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The Fossil Record and Evolution of Bovidae: State of the Field Faysal

The Fossil Record and Evolution of Bovidae: State of the Field Faysal

Palaeontologia Electronica http://palaeo-electronica.org

The Record and Evolution of : State of the Field

Faysal Bibi, Maia Bukhsianidze, Alan W. Gentry, Denis Geraads, Dimitris S. Kostopoulos, Elisabeth S. Vrba

ABSTRACT

This paper presents a summary of the current state of knowledge on the evolution and fossil record of Bovidae and identifies topics and problems for further study. The earliest records of the different bovid tribes are reviewed, along with their geographic locations. Future work on the fossil bovid record should focus on: the reinvestigation of osteological characters that might distinguish early bovids from stem taxa; the late Mio- cene fossil record with respect to the evolutionary origins and biogeography of the liv- ing bovid tribes; and improved methodologies for reconstructing bovid phylogeny using morphology. This paper reflects the main themes discussed at the meeting of the Revealing Hominid Origins Initiative Bovid Analytical Working Group in Addis Ababa, .

Faysal Bibi. Department of Geology and Geophysics, Yale University, PO Box 208109, Connecticut, 06520-8109, USA. [email protected] Current address: Institut international de Paléoprimatologie, Paléontologie humaine, Evolution et Paléoenvironnements, Université de Poitiers, F-86022, POITIERS cedex Maia Bukhsianidze. The National Museum of Georgia, 3 Purtseladze St., 0107 Tbilisi, Georgia. [email protected] Alan W. Gentry. c/o Department of Palaeontology, Natural History Museum, Cromwell Road, London, SW7 5BD, United Kingdom. [email protected] Denis Geraads. CNRS UPR 2147, 44 rue de l’Amiral Mouchez, 75014 Paris, France. [email protected] Dimitris S. Kostopoulos. Aristotle University of Thessaloniki, School of Geology, Laboratory of Geology and , 54124 Thessaloniki, . [email protected] Elisabeth S. Vrba. Department of Geology and Geophysics, Yale University, PO Box 208109, Connecticut, 06520-8109, USA.

KEY WORDS: Bovidae; Biogeography; Phylogeny; Systematics; , late

PE Article Number: 12.3.10A Copyright: Society for Paleontology December 2009 Submission: 15 May 2009. Acceptance: 9 August 2009

Bibi, Faysal, Bukhsianidze, Maia, Gentry, Alan W., Geraads, Denis, Kostopoulos, Dimitris S., and Vrba, Elisabeth S., 2009. The Fossil Record and Evolution of Bovidae: State of the Field. Palaeontologia Electronica Vol. 12, Issue 3; 10A: 11p; http://palaeo-electronica.org/2009_3/169/index.html Bibi et al.: Fossil Bovids

INTRODUCTION France, MN4 in Gentry et al. 1999; ca.18–17Ma in Steininger 1999) has been proposed as a stem Bovids are diverse herbivorous that bovid (see also Azanza and Morales 1994), as has inhabit a wide spectrum of and occur natu- E. noyei (ca.18Ma, Pakistan, Solounias et al. rally over much of the globe. Ranging from the 1995). minus, from the early Miocene of enigmatic and elusive forest of South Pakistan, may be even older than E. noyei, and East and Central to the ubiquitous cat- possibly more primitive in morphology (Ginsburg et tle and other common to most world cul- al. 2001). tures, and from the Himalayan to the Other taxa worth further consideration in rela- to the North American , bovids represent tion to stem bovids include Andegameryx, Amphi- an example of a that owes its success to its moschus, Namibiomeryx, Namacerus, capacity to diversify in the face of selective pres- Pseudoeotragus, and Homoiodorcas. Taxa sures. The evolutionary history of Bovidae is included in Hypsodontinae (Köhler 1993) might be known to have taken place in the Miocene to diphyletic to (i.e., the of) the remainder Recent, or within the last 23 Ma (Figure 1). The of Bovidae (Gentry et al. 1999). Various Miocene Miocene fossil record is both rich and geographi- taxa have been referred to Hypsodontinae (Köhler cally extensive, and as a result the reconstruction 1987). The group is in need of revision, having of the evolutionary history of Bovidae promises to never been treated in much detail, and its phyloge- provide one of the most complete such histories for netic status remains unclear. any large group. Between August 4 and 8, 2008, the Bovid TRIBAL ORIGINS AND RADIATIONS Analytical Working Group of the Revealing Hom- inid Origins Initiative (National Science Foundation, Bovids have long been classified among a USA) met in Addis Ababa for a conference on the dozen or so higher taxa, typically tribes. Simpson’s fossil record and evolution of Bovidae with special (1945) classification divided bovids among 13 emphasis on the late Miocene to early . tribes in five subfamilies. Since that time, work on A summary of the main themes, findings, and rec- bovid systematics (morphological, behavioral, and ommendations of the conference follows. molecular) has revealed a deep bifurcation within Bovidae separating the from all the BOVID ORIGINS remaining bovids (Kingdon 1982; Lowenstein 1986), and has condensed the number of tribes to Recognition of the earliest fossil bovids is not around 10 (e.g., Hassanin and Douzery 1999b). straightforward, nor for that matter is recognition of early or stem members of the different bovid sub- Bovinae (++) . Bovidae is diagnosed by the presence of Bovinae is the sister group to all remaining liv- permanent and unbranched cores with a per- ing bovids and is characterized by numerous mor- manent and unbranched horn sheath (Janis and phological, behavioral, and molecular Scott 1988). Sheaths are rarely preserved as fos- synapomorphies (e.g., Kingdon 1982; Hassanin sils, so the recognition of an early bovid in the fossil and Douzery 1999b; Vrba and Schaller 2000; Bibi record relies on the presence and morphology of 2009). Eotragus sansaniensis (15.2Ma, France, the bony horn core alone. This proposition is prob- Sen 1996), E. noyei (ca.18Ma, Pakistan), and an lematic, however, given that members of other pec- unnamed from the Vihowa Formation oran groups also a very similar horn-core (ca.18Ma, Pakistan) have been tentatively pro- structure (both fossil and living species, e.g., posed as early or stem Bovinae (Solounias and Antilocapra americana) and given the potential Moelleken 1992; Azanza and Morales 1994; absence of horns in female individuals. Dental and Solounias et al. 1995). These would sug- other cranial and postcranial osteological charac- gest that the cladogenetic event separating the ters of potential use for distinguishing the earliest Bovinae from the remainder of Bovidae must have bovids and stem bovids should be further investi- occurred in the early Miocene, or certainly by gated. Teeth and cranial characters of certain spe- 18Ma. cies of Eotragus, long considered the earliest Stem Bovini were likely present in southern bovids, may indicate that certain of these species Asia (Siwaliks) by 9Ma, as represented by Seleno- are basal to (i.e., precede on the ) the portax (Bibi 2007). After this time, early bovines last common ancestor of all the living Bovidae. dispersed first to Africa then to Europe. It is not Eotragus artenensis (Ginsburg and Heintz 1968; clear when or where exactly the crown clade origi-

2 PALAEO-ELECTRONICA.ORG

R Bovini Tragelaphini Boselaphini Reduncini Aepyceros Alcelaphini Hippotragini Caprini e melampus n stoce i e Pl

19 e n oce li P

Tragoportacini 5Ma

18

14 15 16 17 e n 11 oce i 12 13 M

Late 10 9

8 10Ma

e 7 n oce Mi e

iddl Bovinae 6 M 5 15Ma 4

Antilopinae

3 Bovidae Crown Group 2 1 Miocene

y Stem Group

Earl Ghost Lineage

20Ma

FIGURE 1. Phylogenetic diagram charting the evolution of the major bovid clades. Solid numbered cir- cles represent fossils assignable to crown or stem groups, while open circles represent fossil taxa of potential phylogenetic affinity. Numbers refer to the following: 1, Eotragus artenensis, Eotragus noyei, Eotragus minus; 2, Eotragus noyei, Vihowa Bovidae indet.; 3, Pseudoeotragus seegrabensis; 4, Eotra- gus sansaniensis; 5, Caprotragoides, Gentrytragus, Tethytragus; 6, Fort Ternan Gazella sp.; 7, Ngorora ‘Pachytragus aff. solignaci’; 8, Pachytragus solignaci; 9, Djebel Krechem Hippotragini? indet.; 10, Seleno- portax; 11, Pheraios chryssomallos; 12, Siwalik Reduncini; 13, /Pachytragus; 14, Lothagam Reduncini; 15, Lothagam Aepyceros; 16, Lothagam Alcelaphini; 17, Toros Menalla & Lothagam Hip- potragini; 18, Lukeino cf. spekii; 19, Omo & Koobi Fora Aepyceros melampus. Bovid silhou- ettes adapted from Kingdon (1982).

3 Bibi et al.: Fossil Bovids

nated as the fossil taxon closest to the most recent referred to “Boselaphini.” Crown Boselaphini has a common ancestor of all living bovines has not yet poor fossil record, with a handful of Asian localities been identified. The earliest African record of a of imprecise Plio-Pleistocene age yielding fossil bovine is at Toros-Menalla (Vignaud et al. 2002), Boselaphus and Proboselaphus (Rütimeyer 1878; Chad, at around 7Ma. By the end of late Miocene, Matsumoto 1918; Pilgrim 1939). We know of no several western Eurasian “boselaphines” begin to fossil record for Tetracerus. The Miocene fossils exhibit morphology suggestive of the bovine condi- traditionally attributed to “Boselaphini” may be tion but the earliest European record of a bovine is divided among independent clades and stem in the early (Gromolard 1980). The ability groups of one or more living bovid tribes (Pilgrim to confirm with certainty a southern Asian origin of 1939; Moyà-Solà 1983; Thomas 1984b; Bibi 2007). Bovini is hampered by the poor African fossil The dominant late Miocene fossil “boselaphines,” record between 10 and 7Ma, whereby the absence both in terms of diversity and geographic range, of Bovini from Africa during this time may be a are species of the genera and result of preservation bias. Once Bovini had origi- . Interrelationships between Tragopor- nated, multiple bovine lineages become evident tax and Miotragocerus remain unclear, with diagno- but good hypotheses of between- and within-lin- ses so far proposed to distinguish the two genera eage relationships are lacking. The water being difficult to consistently apply in a wide con- most likely emerged from the Pliocene text. Species of these genera range throughout the Proamphibos (Pilgrim 1939) and the African Syn- late Miocene, and are found across from cerus from a Ugandax (Gentry and Gentry 1978) China to Spain, in the Siwaliks, and to the far south though the precise limits of the latter , along of South Africa. Tragoportax, Miotragocerus, and with that of Simatherium, are still unclear. The ori- other taxa such as , have been gins of and Bison and the interrelationships of united under the name Tragocerina or Tragocerini the many fossil species seemingly relevant to them (Sokolov 1953; Korotkevich 1981; Thomas 1984a, remain problematic. The hypothesis that 1984b). However, by the rules of the International gave rise to Bos (Martínez-Navarro et al. 2007) Code on Zoological Nomenclature (Article 39), a requires further evidence. -group taxon name may not be based on a The history of the Tragelaphini as known so genus found to be a junior homonym—in this case far is almost entirely African. The earliest tragela- Tragocerus, previously occupied for a beetle. As a phine that is probably a crown group member (i.e., result we adopt for this fossil group the name not a stem representative) is recognized from the Tragoportacini, appropriately based on Tragopor- Lukeino Formation (Thomas 1980), , dated tax—this being the genus in which we now place to between 6.56–5.72Ma (Deino et al. 2002). the old type species T. amalthea, of Trago- Tragelaphini are entirely absent from the fossil cerus—for a fossil group uniting Tragoportax, record of Chad (Geraads et al. 2001; Vignaud et al. Miotragocerus, and allies. 2002) and not well represented in the North African /Aegodontia record (Geraads et al. 1998). The Lukeino tragela- phine already the main horn core synapo- The clade including all bovids that are not morphies of the group, and no consensus was Bovinae is known as either Antilopinae (Kingdon reached on the possible existence of any stem 1982; Hassanin and Douzery 1999b) or Aegodon- group lineages or potential ancestors for Tragela- tia (Schlosser in Zittel 1911:499, sensu Vrba and phini. The stem tragelaphine lineage may have Schaller, 2000). The former term is more appropri- evolved either in Africa or Eurasia. Pheraios chrys- ate in the context of phylogenetic nomenclature but somallos (Kostopoulos and Koufos 2006), from the is problematic for the Linnean system, as it would late Miocene of Greece, has been proposed as a incorporate other subfamilial designations (e.g., possible stem tragelaphine. ). ‘Antilopinae’ (sensu Simpson 1945) has Boselaphini is a large, diverse, and, as cur- also been previously used to unite the Antilopini + rently understood, non-monophyletic group. A revi- , but the non- of the latter sion of the systematics of taxa variously grouped group (see below) means such a designation is as Boselaphini is warranted. The crown clade—i.e. now probably unnecessary, leaving the name avail- the two living boselaphines, Boselaphus tragocam- able for redefinition as the sister-group to Bovinae. elus and Tetracerus quadricornis, along with their Aegodontia has been recently redefined, but is immediate ancestors—is most likely separate from ambiguous in that this group would include bovids the majority of Miocene fossil forms generally that are not ‘aegodont’ (‘goat-toothed’) as originally

4 PALAEO-ELECTRONICA.ORG diagnosed. This clade is supported by numerous The old name Pseudotragini (sensu Solounias and morphological, behavioral, and molecular synapo- Moelleken 1992) has been proposed as a para- morphies (e.g., Kingdon 1982; Hassanin and Dou- phyletic group that includes possible stem zery 1999b; Vrba and Schaller 2000; Bibi 2009). caprines. Pachytragus solignaci (, MN9/10 Pseudoeotragus seegrabensis from Austria in Geraads 1989; ca.10Ma in Steininger 1999) has may be an early stem member of Antilopinae more specifically been proposed as a potential (Azanza and Morales 1994 figs. 7-8), with an age early crown caprine (Solounias and Moelleken of 18–17Ma (MN4, Made 1989; Agustí et al. 2001). 1992). “Ovibovini” is non-monophyletic, with phylo- The oldest substantiated African occurrences genetic studies partitioning Budorcas and Ovibos of Reduncini occur at Lothagam (Harris 2003) and among Caprinae, (Bouvrain and de Bonis 1984; Toros-Menalla (Vignaud et al. 2002). These may Ropiquet and Hassanin 2005). prove younger than the earliest occurrence in the The earliest confirmed hippotragines are Siwaliks, which may be 8Ma or older (Barry et al. known from Toros-Menalla, Chad, aged to about 2002). The cranium attributed to Pachytragus aff. 7Ma (Geraads et al. 2008). The Chadian fossils solignaci from the late middle or early late Miocene demonstrate that hippotragines present a mix of of Ngorora, Kenya (Thomas 1981), deserves re- derived and primitive characters that suggestively examination as a possible reduncine. The problem place them at the stem of the hippotragine clade. of the relatively poor African late Miocene fossil The maximal age of the most recent common record is again pertinent here. ancestor of all living Hippotragini (crown clade) is In line with recent phylogenetic studies, it was as a result set to around 7Ma. agreed that “Neotragini” is not monophyletic. The The oldest Alcelaphini are known from majority of “Neotragini” are paraphyletic with between about 7.5–6.5Ma from Lothagam, Kenya respect to Antilopini (Gentry 1992; Rebholz and (Harris 2003; McDougall and Feibel 2003). Alcela- Harley 1999). The most species-rich and wide- phini are virtually absent at Toros-Menalla, and not spread antilopine is Gazella. Fossils attributed to recorded from Mpesida (Thomas 1980) and Mio- this taxon go back to the late mid-Miocene of Fort Pliocene Middle Awash deposits (Haile-Selassie et Ternan, Kenya (ca. 14Ma, Gentry 1970; Shipman al. 2004; Haile-Selassie et al. 2009). Vrba’s (1997) et al. 1981). phylogenetic study suggests the origins of crown The origins of the , Aepyceros, remain Alcelaphini are to be found prior to 5Ma. unknown. The impala is first known from late Mio- cene African localities, with the oldest secure APPROACHES TO SYSTEMATICS records coming from Lothagam and Toros-Menalla Bovids, being diverse that inhabit a (Harris 2003; Geraads et al. 2008). A study of spectrum of habitats from arid environments to impala origins should investigate a possible con- wetlands, present an excellent resource with which nection with European and North African spiral- to formulate and test evolutionary and paleoeco- horned antilopines. The living species, A. melam- logical hypotheses. At the same time, their great pus, is first recorded in the latest Pliocene, at Koobi diversity presents a serious challenge to morpho- Fora (Harris 1991) and possibly Omo (Gentry logical systematics. Homoplasy in bovid evolution 1985). is evidently rampant (Gentry 1992), and the appli- The Hippotragini + Alcelaphini + Caprinae cation of comprehensive morphological cladistic together form a monophyletic clade, substantiated methods to bovid systematics requires much more by numerous recent phylogenetic studies (e.g., work, particularly as concerns the difficult task of Vrba and Schaller 2000; Ropiquet and Hassanin character formulation and coding. Over recent 2005). Fossil taxa such as Protoryx and Pachytra- , morphological cladistic analyses using a gus, debated to be either caprine or hippotragine selective set of characters and taxa (in contrast to (Bosscha Erdbrink 1988; Gentry 2000) may actu- ‘total’ methods) have had greater success in pro- ally lie closer to a common ancestor of both these ducing phylogenies commensurate with the fossil clades. Taxa such as Tethytragus, Gentrytragus, and molecular data (e.g., Vrba 1997; Geraads et and Caprotragoides may also lie in various posi- al. 2008). Future phylogenetic work would benefit tions in, or close to, the stem group of the larger greatly from increased incorporation of ontogenetic clade Caprini+Hippotragini+Alcelaphini (Azanza data (e.g., Vrba et al. 1994). and Morales 1994; Gentry 2000). The relationships Molecular systematics has over the last 20 of these three genera to one another are still not years provided significant insight into the evolution- known, particularly of Tethytragus to the other two. ary history of, and interrelationships within, Bovi-

5 Bibi et al.: Fossil Bovids

dae. Molecular systematic work provided strong Bovids are diagnostic components of biogeo- evidence for the existence of Antilopinae/Aegodon- graphic zones defined on the basis of whole fossil tia (Lowenstein 1986; Hassanin and Douzery faunas. For example, the fossil bovids from the late 1999b), the polyphyly of Neotragini (Hassanin and Miocene Baynunah Formation (United Arab Emir- Douzery 1999b; Matthee and Robinson 1999; Reb- ates, Gentry 1999) suggest an overlapping of the holz and Harley 1999) and Ovibovini (Groves and North African, Greco-Iranian (=Sub-Paratethyan), Shields 1997; Hassanin et al. 1998; Ropiquet and and Oriental (south Asian) Provinces. These tradi- Hassanin 2005). Molecular phylogenetic work has tional biogeographic provinces, outlined for the late also advanced knowledge of living but enigmatic Miocene by Bernor (1984), continue to remain rele- bovid species. Notable among these is the case of vant for discussion today, as do the topics the , Pseudoryx nghetinhensis, a goat-like addressed by past syntheses of bovid paleobio- living in the forests of and , geography (Gentry 1968; Thomas 1984a). only first described in 1993 (Dung et al. 1993; Systematic and paleontological work of the Kemp et al. 1997). Morphological comparisons last two decades has significantly improved the (Thomas 1994) suggested inclusion within the resolution at which bovid paleobiogeography can Caprini while molecular work proposed affinity to formulate and test hypotheses. For one, the pro- Bovinae and Bovini (Dung et al. 1993; Hassanin duction of more robust phylogenies, particularly by and Douzery 1999a). A ‘total evidence’ approach way of molecular methods, has allowed for the using both morphological and molecular characters rapid development of phylogeographic studies. supported the previous contentions of the molecu- Such studies have worked to reconstruct ancestral lar evidence, that the saola was in fact a member areas for higher clades (e.g., Hassanin and Ropi- of Bovinae and even nested deep within Bovini quet 2004) as well as the geographic range expan- (Gatesy and Arctander 2000). Pseudonovibos spi- sions and contractions of individual species (e.g., ralis, the linh duong, is another recently described Arctander et al. 1999). Additionally, the fossil bovid (Peter and Feiler 1994) of uncertain status. record has been sufficiently improved to allow for Pseudonovibos spiralis is known only from a num- new appraisal of late Miocene biogeographic ber of horn sheaths collected from Vietnam, and trends. Higher resolution paleobiogeographical controversy has raged over whether the character- studies are now possible that reveal developing istically spiraled and annulated horns are indeed zoogeographies both among and within biogeo- indicative of a new species, or whether they are graphic provinces in the late Miocene. For exam- simply modified horn sheaths of , Bos taurus. ple, marked differences appear to have Molecular phylogenetic studies have made a differentiated the Southern Balkan vs. Anatolian strong case that Pseudonovibos spiralis is simply a faunas of the late Miocene Greco-Iranian Province of Bos taurus (Peter and Feiler 1994; (Kostopoulos 2009). Olson and Hassanin 2003 and references therein). The presence of a monophyletic Caprini+Alcelaphini+Hippotragini (unnamed clade) BIOGEOGRAPHY provides a good opportunity for paleobiogeo- graphic studies as the living Caprini are almost The wide geographic range of Bovidae, and entirely Eurasian while recent Alcelaphini and Hip- the fact that many bovid clades are restricted in potragini are African (the latter are present in Ara- their distributions, makes this group ideal for bio- bia as well). On the basis of fossil and molecular geographic investigation. The Miocene evolution of phylogenetic evidence, Ropiquet and Hassanin Bovidae took place against the backdrop of major (2005) proposed that Caprini originated on Medi- climatic and physiographic changes, including the terranean islands in the late Miocene. In addition to closure of the Tethys Gateway (Berggren 1972), major cladogenetic and dispersal events, the opening of the East African and Red Sea rift val- record of rare and isolated occurrences of bovid leys (Ghebreab 1998), gradual global cooling taxa outside their expected geographic range is (Zachos et al. 2001), the Messinian salinity event another important indicator of biogeographic inter- (Hsü et al. 1973), and the global expansion of C 4 change. For example, caprines have a very limited grasslands at low latitudes (Cerling et al. 1997). As record of presence in Africa, and Plio-Pleistocene such, the search for correspondence between fossil caprines like Bouria and are global environmental changes and major biogeo- taken to be immigrants southwards from Eurasia. graphic events in the evolutionary history of Bovi- The age of occurrence of these fossil caprines in dae is a main focus of any synthetic treatment of Africa happens also to correspond to pronounced the bovid fossil record.

6 PALAEO-ELECTRONICA.ORG

global cooling events (Vrba 1995). Both Alcelaphini The environmental and ecological dynamics and Hippotragini have a limited Plio-Pleistocene responsible for the evolution and distribution of record (Pilgrim 1939) outside of Africa, namely in early hominid taxa are rarely to be reconstructed the Siwaliks, and Alcelaphini may have also from the hominids themselves. That bovids are occurred in the late Miocene of Italy with Marem- ecologically diverse and present in abundance at mia (e.g., Thomas 1984a). Additional possible Afri- hominid sites means this clade provides some of can occurrences in Europe include possible the strongest evidence with which to propose and reduncines from late Miocene Sicily and Turkey test hypotheses relating evolution to envi- (Seguenza 1902; Köhler 1987; Gentry 1999). Late ronmental changes. To those interested in the con- Miocene links between sub-Saharan Africa and text of early hominid evolution, these are southern Asia are also apparent in the shared interesting times as multiple independent fieldwork presence of bovid taxa such as the reduncine efforts, in Africa and elsewhere, target the period porrecticornis (see Haile-Selassie et al. between 5 and 10Ma. This span of time is much 2009). In light of the apparent barriers to exchange better known from Europe and will soon be better between southern Asia and the Greco-Iranian understood from southern and central Asia with Province (Beden and Brunet 1986), shared occur- work currently in progress. The grand task of rences of Bovidae between the Siwaliks and parts threading together the inter-continental picture of of Africa begs further investigation. evolutionary, biogeographic, and climatic changes will prove to be a major objective of those working BOVIDS IN THE CONTEXT OF on bovid paleontology and systematics in the years HOMINID ORIGINS to come. The oldest hominids known are Sahelanthro- ACKNOWLEDGMENTS pus tchadensis from Chad (ca.7Ma, Lebatard et al. 2008), Ardipithecus kadabba from Ethiopia (5.7Ma, Our meeting was made possible by funding WoldeGabriel et al. 2001), and Orrorin tugenensis from the Revealing Hominid Origins Initiative (PIs: from Kenya (ca.6Ma, Pickford and Senut 2001). T. White and the late F.C. Howell) under the U.S. Though these may be the three most popular National Science Foundation Award #BCS- actors on the late Miocene stage, it is on the basis 0321893. We are grateful to the National Museum of the remainder of the fossil fauna that the contex- of Ethiopia for welcoming us and to B. Asfaw and T. tual picture (climatic, vegetational, and biogeo- White for their help in conference organization. graphic) is drawn. A major impediment to F.Bibi was supported by a National Science Foun- increasing the resolution of the paleontological ‘big dation Graduate Research Fellowship. picture’ in the late Miocene is the limited nature of the African fossil record prior to 7Ma. Even with REFERENCES such limitations, Bovidae, often the most abun- Agustí, J., Cabrera, L., Garces, M., Krijgsman, W., Oms, dantly represented mammalian group in these fos- O., and Pares, J.M. 2001. A calibrated mammal sil contexts, have the greatest potential to inform scale for the Neogene of Western Europe. State of the biogeographic and climatic contexts of early the art. Earth-Science Reviews, 52:247-260. hominid origins. For example, the absence of Arctander, P., Johansen, C., and Coutellec-Vreto, M.A. tragelaphines from the late Miocene and Pliocene 1999. Phylogeography of three closely related Afri- Chadian fossil record (Geraads et al. 2001; Vig- can bovids ( Alcelaphini). Molecular Biology and naud et al. 2002) hints at deep biomic discontinui- Evolution, 16:1724-1739. ties between the Lake Chad basin and the entire Azanza, B., and Morales, J. 1994. Tethytragus nov. gen East African Rift Valley throughout this time. The et Gentrytragus nov. gen. Deux nouveaux Bovidés abundance of Tragelaphus at the 4.4Ma Aramis (Artiodactyla, Mammalia) du Miocène moyen. Pro- ceedings, Koninklijke Nederlandse Akademie Van hominid site provides a key indication that Ardipith- Wetenschappen, B 97:249-282. ecus ramidus inhabited a wooded environment Barry, J.C., Morgan, M.E., Flynn, L.J., Pilbeam, D., Beh- (WoldeGabriel et al. 1994). Changing proportions rensmeyer, A.K., Raza, S.M., Khan, I.A., Badgley, C., of alcelaphines and antilopines at late Pliocene Hicks, J., and Kelley, J. 2002. Faunal and environ- hominid sites provide a main support for the mental change in the late Miocene Siwaliks of north- hypothesis that Homo and Paranthropus originated ern Pakistan. Paleobiology, 28(S2):1-71. during a major drying trend between 2.8–2.0Ma Beden, M., and Brunet, M. 1986. Faunes de mammifères (Vrba 1985; Bobe and Eck 2001). et paléobiogéographie des domaines indiens et péri- indiens au Néogène. Sciences de la Terre, 47:61-87.

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