features in Australopithecus robustus, Australopithecus boisei and Homo hablis: Support for the Skelton-McHenry-Drawhorn hypothesis. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp. 293-308. New York: Aldine de Gruyter.

Tobias, P.V. 1991. Olduvai Gorge: Volume 4, The Skulls, Endocasts and Teeth of Homo habilis, Parts I-IV & Parts V-IX. Cambridge: Cambridge University Press.

Tobias, P.V. 1994. The craniocerebral interface in early hominids. In, (Corruccini, R.S. & Ciochon, R.L., Eds.) Integrative paths to the past: Paleoanthropological advances in honor of F. Clark Howell, pp. 185-203. USA: Prentice Hall.

Tosi, A.J., Buzzard, P.J., Morales, J.C. & Melnick, D.J. 2002a. Y-chromosome data and tribal affiliations of Allenopithecus and Miopithecus. IJP 23, 1287-1399.

Tosi, J.T., Buzzard, P.J., Morales, J.C. & Melnick, D.J. 2002b. Y-Chromosomal window onto the history of Terrestrial Adaptation in the Cercopithecini. In, (Glenn, M.E. & Cords, M., Eds.) The Guenons: Diversity and adaptation in African monkeys, pp. 15-26. New York: Kluwer Academic / Plenum Publishers.

Tosi, A.J., Disotell, T.R., Morales, J.C. & Melnick, D.J. 2003a. Cercopithecine Y-chromosome data provide a test of competing morphological evolutionary hypotheses. MPE 27, 510-521.

Tosi, A., Morales, J.C., & Melnick, D.J. 2003b. Paternal, maternal, and biparental molecular markers provide unique windows onto the evolutionary history of macaque monkeys. Evolution 57(6), 1419-1435.

Tosi, A., Melnick, D.J. & Disotell, T. 2004. Sex chromosome phylogenetics indicate a single transition to terrestirality in the guenons (Tribe Cercopithecini). JHE 46, 223-237.

Trinkaus, E. 2007. European early modern humans and the fate of the Neandertals. PNAS 104, 7367-7372.

Turner, T.R., Maiers, J.E. & Mott, C.S. 1988. Population differentiation in Cercopithecus monkeys. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 140-149. UK: Cambridge Uni. Press.

Turner, T.R., Anapol, F. & Jolly, C.J. 1997. Growth, development, and sexual dimorphism in vervet monkeys (Cercopithecus aethiops) at four sites in Kenya. AJPA 103, 19-35.

Turner, A. 1986. Species, speciation and Human evolution. Human Evolution 1(5), 419-430.

Turner, A. & Wood, B. 1993. Taxonomic and geographic diversity in robust australopithecines and other African larger mammals. JHE 24, 147-168.

Turner, A. 1995. Plio-Pleistocene correlations between climatic and evolution in terrestrial mammals: the 2.5 ma event in Africa and Europe. Acta zool. Cracov. 38, 45-58.

Turner, A. 1999. Evolution in the African Plio-Pleistocene Mammalian Fauna: Correlation and causation. In, (Bromage, T.G. & Schrenk, F., Eds.) African biogeography, climate change, & human evolution, pp. 76-90. New York: Oxford University Press.

Turner, T., Maiers, J.E. & Mott, C.S. 1988. Population differentiation in Cercopithecus monkeys. In, (Gautier-Hion, A., Bourliere, F., Gautheir, J-P. & Kingdon, J., Eds.) A Primate Radiation: evolutionary biology of the African Guenons, pp. 140-149. UK: Cambridge Uni. Press.

Tutin, C.E.G., Fernandez, M., Rogers, M.E., Williamson, A. & Mcgrew, W.C. 1991. Foraging profiles of sympatric lowland gorillas and chimpanzees in the Lope Reserve, Gabon. Phil. Tran: Biol. Sci. 334, 179-185.

481

Tutin, C.E.G. & Oslisy, R. 1995. Homo, Pan and Gorilla: co-existence over 60 000 years at Lope in central Gabon. JHE 28, 597-602.

Tuttle, R. & Basmajian, J.V. 1974. Electromyography of brachial muscles in Pan gorilla and hominoid evolution. AJPA 41, 71-90.

Tuttle, R.H., Hallgrimsson, B & Stein, T. 1998. Heel, squat, stand, stride: Function and evolution of hominoid feet. In, (Strasser, E., Fleagle, J., Rosenberger, A. & McHenry, H., Eds.) Primate locomotion: Recent advances, pp. 435-448. New York: Plenum Press.

Tuttle, R.H. 2006a. Seven decades of East African Miocene Anthropoidea. In, (Ishida, H., Tuttle, R., Pickford, M., Ogihara, N. & Nakatsukasa, M., Eds.) Human Origins and Environmental Background, pp. 15- 30. USA: Springer.

Tuttle, R.H. 2006b. Neontological perspectives on East African Middle and Late Miocene Anthropoidea. In, (Ishida, H., Tuttle, R., Pickford, M., Ogihara, N. & Nakatsukasa, M., Eds.) Human Origins and Environmental Background, pp. 209-225. USA: Springer.

Tuttle, R.H. 2006c. Are human beings apes, or are apes people too? In, (Ishida, H., Tuttle, R., Pickford, M., Ogihara, N. & Nakatsukasa, M., Eds.) Human Origins and Environmental Background, pp. 249-258. USA: Springer.

Uchida, A. 1996a. What we don’t about great ape variation. TREE 11, 163-168.

Uchida, A. 1996b. Craniodental variation among the great apes. Peabody Museum Bulletin 4, pp. 1-184.

Uddin, M., Wildman, D.E., Liu, G., Xu, W., Johnson, R.M., Hof, P.R., Kapatos, G., Grossman, L.I. & Goodman, L.I. 2004. Sister grouping of chimpanzees and humans as revealed by genome-wide phylogenetic analysis of brain gene expression profiles. PNAS 101, 2957-2962.

Ungar, P.S., Grine, F.E., Teaford, M. & El Zaatari, S. 2006. Dental microwear and diets of African early Homo. JHE 50, 78-95.

Ungar, P. 2004. Dental topography and diets of Australopithecus afarensis and early Homo. JHE 46, 605-622.

Valentine, J.W. & May, C.L. 1996. Hierarchies in biology and paleontology. Paleobiology 22, 23-33.

Van der Kuyl, A. C., Kuiken, C. L., Dekker, J. T., and Goudsmit, J. (1995). Phylogeny of African monkeys based upon mitochondrial 12S rRNAsequences. J. Mol. Evol. 40(2): 173–180.

Van der Kuyl, A. C., Dekker, J. T., and Goudsmit, J. (2000). Primate genus Miopithecus: Evidence for the existence of species and subspecies of dwarf guenons based on cellular and endogenous viral sequences. Mol. Phylogenet. Evol. 14(3): 403–413.

Van Geel, M., Eichler, E.E., Beck, A.F., Shan, Z., Haaf, T., van der Maarel, M., Frants, R.R. & de Jong, P.J. 2002. A cascade of complex subtelomeric duplications during the evolution of the hominoid and Old World monkey genomes. Amer. J. Hum. Genet. 70, 269-278.

Van Gelder, R.G. 1977. Mammalian hybrids and generic limits. Amer. Museum Novitates 2635, 1-25.

Van Schaik, C.P., van Noordwijk, M.A. & Nunn, C.L. 1999. Sex and social evolution in primates. In, (Lee, P.C., Ed.) Comparative primate socioecology, pp.204-240. UK: Cambridge University Press.

Van Valen, L. 1971. Adaptive zones and the orders of mammals. Evolution 25(2), 420-428.

482 Vekua, A., Lordkipanidze, D., Rightmire, G.P., Agusti, J., Ferring, R., Maisuradze, G., Mouskhelishvili, A. Nioradze, M., Ponce de Leon, M., Tappen, M., Tvalchrelidze, M. & Zollikofer, C. 2002. A new skull of Early Homo from Dmanisi, Georgia. Science 297, 85-89.

Verhaegen, M.J.B. 1994. Australopithecines: Ancestors of the African apes? Human Evolution 9, 121-139.

Verhaegen, M. 1996. Morphological distance between australopithecine, human and ape skulls. Human Evolution 11, 35-41.

Verheyen, W.N. 1962. Contribution a la craniologie compare des primates: les genres Colobus Illiger 1811 et Cercopithecus Linne 1758. Koninklijk Museum Voor Midden-Afrika-Tervuren, Belgie. Annalen – Reeks IN 8 - Zoologische Wetenschappen – nr 105.

Vervaecke, H. & van Elsacker, L. 1992. Hybrids between common chimpanzee (Pan troglodytes) and pygmy chimpanzee (Pan paniscus) in captivity. Mammalia 56, 667-669.

Videan, E.N. & McGrew, W.C. 2002. Bipedality in chimpanzee (Pan troglodytes) and bonobo (Pan paniscus): Testing hypotheses on the evolution of bipedalism. AJPA 118, 184-190.

Vignaud, P., Duringer, P., Mackaye, H.T., Likius, A., Blondel, C., Boisserie, J-R., de Bonis, L., Eisenmann, V., Etienne, M-E., Geraads, D., Guy, F., Lehman, T., Lihoreau, F., Lopez-Martinez, N., Mourer-Chauvire, C., Otero, O., Rage, J-C., Schuster, M., Viriot, L., Zazzo, A. & Brunet, M. 2002. Geology and paleontology of the Upper Miocene Toros-Menalla hominid locality, Chad. Nature 418, 152-155.

Vogel, C. 1966. Morphologische studien am gesichtsschadel catarrhiner Primaten. Biblioth. primat., Vol. 4, Basel.

Vogel, C. 1968. The phylogenetical evaluation of some characters and some morphological trends in the evolution of the skull in catarrhine primates. In Taxonomy and Phylogeny of Old World Primates with References to the Origin of Man, pp. 21-55. Torino, Italy: Casa Editrice Libraria - Rosenberg & Sellier.

Vrba, E.S. 1985. Ecological and adaptive changes associated with early hominid evolution. In, (E. Delson, Ed.) Ancestors: the Hard Evidence, pp. 63-71. New York: Alan R. Liss, Inc.

Vrba, E.S. 1988. Late Pliocene climatic events and hominid evolution. In (F.E. Grine, Ed.) The Evolutionary history of the Robust Australopithecines, pp. 405-426. New York: Aldine Publishing Company.

Vrba, E.S. 1992. Mammals as a key to evolutionary theory. J. Mamm. 73, 1-28.

Vrba, E.S., Denton, G.H. & Prentice, M.L. 1989. Climatic influences on early hominid evolution. Ossa 14, 127-156.

Vrba, E.S. 1999. Habitat theory in relation to the evolution in African Neogene biota and hominids. In, (Bromage, T.G. & Schrenk, F., Eds.) African biogeography, climate change, & human evolution, pp. 19-33. New York: Oxford University Press.

Wahungu, G.M. 1998. Diet and habitat overlap in two sympatric primate species, the Tana crested mangabey Cercocebus galeritus and yellow baboon Papio cynocephalus. Afr. J. Ecol. 36, 159-173.

Walker, A., Leakey, R.E.F., Harris, J.M & Brown, F.H. 1986. 2.5-Myr Australopithecus boisei from west of Lake Turkana, Kenya. Nature 322, 517-522.

Walker, A. & Leakey, R.E.F. 1988. The evolution of Australopithecus boisei. In, (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.247-258. New York: Aldine de Gruyter.

483 Walker, A. 1976. Remains attributable to Australopithecus in the East Rudolf Basin. In, (Coppens, Y., Howell, F.C., Isaac, G.Ll. & Leakey, R.E.F., Eds.) Earliest man and environments in the Lake Rudolf basin, pp. 484-489. Chicago: University of Chicago Press.

Walker, R., Hill, K., Burger, O. & Hurtado, M. 2006. Life in the slow lane revisited: Ontogenetic separation between chimpanzees and humans. AJPA 129, 577-583.

Wang, W., Michaei, R. J. F., Zhang, Y. P., Liu, Z. M., Wei, Y., Huang, H. Q., Hu, H. G., Xie, Y. X., Wu, D. H., and Melnick, D. J. 1997. A phylogeny of Chinese leaf monkey using mitochondrial ND3-ND4 gene sequences. Int. J. Primatol. 18(3): 305–320.

Wang, W., Jiang, X. & Li, D. 1998. Classification and distribution of the extant subspecies of Golden Snub- nosed monkey. In (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub-Nosed Monkeys, pp. 53- 64. Singapore: World Scientific.

Watanabe, H., Fujiyama, A., Hattori, M., Taylor, T.D., Toyoda, A., Kuroki, Y., Noguchi, H., BenKahla, A., Lehrach, H., Sudbrak, R., Kube, M., Taenzer, S., Galgoczy, P., Platzer, M., Scharfe, M., Nordsiek, G., Blöcker, H., Hellmann, I., Khaitovich, P., Pääbo, S., Reinhardt, R., Zheng, H.-J., Zhang, X.-L., Zhu, G.-F., Wang, B.-F., Fu, G., Ren, S.-X., Zhao, G.-P., Chen, Z., Lee, Y.-S., Cheong, J.-E., Choi, S.-H., Wu, K.-M., Liu, T.-T., Hsiao, K.-J., Tsai, S.-F., Kim, C.-G., Ota, S., Kitano, T., Kohara, Y., Saitou, N., Park, H.-S., Wang, S.-Y., Yaspo, M.-L. & Sakaki, Y. 2004. DNA sequence and comparative analysis of chimpanzee chromosome 22. Nature 429, 382 – 388.

Ward, C.V. 1997. Functional anatomy and phyletic implications of the hominoid trunk and hindlimb. In (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 101-130. New York: Plenum Press.

Ward, C.V. 1998. Afropithecus, Proconsul, and the primitive hominoid skeleton. In, (Strasser, E., Fleagle, J., Rosenberger, A. & McHenry, H., Eds.) Primate locomotion: Recent advances, pp. 337-352. New York: Plenum Press.

Ward, C., Leakey, M. & Walker, A. 1999. The new hominid species Australopithecus anamensis. Evolutionary Anthropology 7, 197-205.

Ward, C.V. 2002. Interpreting the Posture and Locomotion of Australopithecus afarensis: Where Do We Stand? Yearbook of Physical Anthropology 45:185–215.

Ward, S. 1997. The taxonomy and phylogenetic relationships of Sivapithecus revisited. In (Begun, D.R., Ward, C.V. & Rose, M.D., Eds.) Function, phylogeny and fossils: Miocene hominoid evolution and adaptation, pp. 269-290. New York: Plenum Press

Ward, S., Brown, B., Hill, A., Kelley, J. & Downs, W. 1999. Equatorius: A new hominoid genus from the Middle Miocene of Kenya. Science 285, 1382-1386.

Ward, S.C. & Duren, D.L. 2002. Middle and Late Miocene African hominoids. In (W.C. Hartwig, Ed.) The Primate Fossil Record, pp. 385-398. UK: Cambridge University Press.

Waser, P. 1977. Feeding, ranging and group size in the mangabey Cercocebus albigena. In (Clutton-Brock, T.H., Ed.) Primate Ecology: Studies of feeding and ranging behaviour in lemurs, monkeys and apes, pp. 183- 222. London: Academic Press.

Washburn, S.L. 1944. The genera of the Malaysian Langurs. Journal of Mammalogy 25(3), 289-294.

Watkins, B.T. 2003. Hand bone ratios and their utility in predicting general substrate use in primates. Cour. Forsch.-Inst. Senckenberg 243, 47-59.

484 Watson, E.E., Easteal, S. & Penny, D. 2001. Homo Genus: A review of the classification of humans and the great apes’, In (P.V. Tobias, M.A. Raath, J. Moggi-Cecchi & G.A. Doyle, Eds.) Humanity from African Naissance to Coming Millenia, pp. 307-318. Firenze University Press and Witwatersrand University Press.

Wayne, R.K., van Valkenburgh, B. & O’Brien, S.J. 1991. Molecular distance and divergence time in carnivores and primates. Mol. Biol. Evol. 8(3), 297-319.

Weaver, T.D., Roseman, C.C. & Stringer, C.B. 2007. Were neandertal and modern human cranial differences produced by natural selection or genetic drift? JHE 53, 135-145.

Weber, J., Czarnetzki, A. & Pusch, C.M. 2005. Comment on "The Brain of LB1, Homo floresiensis". Science 310: 236.

Weidenreich, F. 1939. The relationship between Pithecanthropus and Sinanthropus. Nature 144, 926-929.

Weidenreich, F. 1943. The skull of Sinanthropus pekinensis; a comparative study on a primitive hominid skull. Palaeontologia Sinica New series D. no. 10; Whole series No. 127, 1-289.

Weidenreich, F. 1946. Generic, specific and subspecific characters in human evolution. AJPA 4, 413-431.

Weiner, J.S. 1958. The pattern of evolutionary development of the genus Homo. S. Afr. J. med Sci. 23, 111- 120.

Weitzel, V., Yang, C.M. & Groves, C.P. 1988. A catalogue of primates in the Singapore Zoological Reference collection, Department of Zoology, National University of Singapore. A Journal of Zoology of Southeast Asia 36(1), 1-167.

White, T.D. 1985. The hominids of Hadar and Laetoli: An element-by-element comparison of the dental samples. In, (E. Delson, Ed.) Ancestors: the Hard Evidence, pp. 138-152. New York: Alan R. Liss, Inc.

White, T.D. 2000. Human Osteology, 2nd ed. San Diego: Academic Press.

White, T.D. 2002. Earliest hominids. In (W.C. Hartwig, Ed.) The Primate Fossil Record, pp. 407-418. UK: Cambridge University Press.

White, T. 2003. Early hominids – diversity or distortion? Science 299, 1994-1997.

White, T.D. 2006. Early hominid femora: The inside story. Comptes Rendus Palevol 5, 99-108.

White, T.D., Johanson, D.C. & Kimbel, W.H. 1981. Australopithecus afarensis: its phyletic position reconsidered. SAJS 77, 445-470.

White, T.D., Suwa, G., Hart, W.K., Walter, R.C., Woldegabriel, G., de Heinzelin, J., Clark, J.D., Asfaw, B. & Vrba, E. 1993. New discoveries of Australopithecus at Maka in Ethiopia. Nature 366, 261-265.

White, T.D., Suwa, G. & Asfaw, B. 1994. Australopithecus ramidus, a new species of early hominid from Aramis, Ethiopia. Nature 371, 306-312.

White, T.D., Suwa, G. & Asfaw, B. 1995. Corrigendum: Australopithecus ramidus, a new species of early hominid from Aramis, Ethiopia. Nature 375, 88.

White, T.D. Asfaw, B., DeGusta, D., Gilbert, H., Richards, G.D., Suwa, G. & Howell, F.C. 2003. Pleistocene Homo sapiens from Middle Awash, Ethiopia. Nature 423, 742-747.

White, T.D., WoldeGabrie, G., Asfaw, B., Ambrose, S., Beyene, Y., Bernor, R.L., Boisserie, J-R., Currie, B., Gilbert, H., Haile-Selassie, Y., Hart, W.H, Hlusko, L.J., Howell, F.C., Kono, R.T., Lehmann, T., Louchart,

485 A., Lovejoy, C.O., RenneP.R., Saegusa, H., Vrba, E.S., Wesselman, H. & Suwa, G. 2006. Asa Issie, Aramis and the origin of Australopithecus. Nature 440, 883-889.

Whitehead, P.F., Sacco, W.K. & Hochgraf, S.B. 2005. A Photographic Atlas for Physical Anthropology. USA: Morton Publishing Company.

Whiten, A., Goodall, J., McGrew, W.C., Nishida, T., Reynolds, V., Sugiyama, Y., Tutin, C.E.G., Wrangham, R.W. & Boesch, C. 1999. Culture in Chimpanzees. Nature 399, 682-685.

Whittaker, D.J., Ting, N. & Melnick, D.J. 2006. Molecular phylogenetic affinities of the simakobu monkey (Simias concolor). MPE 39, 887-892.

Wildman, D., Uddin, M., Liu, G., Grossman, L.I., & Goodman, M. 2003. Implications of natural selection in shaping 99.4% nonsynonymous DNA identity between humans and chimpanzees: Enlarging the genus Homo. PNAS 100, 7181-7188.

Williams, S.A. & Goodman, M. 1989. A statistical test that supports a human/chimpanzee clade based on noncoding DNA sequence data. Mol. Biol. Evol. 6, 325-330.

Williams, F.L., Godfrey, L.R. & Sutherland, M.R. 2003. Diagnostic heterochronic perturbations in the craniofacial evolution of Homo (Neandertals and modern humans) and Pan (P. troglodytes and P. paniscus). In, (J.L. Thompson, G.E. Krovitz & A.J. Nelson, Eds.) Patterns of Growth and Development in the Genus Homo, pp. 295-319. Cambridge: Cambridge University Press.

Williamson, D.K. & Dunbar, R. 1999. Energetics, time budgets and group size. In, (Lee, P.C., Ed.) Comparative primate socioecology, pp. 320-338. UK: Cambridge University Press.

Willis, M.S. & Swindler, D.R. 2004. Molar size and shape variations among Asian colobines. AJPA 125, 51- 60.

Wilson, A.C., Kunkel, J.G. & Wyles, J.S. 1984. Morphological distance: an encounter between two perspectives in evolutionary biology. Evolution 38, 1156-1159.

Wimmer, R., Kirsch, S., Rappold, G.A. & Schempp, W. 2002. Direct evidence for the Homo-Pan clade. Chromosome Research 10, 55-61.

Woldegabriel, G., Haile-Selassie, Y., Renne, P.R., Hart, W.K., Ambrose, S.H., Asfaw, B., Heiken, G. & White, T. 2001. Geology and paleontology of the Late Miocene Middle Awash Valley, Afar rift, Ethiopia. Nature2, 175-178. 41

Wolfheim, J.H. 1983. Primates of the World. Seattle: Uni. Of Washington Press.

Wolpoff, M.H. 1971. Metric Trends in Hominid Dental Evolution. Case Western Reserve University in Anthropology 2. Cleveland.

Wolpoff, M.H. 1978. Analogies and interpretation in paleoanthropology. In (C. Jolly, Ed.) Early Hominids of Africa, pp. 263-503. Palgrave Macmillan.

Wolpoff, M.H. 1980. Cranial remains of Middle Pleistocene European hominids. JHE 9, 339-358.

Wolpoff, M.H. 1999. Paleoanthropology, 2nd ed. USA: McGraw-Hill.

Wolpoff, M.H.; and Wood, B. & Collard, M. 1999. The Systematics of Homo; and response. Science 284, p. 1773.

Wolpoff, M.H. & Lovejoy, C.O. 1975. A rediagnosis of the genus Australopithecus. JHE 4, 275-276.

486

Wolpoff, M.H., Hawks, J., Frayer, D.W. & Huntley, K. 2001. Modern human ancestry at the peripheries: a test of the replacement theory. Science 291, 293-297.

Wolpoff, M.H., Senut, B., Pickford, M. & Hawks, J.; Brunet, M. 2002. Sahelanthropus or “Sahelpithecus”. Nature 419, 581-582.

Wolpoff, M.H. & Lee, S.-H. 2006. Variation in the Habiline Crania – Must it be Taxonomic? Human Evolution 21, 71-84.

Wolpoff, M.H., Hawks, J., Senut, B., Pickford, M. & Ahern, J. 2006. An ape or the Ape: Is the Toumai cranium TM 266 a hominid. PaleoAnthropology 2006, 36-50.

Wood, B. 1976. Remains attributable to Homo in the East Rudolf Succession. In (Y. Coppens, F.C. Howells, G.Ll. Isaac & R.E.F. Leakey, Eds.) Earliest Man and Environments in the Lake Rudolf Basin: Stratigraphy, Paleoecology and Evolution, pp. 490-506. Chicago: University of Chicago Press.

Wood, B. 1988. Are “Robust”Australopithecines a monophyletic group? In (Grine, F.E., Ed.) Evolutionary history of the “Robust Australopithecines”, pp.269-284. New York: Aldine de Gruyter.

Wood, B. 1991. Koobi Fora research project Vol. 4 Hominid cranial remains. Oxford: Clarendon Press.

Wood, B. 1992. Origin and evolution of the genus Homo. Nature 355, 783-790.

Wood, B. 1993. Early Homo: How many species? In (W.H. Kimbel & B.M. Lawrence, Eds.) Species, Species Concepts, and Primate Evolution, pp. 485-522. New York: Plenum Press.

Wood, B. 1999. ‘Homo rudolfensis’ Alexeev, 1986-fact or phantom? JHE 36, 115-118.

Wood, B. & Chamberlain, A.T. 1986. Australopithecus: grade or clade? In (Wood, B., Martin, L. & Andrews, P., Eds.) Major topics in primate and human evolution, pp. 220-265.

Wood, B. & Collard, M. 1999a. The human genus. Science 284, 65-71.

Wood, B. & Collard, M. 1999b. The changing face of the genus Homo. Evolutionary Anthropology 8, 195- 207.

Wood, B. & Richmond, B.G. 2000. Human evolution: taxonomy and paleobiology. Journal of Anatomy 196, 19-60.

Wood, B. & Collard, M. 2001a. The meaning of Homo. Ludus Vitalis 9, 63-74.

Wood, B. & Collard, M. 2001b. Evolving interpretations of Homo. In (P.V. Tobias, M.A. Raath, J. Moggi- Cecchi & G.A. Doyle, Eds.) Humanity from African Naissance to Coming Millenia, pp. 141-146. Firenze University Press and Witwatersrand University Press.

Wood, B. & Strait, D. 2004. Patterns of resource use in early Homo and Paranthropus. JHE 46, 119-162.

Wooding, S., Bufe, B., Grassi, C., Howard, M.T., Stone, A.C., Vazquez, M., Dunn, D.M., Meyerhof, W., Weiss, R.B. & Bamshad, M.J. 2006. Independent evolution of bitter-taste sensitivity in humans and chimpanzees. Nature 440, 930-934.

Wu, Bao-Qt. 1993. Patterns of Spatial dispersion, locomotion and foraging behavior in three groups of the Yunnan snub-nosed langur (Rhinopithecus bieti). Folia. Primatol. 60, 63-71.

487 Xing, J., Wang, H., Han, K., Ray, D.A., Huang, C.H., Chemnick, L.G., Stewart, C-B., Disotell, T.R., Ryder, O.A. & Batzer, M.A. 2005. A mobile element based phylogeny of Old World monkeys. MPE 37, 872-880.

Xu, X. & Arnason, U. 1996. The mitochondrial DNA molecule of Sumatran Orangutan and a molecular proposal for two (Bornean and Sumatran) species of orangutan. Journal of Molecular Evolution 43, 431-437.

Yamakoshi, G. 1998. Dietary responses to fruit scarcity of wild chimpanzees at Bossou, Guinea: Possible implications for ecological importance of tool use. AJPA 106, 283-295.

Yamakoshi, G. 2004. Evolution of complex feeding techniques in primates: Is this the origin of great ape intelligence? In (Russon, A.E. & Begun, D.R., Eds.) The Evolution of Thought: Evolutionary origins of Great Ape Intelligence, pp. 140-171. UK: Cambridge Uni. Press.

Yeager, C.P. 1989. Feeding ecology of the Proboscis monkey. IJP 10, 497-530.

Yeager, C.P. & Kirkpatrick, R.C. 1998. Asian colobine social structure: ecological and evolutionary constraints. Primates 39(2), 147-155.

Yeager, C.P. & Kool, K. 2000. The behavioral ecology of Asian colobines. In, (Whitehead, P.F. & Jolly, C.J., Eds.) Old World Monkeys, pp. 496-521. UK: Cambridge Uni. Press.

Yerkes, R.M. & Yerkes, A.M. 1970. The Great Apes: A Study of Anthropoid Life. New Haven: Yale University Press.

Yirga, S. 2002. Hind limb bones and locomotion in the old world monkeys. SINET: Ethiop. J. Sci. 25(2), 205- 226.

Young, N.M. & MacLatchy, L. 2004. The phylogenetic position of Morotopithecus. JHE 46(2), 163-184.

Young, N.M. 2005. Estimating hominoid phylogeny from morphological data: Character choice, phylogenetic signal and postcranial data. In (Lieberman, D.E., Smith, R.J., Kelley, J., Eds.) Interpreting the past: Essays on human, primate, and mammal evolution in honor of David Pilbeam, p. 1-13. Boston: Brill Academic Publishers, Inc.

Yunis, J.J. & Prakash, O. 1982. The origin of man: A chromosomal pictorial legacy. Science 215, 1525-1530.

Zhang, Y. P., & Ryder, O.A. 1998. Mitochondrial cytochrome b gene sequences of old world monkeys: with special reference on evolution of Asia Colobines. Primates 39(1), 39–49.

Zhang, Y. P., & Shi, L. M. 1993. Phylogenetic relationships of macaques as inferred from restriction endonuclease analysis of mitochondrial DNA. Folia Primatol. 60, 7–17.

Zhang, Y., & Ryder, O.A. 1998. Mitochondrial Cytochrome b gene sequences of Langurs: Evolutionary inference and conservation relevance. In (Jablonski, N.G, Ed.) The Natural History of the Doucs and Snub- Nosed Monkeys, pp. 65-78. Singapore: World Scientific.

Zhilman, A.L., Cronin, J.E., Cramer, D.L. & Sarich, V. 1978. Pygmy chimpanzee as a possible prototype for the common ancestor of humans, chimpanzees and gorillas. Nature 275, 744-746.

Zhilman, A.L. & Cramer, D.L. 1978. Skeletal differences between pygmy (Pan paniscus) and common chimpanzees (Pan troglodytes). Folia primatol. 29, 86-94.

Zhilman, A.L. & Lowenstein, J.M. 1983. Ramapithecus and Pan paniscus: significance for human origins. In (R.L. Ciochon & R.S. Corruccini, Eds.) New Interpretations of Ape and Human Ancestry, pp. 677-694. New York: Plenum Press.

488 Ziegler, T., Hodges, K., Winkler, P. & Heistermann, M. 2000. Hormonal correlates of reproductive seasonality in wild female Hanuman langurs. AJP 51, 119-134.

Ziegler, T., Abegg, C., Meijaard, E., Perwitasari-Farajallah, D., Walter, L., Hodges, J.K. & Roos, C. 2007. Molecular phylogeny and evolutionary history of the Southeast Asian macaques forming the M. silenus group. MPE 42, 807-816.

Zischler, H., Geisert, H., von Haeseler, A. & Paabo, S. 1995. A nuclear ‘fossil’ of the mitochondrial D-loop and the origin of modern humans. Nature 378, 489-492.

Zollikofer, C.P.E., Ponce de Leon, M.S., Lieberman, D.E., Guy, F., Pilbeam, D., Likius, A., Mackaye, H.T., Vignaud, P. & Brunet, M. 2005. Virtual cranial reconstruction of Sahelanthropus tchadensis. Nature 434, 755-759.

Zuckerman, S., Ashton, E.H., Flinn, R.M., Oxnard, C.E. & Spence, T.F. 1973. Some locomotor features of the pelvic girdle in primates. Symp. Zool. Soc. Lond. 33, 71-165.

Abbreviations: AJPA: American Journal of Physical Anthropology AJP: American Journal of Primatology EA: Evolutionary Anthropology JHE: Journal of Human Evolution IJP: International Journal of Primatology MPE: Molecular Phylogenetics and Evolution PNAS: Proceedings of the National Academy of Sciences, USA TREE: Trends in Ecology and Evolution

489 Appendix:

Hominidae (Gray, 1825) “Fam. 1. HOMINIDÆ. Cutting teeth four above and below; grinders 5-5 above and below; nostrils separated by narrow septum. †Tail none. 1. Hominina, Homo. 2. Simiina. Troglodytes, Geoff. Simia, Lin. Hylobates, Illiger. ††Tail long or short. 3. Presbytina. Presbytes, Eschy. 4. Cercopithecia. Lasiopyga, Illig. Cercopithecus, Lin. Cercocebus, Geoff. Macacus. 5. Cynocepalina. Cynocephalus, Brisson. Papio, Brisson” (p. 338).

Hominidae (Campbell, 1962) “The Hominidae are characterized by a combination of the following main evolutionary trends: 1) Bipedalism, together with associated skeletal changes. 2) Changes in the position of the head on the vertebral column resulting from the erect posture, and associated modifications of the face, neck and jaws. 3) Reduction and modification of the teeth. Hands and tools replaced the use of teeth for many tasks. 4) Fœtalization: that is, the retardation of both ontogenetic development and the onset of sexual maturity, which have resulted in a prolonged juvenile learning phase and greater plasticity of behavior. 5) Lack of specialization, making possible the exploitation of a variety of environments. In particular the dentition and digestion are omnivorous, and the hand versatile, and prehensile. 6) The development of social life, tool-making and speech. 7) The increase in the absolute and relative size of the brain, and in particular the frontal region” (p. 225).

490 Hominidae (Le Gros Clark, 1964) “[A] subsidiary radiation of the Hominoidea distinguished from the Pongidae1 by the following evolutionary trends; progressive skeletal modifications in adaptation to erect bipedalism, shown particularly in a proportionate lengthening of the lower extremity, and changes in the proportions and morphological details of the pelvis, femur , and pedal skeleton related to mechanical requirements of erect posture and gait and to the muscular development associated therewith; preservation of well developed pollex; ultimate loss of opposability of hallux; increasing flexion of Basicranial axis associated with increasing cranial height; relative displacement forward of the occipital condyles; restriction of the nuchal area of occipital squama, associated with low position of inion; consistent and early ontogenetic development of a pyramidal mastoid process; reduction of subnasal prognathism, with ultimate early disappearance (by fusion) of facial component of premaxilla; diminution of canines to a spatulate form, interlocking slightly or not at all and showing no pronounced sexual dimorphism; disappearance of diastema; replacement of sectorial first lower premolars by bicuspid teeth (with later secondary reduction of lingual cusp); alteration in occlusal relationships, so that all the teeth tend to become worn down to a relatively flat even surface at an early stage of attrition; development of an evenly rounded dental arcade; marked tendency in later stages of evolution to a reduction in size of the molar teeth; progressive acceleration in the replacement of deciduous teeth in relation to the eruption of permanent molars; progressive “molarization” of the first deciduous molar; marked and rapid expansion (in some of the terminal products of the hominid sequence of evolution) of the cranial capacity, associated with reduction in size of jaws and area of attachment of masticatory muscles and the development of a mental eminence” (pp. 119-120).

1 “Family Pongidae-a subsidiary radiation of the Hominoidea distinguished from the Hominidae by the following evolutionary trends: progressive skeletal modifications in adaptations to arboreal brachiation, shown particularly in a proportionate lengthening of the upper extremity as a whole and of its different segments; acquisition of a strong opposable hallux and modification of morphological details of limb bones for increased mobility and for muscular developments related to brachiation; tendency to relative reduction of pollex; pelvis retaining the main proportions characteristic of quadrupedal mammals; marked prognathism, with late retention of facial component of premaxilla and sloping symphysis; development (in larger species) of massive jaws associated with strong muscular ridges on the skull; nuchal area of the occiput becoming extensive, with relatively high position of inion; occipital condyles retaining a backward position well behind the level of the auditory apertures; only a limited degree of flexion of basicranial axis associated with maintenance of low cranial height; cranial capacity showing no marked tendency to expansion; progressive hypertrophy of incisors with widening of symphyseal region of the mandible and ultimate formation of “simian shelf”; enlargement of strong conical canines interlocking in diastema and showing distinct sexual dimorphism; accentuated sectorialization of first lower premolar with development of strong anterior root; postcanine teeth preserving a parallel or slightly forward divergent alignment in relatively straight rows; deciduous molar retaining a predominantly unicuspid form; no acceleration in eruption of permanent canine” (pp. 120-121).

491 Hominidae (Pilbeam, 1968): “It is generally agreed that there are two principal sets criteria which must be satisfied if a fossil primate is to be considered a hominid. The requirements are, first, evidence of habitual upright bipedalism as the chief method of locomotion, and, second, the presence of teeth which are essentially human in form. It has become almost a matter of anthropological dogma that the earliest hominids differed from there pongid ancestors because of subtle locomotor changes in the direction of bipedalism. Hominids are also characterized dentally by a particular type of occlusal pattern in the cheek teeth, vertically implanted incisors and small non-projecting canines morphologically resembling the incisors. The small canines are, at least in part, responsible for the fact that hominid teeth are arranged in a parabolic dental arcade. In apes the incisors are large and procumbent, and – particularly in males – the canines are projecting, pointed tusks. The cheek teeth rows are parallel. The basic differences between dentitions of man and apes are in canine size and in the fact that both male and female hominids have small canines. The second of anthropology’s central dogmas is that reduction in hominid male canine size is correlated with the use of weapons (tools) in intra and inter-group feuding and in fighting between species. Clearly, then, these morphological complexes, associated with bipedalism and changes in tooth function, reflect marked shifts in behavior between pongids and hominids” (pp. 1335-1336).

492 Sahelanthropus tchadensis Brunet et al, 2002 “Cranium with an orthognathic face showing weak subnasal prognathism, a small ape- size braincase, a long and narrow basicranium, and characterized by the following morphology: the upper part of the face wide relative to a mediolaterally narrow and anteroposteriorly short lower face; a large canine fossa; a small and narrow U-shaped dental arch; orbits separated by a very wide interorbital pillar and crowned with a large, thick and continuous supraorbital torus; a flat frontal squama with no supratoral sulcus but with a marked postorbital constriction; a small, posteriorly located sagittal crest and a large nuchal crest (at least, in presumed males); a flat and relatively long nuchal plane with a large external occipital crest; a large mastoid process; small occipital condyles; a short, anteriorly narrow basioccipital; the long axis of the petrous temporal bone oriented roughly 30° relative to the sagittal plane; the biporion line touching the basion; a round external auditory porus; a broad glenoid cavity with a large postglenoid process; a robust and superoinferiorly short mandibular corpus associated with a wide extramolar sulcus; a large, anteriorly opening mental foramen centred beneath lower teeth P4–M1, below midcorpus height; relatively small incisors; distinct marginal ridges and multiple tubercles on the lingual fossa of upper I1; small (presumed male) upper canines longer mesiodistally than buccolingually; upper and lower canines with extensive apical wear; no lower c–P3 diastema; upper and lower premolars with two roots; molars with low 3 rounded cusps and bulbous lingual faces, M triangular and M3 rounded distally; enamel thickness of cheek teeth intermediate between Pan and Australopithecus. Sahelanthropus is distinct from all living great apes in the following respects: relatively smaller canines with apical wear, the lower showing a full occlusion above the well-developed distal tubercle, probably correlated with a non-honing C–P3 complex (P3 still unknown). Sahelanthropus is distinguished as a hominid from large living and known fossil hominoid genera in the following respects: from Pongo by a non-concave lateral facial profile, a wider interorbital pillar, superoinferiorly short subnasal height, an anteroposteriorly short face, robust supraorbital morphology; from Gorilla by smaller size, a narrower and less prognathic lower face, no supratoral sulcus, and smaller canines and lower-cusped cheek teeth; from Pan by an anteroposteriorly shorter face, a thicker and more continuous supraorbital torus with no supratoral sulcus, a relatively longer braincase and narrower basicranium with a flat nuchal plane and a large external occipital crest, and cheek teeth with thicker enamel; from Samburupithecus by a more anteriorly and higher-placed zygomatic process of the maxilla, smaller cheek teeth with lower cusps and without lingual cingula, and smaller upper premolars and M3; from Ouranopithecus by smaller size, a superoinferiorly, anteroposteriorly and mediolaterally shorter face, relatively thicker continuous supraorbital torus, markedly smaller but mesiodistally longer canines, apical wear and large distal tubercle in lower canines, and thinner postcanine enamel; from Sivapithecus by a superoinferiorly and anteroposteriorly shorter face with non-concave lateral profile, a wider interorbital pillar, smaller canines with apical wear, and thinner cheek-teeth enamel; from Dryopithecus by a less prognathic lower face with a wider interorbital pillar, larger supraorbital torus, and thicker postcanine enamel. Sahelanthropus is also distinct from all known hominid genera in the following respects: from Homo by a small endocranial capacity (preliminary estimated range 320–380 cm3) associated with a long flat nuchal plane, a longer truncated triangle- shaped basioccipital, a flat frontal squama behind a robust continuous and undivided supraorbital torus, a large central upper incisor, and non-incisiform canines; from Paranthropus by a convex facial profile that is less mediolaterally wide with a much smaller malar region, no frontal trigone, the frontal squama with no hollow posterior to glabella, a smaller, longer and narrower braincase, the zygomatic process of the maxilla positioned more posterior relative to the tooth row, and markedly smaller cheek teeth; from Australopithecus by a less prognathic lower face (nasospinale–prosthion length shorter at least in presumed males) with a smaller malar (infraorbital) region and a larger, more continuous supraorbital torus, a relatively more elongate braincase, a relatively long, flat nuchal plane with a large external occipital crest, non-incisiform and mesiodistally long canines, and thinner cheek-teeth enamel; from Kenyanthropus by a

493 narrower, more convex face, and a narrower braincase with more marked postorbital constriction and a larger nuchal crest; from Ardipithecus by upper I1 with distinctive lingual topography characterized by extensive development of the crests and cingulum; less incisiform upper canines not diamond shaped with a low distal shoulder and a mesiodistal long axis, bucco-lingually narrower lower canines with stronger distal 1 tubercle, and P4 with two roots; from Orrorin by upper I with multiple tubercles on the lingual fossa, and non-chimp-like upper canines with extensive apical wear” (pp. 145-146). Orrorin tugenensis Senut et al, 2002 “Jugal teeth smaller than australopithecines; large upper central incisor with thick enamel; mandibular corpus relatively deep below M3; thick enamel on lower cheek teeth; femur with spherical head rotated anteriorly, neck elongated and oval in section; lesser trochanter medially salient; humerus with vertical brachioradialis crest; proximal manual phalanx curved; dentition small relative to body size” (p. 139).

Ardipithecus ramidus White et al, 1995 “Less postcanine megadontia, with upper and lower canines larger relative to the postcanine teeth; lower first deciduous molar narrow and obliquely elongate, with large protoconid, small and distally placed metaconid, no anterior fovea, and small, low talonid with minimal cuspule development; temporomandibular joint without definable articular eminence, absolutely and relatively thinner canine and molar enamel; lower third premolar more strongly asymmetrical, with dominant, tall buccal cusp and steep, posterolingually directed transverse crest; upper third premolar more strongly asymmetric, with relatively larger, taller, more dominant buccal cusp” (p. 88).

Ardipithecus kadabba Haile-Selassie et al, 2004 “A. kadabba differs from fossil and extant apes by features enumerated in [Haile-Selassie, 2001] and by the presence of a clearly defined anterior fovea of the lower P3, demarcated by a foldlike buccal segment of the mesial marginal ridge. It differs from extant apes and O. tugenensis by its more circular upper canine crown outline in occlusal view. In addition to the features distinguishing A. r. ramidus from A. r. kadabba, A. kadabba also differs from A. ramidus by the morphology of the upper canine (more basal termination of the mesial and distal apical crests) and morphology of the lower P3 (more asymmetrical crown outline and relatively smaller anterior fovea)” (p. 1504).

494 Australopithecus Dart, 1925 (Le Gros Clark, 1964) “[A] genus of the hominidae distinguished by the following characters: relatively small cranial capacity, ranging from about 450 to well over 600 cc.; strongly built supra-orbital ridges; a tendency in individuals of larger varieties for the formation of a low sagittal crest in the frontoparietal region of the vertex of the skull (but not associated with a high nuchal crest); occipital condyles well behind the mid-point of the cranial length but on a transverse level with the auditory apertures; nuchal area occiput restricted, as in Homo; consistent development (in immature as well as mature skulls) of a pyramidal mastoid process of typical hominid form and relationships; mandibular fossa constructed on the hominid pattern but in some individuals showing a pronounced development of the postglenoid process; massive jaws, showing considerable individual variation in respect of absolute size; mental eminence absent or slightly indicated; symphyseal surface relatively straight and approaching the vertical; dental arcade parabolic in form with no diastema; spatulate canines wearing down flat from the tip only; relatively large premolars and molars; anterior lower premolar bicuspid with subequal cusps; pronounced molarization of first deciduous molar; progressive increase in size of permanent lower molars from first to third; the limb skeleton (so far as it is known) conforming in its main features to the hominid type but differing from Homo in a number of details, such as the forward prolongation of the region of the anterior superior spine of the ilium and a relatively small sacroiliac surface, the relatively low position (in some individuals) of the ischial tuberosity, and the marked forward prolongation of the intercondylar notch of the femur” (Le Gros Clark, 1964, p. 168).

Australopithecus Dart, 1925 (Tobias, 1967) “A genus of the Hominidae distinguished by the following characters: relatively small cranial capacity, with an average of about 500 c.c. and an estimated population range of about 360 to about 640 c.c.; a relatively thin-walled cranium rendered robust in parts by strong ectocranial superstructures and by marked pneumatization; strongly-built supraorbital ridges; moderate to fairly high orbits, with a lower mean height than in pongids; a tendency in individuals with larger cheek-teeth for the formation of a low sagittal crest in the frontoparietal region of the calvaria (but the sagittal crest is not continuous with either the nuchal crest or the occipital torus, whichever is present); occipital condyles well anteroposterior midpoint of the cranial length, but in the same coronal plane as the external acoustic apertures; foramen magnum well forward on the base of the cranium; planum nuchale of occipital bone rising only a short distance above the F.H. and generally facing downwards much more then backwards; inion low and generally close to the Frankfurt plane; a low nuchal crest in heavier-toothed forms, and a slight occipital torus in moderate-toothed forms; consistent development (in immature and mature crania) of a pyramidal mastoid process of typical hominine form and relationships; a mandibular fossa which is shallow and mediolaterally broad , but is otherwise constructed on the hominid pattern, especially slopes and curvature of the anterior wall and the upward slope of the preglenoid plane, but with a pronounced entoglenoid process and, in some individuals, a moderate development of the postglenoid process; porion elevated in position above the nasion-opisthion line; massive and robust jaws, showing marked individual variation in respect of absolute size; mental eminence absent or slightly indicated; symphyseal surface relatively straight and retreating; contour of internal mandibular arch V-shaped or blunt U-shaped; dental arcade parabolic in form with no diastema; moderate-sized, spatulate canines wearing down flat from the tip only; relatively large premolars and molars, the enlargement being no more marked in the bucco-lingual diameter of the crown; lower anterior premolar bicuspid with subequal cusps; pronounced molarization of the lower first deciduous molar; progressive increase in size of permanent lower molars from first to third, but M3 is commonly smaller than M2; the limb skeleton (so far as it is known) conforming in its main features to the hominid type but differing from Homo in a number of details, such as the forward

495 prolongation of the region of the anterior superior iliac spine and a relatively small sacroiliac surface, the relatively low position (in some individuals) of the ischial tuberosity, and the marked forward prolongation of the intercondylar notch of the femur” (p. 234).

Australopithecus Dart, 1925 (Wolpoff & Lovejoy, 1975) “Large cranial capacity, averaging about 500 cm3 (range from 400 to 800 cm3); heavily pneumatized cranial base; weakly to moderately developed supraorbital torus; extremely well developed muscles of mastication frequently leading to the development of a low sagittal crest generally beginning anterior to or at bregma in males or more robust individuals, widely flared zygomatics and large temporal fossae, an anterior position of the zygomatic process of the maxilla with the anterior root of the zygoma arising between C and M1 and a tendency for the appearance of a frontal trigone; massive jaws, especially in robustness of the corpus, but varying considerably in absolute size; a vertical ascending ramus and symphyseal region; superior and inferior transverse tori present with a horizontally elongated post incisive plane. Massive postcanine dentition relative to body size with a tendency towards multiple or accessory cuspation in both deciduous and permanent posterior teeth (dm1 consistently molarized). Moderately large anterior dentition with distinct sexual dimorphism in canine size; progressive molar size increase from M1 to M3; large and prognathic face; endocasts distinguished from pongids by a posterior placement of the lunate sulcus; pelvic birth canal more restricted than Homo, with commensurate lateral iliac flare and elongated femoral neck” (pp. 275-276).

Australopithecus Dart, 1925 (Groves, 1989) “A genus of the Hominini with the following states synapomorphous with Homo: mesial buccal groove more conspicuous than distal on P3, equally developed on P4; hypoconulids reduced on lower molars; nasals raised from plane of face; upper end of nasal aperture level with inferior orbital margin; a sill separating naso-alveolar clivus from nasal floor; transverse buttress reduced; face height less than 75 per cent of the upper face breadth; foramen ovale posterior to margin of pterygoid plate; digastric scar in notch on basicranium; Biporionic breadth greater than biparietal; frontal angle greater than 35o; femoral distal epiphysis index greater than 75; femoral intercondylar notch wider than high; femoral condyles symmetrical; tibial spine index greater than 50; humero-femoral index less than 75; intertrochanteric line present; femoral shaft diameter less than 60 per cent of head diameter. And with the following autapomorphic states: I1 crown area more than 169 percent that of I2; transverse facial buttress absent; zygomatic prominence bulbous (convergent with Paranthropus); occipital index greater than 70; femoral platymeria reduced; pelvic torsion apparently increased. This genus shares so many synapomorphies with Homo that it becomes almost a matter of taste whether to keep it separate or to combine the two” (pp. 256-257).

496 Kenyanthropus platyops Leakey et al, 1995 “Transverse facial contour flat at a level just below the nasal bones; tall malar region; zygomaticoalveolar crest low and curved; anterior surface of the maxillary zygomatic process positioned over premolars and more vertically orientated than the nasal aperture and nasoalveolar clivus; nasoalveolar clivus long and both transversely and sagittally flat, without marked juga; moderate subnasal prognathism; incisor alveoli parallel with, and only just anterior to, the bicanine line; nasal cavity entrance stepped; palate roof thin and flexed inferiorly anterior to the incisive foramen; upper incisor (I1 and I2) roots near equal in size; upper premolars (P3, P4) mostly three-rooted; upper first and second molars (M1 and M2) small with thick enamel; tympanic element mediolaterally long and lacking a petrous crest; external acoustic porus small. Kenyanthropus can be distinguished from Ardipithecus ramidus by its buccolingually narrow M2, thick molar enamel, and a temporal bone with a more cylindrical articular eminence and deeper mandibular fossa. It differs from A. anamensis, A. afarensis, A. africanus and A. garhi in the derived morphology of the lower face, particularly the moderate subnasal prognathism, sagittally and transversely flat nasoalveolar clivus, anteriorly positioned maxillary zygomatic process, similarly sized I1 and I2 roots, and small M1 and M2 crowns. From A. afarensis it also differs by a transversely flat midface, a small, external acoustic porus, and the absence of an occipital/marginal venous sinus system, and from A. africanus by a tall malar region, a low and curved zygomaticoalveolar crest, a narrow nasal aperture, the absence of anterior facial pillars, a tubular, long and crestless tympanic element, and a small, external acoustic porus. Kenyanthropus lacks the suite of derived dental and cranial features found in Paranthropus aethiopicus, P. boisei and P. robustus, and the derived cranial features of species indisputably assigned to Homo (For example, H. erectus s.l. and H. sapiens, but not H. rudolfensis and H. habilis)” (p. 433).

497 Paranthropus Broom, 1938 “The skull is that of a large ape, larger than most male chimpanzees and nearly as large as most female gorillas; but it differs very greatly from both the living African anthropoids. […] The glenoid cavity and its relations to the tympanic bone are of exceptional interest. In the gorilla, the chimpanzee, the orang and the gibbon, the outer part of the tympanic is situated behind the posterior glenoid process. In man, the tympanic is situated mainly below the glenoid process, and even at its out part it forms the posterior nonarticular part of the glenoid cavity. In the new fossil ape, the condition of the glenoid and tympanic is almost exactly as in man, though the parts are very much larger. The occipital condyle is in practically the same plane as the external auditory meatus and thus farther forward than in the gorilla and the chimpanzee; which appears to indicate that the ape walked somewhat more erect than the living anthropoids. From the portion of the brain case preserved, I estimate the volume of the brain to have been about 600 c.c. The face is remarkably flat and much shorter than in the gorilla. A curious bony ridge runs down from the inner border of the large infraorbital foramen. The molar teeth […] differ considerably in shape from those in Plesianthropus transvaalensis [Au. africanus), and the 2nd premolar is about half as large again as in the Sterkfontein ape. The upper canine had been relatively remarkably small, and the incisors, of which we have much of the sockets preserved, were also relatively small. The palate is relatively short and broad, and owing to the small size of the incisors and canines the anterior part is narrowed, and the teeth are arranged more as in man than in any of the living anthropoids. […] The incisors which are lost have been relatively very small, and the lateral ones are scarcely larger than the central. […] It is clearly very unlike the canine of Plesianthropus transvaalensis. The premolars have rounded crowns without any high well developed cusps as in the living anthropoids, and are thus fairly similar to those of man, but twice as large. The 2nd premolar differs very markedly from that of Plesianthropus transvaalensis, and may thus confidently place the new skull in a new genus and species” (p. 378).

Paranthropus Broom, 1938 (Robinson, 1972) “The Subfamily Paranthropinae is defined as consisting of a group of higher primates having cheek teeth with proportionately large occlusal surfaces of low relief associated with anterior teeth, especially canines, which are proportionately small to extremely small; the canines wear in a manner that the major wear facet either starts on the apex or soon incorporates the apex; the mandible is very robust with proportionately tall and more or less vertical ascending ramus; the face is robust and flat with prominent cheek- bones so that the face is commonly dished; the frontal are is narrow with no more slight convexity so that no forehead exists, and the vertex is relatively low as in pongids; a sagittal crest in the region of the vertex is usual; some evidence of adaptation to erect posture (i.e. more than in pongids), but adaptation is incomplete and does not include a proportionately shortened Ischium and lengthened lower limb – the propulsive mechanism is thus, as in apes, essentially power-oriented” (pp. 6-7)2.

2 Please note, this definition, “includes Zinjanthropus, Meganthropus from Java, and Paraustralopithecus” (p. 6). In addition, Gigantopithecus is also included within the Paranthropine subfamily.

498 Paranthropus Broom, 1938 (Groves, 1989) “A genus of the Hominini with the following derived character states: P4 (in both jaws) expanded, generally broader than either P3 or M1; canines small; DM1, 2 protoconid not mesial to metaconid; DM1 talonid enlarged; DM1 and M1 buccal groove deep, ending in a 1 pit; DM1 fovea anterior reduced to a fissure; M buccal groove very deep: I1lingual marginal ridges reduced; I2 incisive edge without distal slope; lingual relief of canines lost; canines symmetrical, not projecting beyond occlusal line, their tips rounded not pointed; lower molars oval, their cusps centrally placed; Carabelli complex and protostyle 3 reduced or absent; P rounded, its crown not tapering towards occlusal surface; P4 buccal grooves lost; premolar root type B occurs; M1 erupts nearly coevally with I1; pre-maxilla anteriorly straight; curve of Spee enhanced; nasion at glabella; nasals broadened superiorly; vomer inserts on nasal spine; facial buttressing pronounced; petrous axis at 40-55o to bicarotid line; tympanic plate very long; occipito-mastoid crest lost; occipital marginal venous sinus developed; humerus with lateral epicondyles salient. Two of the above characters (relative eruption times of M1 and I1, and petrous axis inclination) converge with Homo” (p. 252).

499 Homo Linneaus, 1758 The major defining feature of Homo sapiens was summed up with the aphorism, ‘Nofce Te ipfum’ (or ‘Nosce Te ipsum’) -‘know thyself’. Other characteristics included diurnal habits and widespread geographic range.

Homo Linneaus, 1758 (Leakey et al, 1964) “A genus of the Hominidae3 with the following characters: the structure of the pelvic girdle and of the hind-limb skeleton is adapted to habitual erect posture and bipedal gait; the fore-limb is shorter than the hind-limb; the pollex is well developed and fully opposable and the hand is capable not only of a power grip but of, at the least, a simple and usually well developed precision grip; the cranial capacity is very variable but is, on average, larger than the range of capacities of the members of the genus Australopithecus, although the lower part of the range of capacities in the genus Homo overlaps with the upper part of the range in Australopithecus; the capacity is (on the average) large relative to body size and ranges from about 600 cc to more than 1600 cc; the muscular ridges on the cranium range from very strongly marked to virtually imperceptible, but the temporal crests or lines never reach the mid-line; the frontal region is without undue post-orbital constriction (such as is common in members of the genus Australopithecus); the supraorbital region of the frontal bone is very variable, ranging from a massive and very salient supra-orbital torus to a complete lack of any supra-orbital projection and a smooth brow region; the facial skeleton varies from moderately prognathous to orthognathous, but is not concave (or dished) as is common in members of the Australopithecinae; the anterior symphyseal contour varies from a marked retreat to a forward slope, while the bony chin may be entirely lacking, or may vary from a slight to a very strongly developed mental trigone; the dental arcade is evenly rounded with no diastema in most members of the genus; the first lower premolar is clearly bicuspid with variably developed lingual cusp; the molar teeth are variable in size, but are in general are small relative to the size of these teeth in the genus Australopithecus; the size of the last upper molar is highly variable, but is generally smaller than second upper molar and commonly also smaller than the first upper molar; the lower third molar is sometimes appreciably larger than the second; in relation to the position in the Hominoidea as a whole, the canines are small, with little or no overlapping after the initial stages of wear, but when compared with those members of the genus Australopithecus, the incisors and canines are not very small relative to the molars and premolars; the teeth in general, and particularly the molars and premolars, are not enlarged bucco-lingually as they are in the genus Australopithecus; the first deciduous lower molars shows a variable degree of molarization” (pp. 7-8).

Homo Linnaeus, 1758 (Groves, 1989) “A genus of the Hominini sharing with Australopithecus numerous synapomorphous states, as listed above, under that genus; and with the following autapomorphic states: P4 4 shapes index below 96; PP shape index above 75; vomer inserts well-behind inferior nasal spine; mandibular fossa deepened, with well-developed articular eminence; cranial vault bones thickened; foramen magnum less back-sloping (convergent with Paranthropus); femoral neck index above 75. In addition, one species (the unnamed Hadar species) is relatively poorly known; the following features are synapomorphic for the remainder of the genus, and may or may not turn out to be applicable to the Hadar species, and so synapomorphic for the genus as a whole: M11 erupts at about the same time as I , alveolar projection reduced, petrous axis 40-55o (these states converge upon Paranthropus); maxillo-alveolar index above 100, palate broader than long; frontal process of maxilla faces laterally; post-orbital constriction reduced; Bizygomatic breadth less than 130% of biorbital; face height less than biorbital breadth” (p. 260).

3 Following the definition of Le Gros Clark, 1964.

500 Homo Linnaeus, 1758 (Wood, 1992) “The resulting Homo clade is defined by the following character state changes at node A4. 1. Increased cranial vault thickness. 2. Reduced post-orbital constriction. 3. Increased contribution of the occipital bone to cranial sagittal arc length. 4. Increased cranial vault height. 5. More anteriorly-situated foramen magnum. 6. Reduced lower facial prognathism. 7. Narrower tooth crowns, particularly mandibular premolars. 8. Reduction in length of the molar tooth row. The two species comprising the original H. habilis hypodigm, H. habilis sensu stricto and H. rudolfensis share a hypothetical ancestor with each other which neither shares `with any other taxon. That sister group is defined by the five character state changes at node B5. 1. Elongated anterior basicranium. 2. Higher cranial vault. 3, 4. Mesiodistally-elongated M1 and M2. 5. Narrow mandibular fossa” (p. 789).

Homo Linnaeus, 1758 (Wood & Collard, 1999a) “We suggest that a fossil species should be included in Homo only if it can be demonstrated that it (i) is more closely related to H. sapiens than it is to the australopiths, (ii) has an estimated body mass that is more similar to that of H. sapiens than to that of the australopiths, (iii) has reconstructed body proportions that match those of H. sapiens more closely than those of the australopiths, (iv) has a postcranial skeleton whose functional morphology is consistent with modern human-like obligate bipedalism and limited facility for climbing, (v) is equipped with teeth and jaws that are more similar in terms of relative size to those of modern humans than to those of the australopiths, and (vi) shows evidence for a modern human-like extended period of growth and development” (1999a. pp.70-71).

Homo Linnaeus, 1758 (Dunsworth & Walker, 2002) In a recent volume (Hartwig, 2002) documenting the entire Primate fossil record, this definition was offered: “A genus of the family Hominidae. Cranial: highly variable cranial capacity, on average, larger than that of Australopithecus, cranial capacity large relative to body size; reduced postorbital constriction; reduced facial prognathism; more anteriorly situated foramen magnum; increased participation of the occipital bone in cranial sagittal arc length; increased cranial vault height. Dental: narrower tooth crowns, particularly mandibular premolars; reduction in length of the molar toothrow; variable molar size but smaller than Australopithecus; in general, teeth are not enlarged buccolingually as in Australopithecus” (p. 427).

4 The node in Wood’s cladogram representing the last common ancestor to H. habilis, H. rudolfensis, H. erectus, H. ergaster and H. sapiens. 5 The node in Wood’s cladogram representing the last common ancestor to only H. habilis and H. rudolfensis. Thus, in this analysis, the remaining characters listed are not pertinent or necessary for the other species of Homo.

501 Pan Oken, 1816 (Hill, 1969b) “Distinguishing characters of the genus are: Stature (vertex-heel) not exceeding 1.7m but body less bulky than in Gorilla and Pongo. Sexual dimorphism in size also less than in Gorilla and Pongo. Pelage coarse, composed mainly of rigid, glossy black hairs, rather sparsely planted, especially on the ventral regions; hairs on the forearms directed towards elbows; face highly prognathous, truncated in front, with highly developed, prominent supraorbital ridges continuous across the midline behind which the forehead recedes at a low vaulted cranium; external ears large, outstanding, not simplified. External nose not raised above the level of the face; margins of nares not markedly expanded. Pectoral limbs long, reaching below the knee, but shorter than in Pongo (orangutan); legs short, but longer than in Pongo; hands slender, longer than foot; manual digits (except pollex, which is short) long, slender, curved; digits II-V of foot not united cutaneously to distal ends of proximal phalanges. Penis long, narrow, laterally compressed without marked terminal glans, meatal orifice small; females with catamenial swelling [Pocock, 1926]. Skull with depressed calva, lacking a sagittal crest or with but a short, low crest posteriorly. Orbits quadrangular with rounded angles; maxillary region prognathous. Nasals reduced, parallel sided. Temporo-frontal contact at pterion. Mastoid area roughened but mastoid process developed. Canines larger in males than females, but not as hypertrophied as in Gorilla. Upper premolar crowns medio-distally short, with buccal cusps higher lingual. Molar 3 smaller by one third than molar 1 or molar 2, with indistinct lingual cingulum. Ribs 13 pairs (average 13.2 compared with 12.86 in Gorilla). Os centrale carpi lacking. Brain absolutely smaller, but relatively larger than in Gorilla, less flattened in frontal region, but not so high as in Pongo. A true inferior frontal sulcus usually present and a vertical diagonal sulcus budded off with it from the s. praecentralis inferior. Branching of parallel sulcus less developed than in Pongo or Gorilla indicative of lesser differentiation in posterior parietal region – correlated with relatively greater lateral extension of occipital operculum [Connolly, 1950]. Diploid chromosome number 48 with probable XX-XY sex chromosome constitution [Young et al, 1960]. The genus is represented by two species, P. troglodytes and P. paniscus, distinguished primarily in size, the latter being paedomorphic dwarf which is further distinguished by its lighter build” (pp. 30-31).

502