Combining isotopic and ecomorphological data to refine bovid paleodietary reconstruction: a case study from the Makapansgat Limeworks hominin locality

The relationship between environmental change and hominin evolution remains obscure. For the most part, this stems from the difficulty of reconstructing ancient hominin habitats. Bovids are among the most frequently utilized paleoenvironmental indicators, but little is known about the habitat preferences of extinct taxa. It is generally assumed that fossil bovids both ate the same things and occupied the same habitats as their closest extant relatives. We test the first part of this assumption by reconstructing the diets of seven bovids from Makapansgat Limeworks, South Africa. Since diet and habitat are linked, these reconstructions have implications for our understanding of fossil bovid habitat tolerances. Ecomorphological and stable carbon isotope analyses are employed, allowing us to take advantage of the strengths and overcome the weaknesses of both. In most cases, fossil bovids did have similar diets to their extant relatives, and probably occupied similar habitats. Gazella vanhoepeni and Aepyceros sp., however, were almost exclusive browsers, and not mixed feeders like their living counterparts.     

Biogeochemical and craniometric investigation of dietary ecology, niche separation, and taxonomy of Plio-Pleistocene cercopithecoids from the Makapansgat Limeworks

Three sympatric fossil cercopithecoid genera (Cercopithecoides, Parapapio, and Theropithecus) occur in Members 3 and 4 at the Makapansgat Limeworks hominin locality, South Africa, and their presence in a single ecosystem suggest a certain degree of ecological and/or dietary differentiation between taxa. Here, we explore the extent of dietary niche separation amongst these taxa using stable isotope (13C/12C, 18O/16O) and trace-element (Sr, Ba, Ca) analyses of fossil tooth enamel. In particular we searched for evidence of subtle niche separation between the more closely related, morphologically similar taxa of the genus Parapapio, as uncertainties exist around their taxonomy and taxonomic identification. Given these uncertainties, craniometric analyses were also performed to ground the dietary interpretations in a morphological context. The results found no clear taxonomic signal in the craniometric data for the Parapapio sample, and further indicate that this sample was no more variable morphologically than a single, geographically circumscribed, extant chacma baboon sample. In contrast, two overlapping dietary ecologies were found within this same Makapansgat Parapapio sample. Additionally, two widely differing dietary ecologies were found within the Cercopithecoides williamsi sample, while results for Theropithecus darti indicate a predominantly C4 diet. Hence, although biogeochemical dietary indicators point towards distinct dietary ecologies within and between fossil genera at Makapansgat, within the genus Parapapio disjunctions exist between the dietary categories and the taxonomic assignment of specimens.     

安徽东至华龙洞遗址洞穴演化与古人类活动

安徽东至华龙洞因发现距今约30万年的古人类头骨化石和大量动物化石及石制品受到学术界广泛关注。本文对华龙洞遗址的地质、地貌、沉积物特点及洞穴演化过程与古人类活动的关系进行分析。华龙洞地处扬子陆块区西北缘,周边呈现低山—丘陵—湖泊平原地貌景观。与华龙洞遗址密切关联的岩溶洞穴,发育在上寒武统微晶灰岩和白云质灰岩岩系内,中更新以来的地壳运动和岩溶发育是其形成的主要营力。华龙洞遗址是一处坍塌的洞穴,其发育大致经历发育初期（中更新世早期甚至更早）—稳定发育期（中更新世中期）—坍塌埋藏期（中更新世中晚期）等三个阶段,岩溶发育和洞外溪谷的侵蚀使得原始洞穴和堆积物一起在重力作用下坍塌。洞穴坍塌沉积物主要包括围岩岩块与碎屑、各种岩溶沉积物和文化遗物,胶结坚硬,不规则地埋藏于裂隙和巨石之间。古人类在遗址的活动时间处在距今约30万年前的稳定发育期,石制品和骨骼表面痕迹证据表明,华龙洞古人类具备依据不同原料的特点采取砸击法与锤击法并用的技术策略;石片边缘的使用痕迹和动物骨骼表面痕迹显示,古人类在遗址可能进行过肢解动物的行为。本研究对揭示长江下游中更新世中晚期古人类演化与适应生存行为具有重要意义。     

The first complete skull of Simopithecus darti from Makapansgat,South Africa,and its systematic position

The skull of a subadult female of Simopithecus darti from the “Upper Phase I” of the Makapansgat Limeworks, South Africa, is described. It is the first complete skull of Simopithecus which is recorded from the lower Pleistocene “Australopithecine cave deposits” of southern Africa. The present skull shows an interesting mixture of primitive and advanced features. In some respects, it is reminiscent of Parapapio, in others it seems to have affinities with the extant Theropithecus gelada rather than with Simopithecus oswaldi. The systematics of the fossil genus Simopithecus and of the tribe Theropithecini Jolly (1966) are discussed.     

Revised age estimates of Australopithecus-bearing deposits at Sterkfontein,South Africa

The Sterkfontein fossil site in South Africa has produced the largest concentration of early hominin fossils from a single locality. Recent reports suggest that Australopithecus from this site is found within a broad paleontological age of between 2.5-3.5 Ma (Partridge [2000] The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 100-125; Partridge et al. [2000a], The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 129-130; Kuman and Clarke [2000] J Hum Evol 38:827-847). Specifically, the hominin fossil commonly referred to as the "Little Foot" skeleton from Member 2, which is arguably the most complete early hominin skeleton yet discovered, has been magnetostratigraphically dated to 3.30-3.33 Ma (Partridge [2000] The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 100-125; Partridge et al. [2000a], The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 129-130). More recent claims suggest that hominin fossils from the Jacovec Cavern are even older, being dated to approximately 3.5 Ma. Our interpretation of the fauna, the archeometric results, and the magnetostratigraphy of Sterkfontein indicate that it is unlikely that any Members yet described from Sterkfontein are in excess of 3.04 Ma in age. We estimate that Member 2, including the Little Foot skeleton, is younger than 3.0 Ma, and that Member 4, previously dated to between 2.4-2.8 Ma, is more likely to fall between 1.5-2.5 Ma. Our results suggest that Australopithecus africanus should not be considered as a temporal contemporary of Australopithecus afarensis, Australopithecus bahrelghazali, and Kenyanthropus platyops.     

The paleoanthropology of Hadar,Ethiopia

Field research at the fossil-bearing deposits in the Afar Depression began in the 1970s. Prior to this, hominin fossils older than 3.0 Mya consisted of only a handful of fragments. During Phase I, the International Afar Research Expedition to Hadar, Ethiopia collected some 240 fossil hominins from Hadar over a time range of 3.0–3.4 Mya. Along with hominin fossils from Laetoli, they were deemed a new species, Australopithecus afarensis. This taxon was posited as the last common ancestor to robust Australopithecus and the Homo lineage in eastern Africa. Phase II research under the Hadar Research Project has added strength to the Phase I results, including the first association of a Homo fossil with stone tools at 2.4 Mya. This presentation is a cursory synopsis of the importance and implications of the hominin fossils recovered at Hadar during over the last 34 years.     

“许家窑人”埋藏地层与时代探讨

“许家窑人”化石发现于河北省阳原县泥河湾盆地西端的侯家窑遗址。该遗址20世纪70年代历经数次发掘,发现古人类化石20件、石制品数万件以及大量的哺乳动物化石。虽然已有大量关于遗址的研究成果发表,但人类化石和考古遗物的出土地层仍有些模糊,文化层时代也存在较大争议;年代数据在20 kaBP到500 kaBP的大范围间变化,影响学者们对“许家窑人”及其文化遗存演化位置的认识。鉴于此,河北省文物研究所（现河北省文物考古研究院）在2007~2012年对该遗址开展了持续的考古调查及发掘,重要目的之一在于廓清遗址埋藏的地貌部位和文化层的分布状况等基本信息。侯家窑遗址的地层包含上、下两个文化层;下文化层以下2~3 m（距地表约14.5 m）处确认了不整合接触面,其下为泥河湾层堆积,之上为河流阶地堆积。综合对20世纪70年代发掘的地层描述、新发掘地层以及人类化石保存状态的认识,“许家窑人”化石应出自遗址地层沉积序列的上文化层中,而非湖相的泥河湾层中。通过对目前遗址已有测年数据的重新梳理,我们认为,上文化层即“许家窑人”化石的时代为200~160 kaBP,主要对应深海氧同位素6阶段早期;下文化层的准确年代仍需验证,目前光释光年龄为198±15 kaBP, ²⁶Al/ ¹⁰Be埋藏年代为240±50 kaBP。     

Mass spectrometric U-series dating of Huanglong Cave in Hubei Province,central China: Evidence for early presence of modern humans in eastern Asia

Most researchers believe that anatomically modern humans (AMH) first appeared in Africa 160-190 ka ago, and would not have reached eastern Asia until ∼50 ka ago. However, the credibility of these scenarios might have been compromised by a largely inaccurate and compressed chronological framework previously established for hominin fossils found in China. Recently there has been a growing body of evidence indicating the possible presence of AMH in eastern Asia ca. 100 ka ago or even earlier. Here we report high-precision mass spectrometric U-series dating of intercalated flowstone samples from Huanglong Cave, a recently discovered Late Pleistocene hominin site in northern Hubei Province, central China. Systematic excavations there have led to the in situ discovery of seven hominin teeth and dozens of stone and bone artifacts. The U-series dates on localized thin flowstone formations bracket the hominin specimens between 81 and 101 ka, currently the most narrow time span for all AMH beyond 45 ka in China, if the assignment of the hominin teeth to modern Homo sapiens holds. Alternatively this study provides further evidence for the early presence of an AMH morphology in China, through either independent evolution of local archaic populations or their assimilation with incoming AMH. Along with recent dating results for hominin samples from Homo erectus to AMH, a new extended and continuous timeline for Chinese hominin fossils is taking shape, which warrants a reconstruction of human evolution, especially the origins of modern humans in eastern Asia.     

First Partial Skeleton of a 1.34-Million-Year-Old Paranthropus boisei from Bed II,Olduvai Gorge,Tanzania

Recent excavations in Level 4 at BK (Bed II, Olduvai Gorge, Tanzania) have yielded nine hominin teeth, a distal humerus fragment, a proximal radius with much of its shaft, a femur shaft, and a tibia shaft fragment (cataloged collectively as OH 80). Those elements identified more specifically than to simply Hominidae gen. et sp. indet are attributed to Paranthropus boisei. Before this study, incontrovertible P. boisei partial skeletons, for which postcranial remains occurred in association with taxonomically diagnostic craniodental remains, were unknown. Thus, OH 80 stands as the first unambiguous, dentally associated Paranthropus partial skeleton from East Africa. The morphology and size of its constituent parts suggest that the fossils derived from an extremely robust individual who, at 1.338±0.024 Ma (1 sigma), represents one of the most recent occurrences of Paranthropus before its extinction in East Africa.     

Stretching the time span of hominin evolution at Kromdraai (Gauteng,South Africa): Recent discoveries

The Plio-Pleistocene locality of Kromdraai B has yielded the type specimen of Paranthropus robustus, as well as 27 additional fossil hominin specimens. In a number of both cranial and dental features, the states shown by the Kromdraai Paranthropus are more conservative when compared to the more derived conditions displayed by both South African conspecifics and the post-2.3 Ma eastern African Paranthropus boisei. Since 2014, we excavated the earliest known infilling of the Kromdraai cave system in a previously unexplored area. This new locality provided as yet 2200 identifiable macrovertebrate fossils, including 22 hominins, all tied in the earliest part of the stratigraphic sequence, representing three distinct depositional periods. Since we report here, for the first time, the occurrence of fossil hominins in Members 1 and 2, our discoveries stretch the time span of hominin evolution at Kromdraai and contribute to a better understanding of the origin of Paranthropus in southern Africa.     

Climatic influences on the evolution of early Homo?

The nature of the human fossil record is less than ideal for the generation of precise correlations between environmental variables and patterns of evolution in specific lineages. Nonetheless, a critical look at what can and cannot be said from individual fossil morphology and the correlation of specific environmental proxies with specific hominin fossils may lead to a greater understanding of the degree of certainty with which we should embrace environmental hypotheses for the evolution of Homo. Climate shifts have been implicated in both the origin of the genus and its dispersal from Africa. Here, I consider three areas in which a climatic influence has been posited to explain evolutionary shifts in the genus Homo: the origin and dispersal of the genus from Africa; geography, climate and body size in early Homo, and the influence of climate-induced sea level rise on morphological isolation in H. erectus. Each of the data sets is far from ideal, and interpretations of each of the data sets are fraught with issues of equifinality. Of the three hypotheses discussed, the clearest link is seen between latitudinal variation (and presumably temperature) and body size in H. erectus. Similarly, climate-induced sea level change seems a reasonable isolating mechanism to explain the pattern of cranial variation in later Asian H. erectus, but the distribution could also reflect incompletely sampled clinal variation. Alternatively, only equivocal support is found for the influence of climate on the differentiation of H. erectus from H. habilis (as proxied by body/brain size scaling), and therefore the dispersal of the genus Homo cannot be as clearly linked to changes in body size and shape as it has been in the past. These preliminary data suggest that an emphasis on understanding local adaptation before looking at global (and specific) level change is critical to elucidating the importance of climatic factors on the evolution of the genus Homo.     

New Australopithecus robustus fossils and associated U-Pb dates from Cooper's Cave (Gauteng, South Africa)

Australopithecus robustus is one of the best represented hominin taxa in Africa, with hundreds of specimens recovered from six fossil localities in the Bloubank Valley area of Gauteng Province, South Africa. However, precise geochronological ages are presently lacking for these fossil cave infills. In this paper, we provide a detailed geological background to a series of hominin fossils retrieved from the newly investigated deposit of Cooper's D (located partway between Sterkfontein and Kromdraai in the Bloubank Valley), including uranium-lead (U-Pb) ages for speleothem material associated with A. robustus. U-Pb dating of a basal speleothem underlying the entire deposit results in a maximum age of 1.526 (±0.088) Ma for Cooper's D. A second U-Pb date of ca. 1.4 Ma is produced from a flowstone layer above this basal speleothem; since this upper flowstone is not a capping flowstone, and fossiliferous sediments are preserved above this layer, some of the hominins might be slightly younger than the calculated age. As a result, we can broadly constrain the age of the hominins from Cooper's D to between 1.5 and approximately 1.4 Ma. Extinct fauna recorded in this comparatively young deposit raise the possibility that the Bloubank Valley region of South Africa represented a more stable environmental refugium for taxa relative to tectonically more active East Africa. The sediments of the deposit likely infilled rapidly during periods when arid conditions prevailed in the paleoenvironment, although it is unclear whether sediment deposition and bone deposition were necessarily contemporaneous occurrences. We reconstruct the paleoenvironment of Cooper's D as predominantly grassland, with nearby woodlands and a permanent water source. The hominin teeth recovered from Cooper's D are all from juveniles and can be confidently assigned to A. robustus. In addition, two juvenile mandibular fragments and an adult thoracic vertebra are tentatively attributed to A. robustus.     

Microwear evidence for Plio-Pleistocene bovid diets from Makapansgat Limeworks Cave, South Africa

Makapansgat Limeworks Cave is a well-known Australopithecus africanus bearing locality that has spawned a considerable amount of paleoecological research because of its hominin component. Most recently, the paleoecology of this Plio-Pleistocene site has been studied by determining the diet and habitat of other extinct taxa, particularly the bovids. The diets of seven bovids (Aepyceros sp., Gazella vanhoepeni, Makapania broomi, Parmularius braini, Redunca darti, Tragelaphus sp. aff. T. angasii, and Tragelaphus pricei) have now been classified using taxonomic uniformitarianism, ecomorphology, stable carbon isotopes, and mesowear analysis. Here, dental microwear is applied to the same bovids for additional comparison and to further elucidate the strengths and weaknesses of each method. The different dietary proxy methods noted provide a temporal continuum, with genetic signals such as ecomorphology and taxonomic uniformitarianism indicating behavioral adaptations over geologic time, while nongenetic data such as stable carbon isotopes and mesowear reflect different aspects of average diet over extended portions of an animal's life, and dental microwear provides dietary snapshots.Microwear separated an extant baseline of ten bovid species into expected dietary categories and the Makapansgat bovids clearly fell into two groups with the same degree of separation as between extant grazers and browsers. The results indicate that a multidisciplinary approach produces a more accurate and robust reconstruction of past diets. In sum, the microwear analysis is in-line with the isotope and mesowear results, which suggest a stronger browsing component than either taxonomic uniformitarianism or ecomorphology imply.     

The Atapuerca sites and their contribution to the knowledge of human evolution in Europe

Over the last two decades, the Pleistocene sites of the Sierra de Atapuerca (Spain) have provided two extraordinary assemblages of hominin fossils that have helped refine the evolutionary story of the genus Homo in Europe. The TD6 level of the Gran Dolina site has yielded about one hundred remains belonging to a minimum of six individuals of the species Homo antecessor. These fossils, dated to the end of the Lower Pleistocene (800 kyr), provide the earliest evidence of hominin presence in Western Europe. The origin of these hominins is unknown, but they may represent a speciation event from Homo ergaster/Homo erectus. The TD6 fossils are characterized by a significant increase in cranial capacity as well as the appearance of a “sapiens” pattern of craniofacial architecture. At the Sima de los Huesos site, more than 4,000 human fossils belonging to a minimum of 28 individuals of a Middle Pleistocene population (ca. 500–400 kyr) have been recovered. These hominins document some of the oldest evidence of the European roots of Neanderthals deep in the Middle Pleistocene. Their origin would be the dispersal out of Africa of a hominin group carrying Mode 2 technologies to Europe. Comparative study of the TD6 and Sima de la Huesos hominins suggests a replacement model for the European Lower Pleistocene population of Europe or interbreeding between this population and the new African emigrants.     

Cercopithecoid primate postcranial fossils from Cooper's D,South Africa

Among several highly fossiliferous localities in the Bloubank Valley (Gauteng, South Africa), the Cooper's Cave System has been known since 1938 and has produced a rich fossil assemblage, including some remains of the early hominin Paranthropus robustus. In 2001, excavations began at a new locality, Cooper's D, which dates to ～1.4-1.5 Ma. Although hominins are relatively rare in the assemblage, remains of cercopithecoid primates are much more common. Craniodental fossils currently indicate the possible presence of at least three large-bodied cercopithecoid primate genera at Cooper's D: Gorgopithecus, Papio, and Theropithecus. In this study, we identify and describe > 100 cercopithecoid primate postcranial fossils representing all regions of the appendicular skeleton. The specimens come from several age classes and size morphs; more than one third of the fossils described are from sub-adult and juvenile individuals. The adult postcranial fossils vary substantially in size, with body masses estimated between 30 and 60 kg (from 16 of the better preserved specimens). The functional morphology of the postcranial remains indicate that these elements come from animals that likely utilized terrestrial substrates, but they remain difficult to definitively attribute to Gorgopithecus, Theropithecus, or Papio given the absence of associated skeletons. The smaller specimens likely belong to Papio while the larger ones can be attributed to the other two genera. Because Cooper's D has also yielded fossils of the early hominin Paranthropus robustus, this raises the question of how these four large-bodied, mostly terrestrial primates sympatrically utilized the landscape.     

Bone breccias, bone dumps, and sedimentary sequences of the western Limeworks, Makapansgat, South Africa

The sediments in the western side of the Makapansgat Limeworks were either precipitated as speleothems, represented in the earlier massive deposits, or were deposited as coarsening clastic sediments, mainly representing later deposits. Between the earlier deposits and the main sedimentary phase, the stratigraphic sequence was inverted twice to a considerable height by the unusual deposition of subaqueous speleothem. Bone-bearing deposits, including the Main Quarry Bone Breccia and the well-known Grey Breccia belong, in time, to the lower part of the clastic deposits called the Red Silts. Australopith fossils have been found in the Grey Breccia dumped material and, in situ, from the dolomite clast breccia on the Main Quarry entrance buttress. Whatever the problems may be in provenancing some of the material from the Limeworks dumps, there is no doubt that the three rows of blocks on the southern side of the dumps belong to the Grey Breccia, and other rows contain red sediment sufficient for them to be safely associated with the Red Silts. There is no reason why this material should not be prepared with confidence as to its stratigraphic provenance. In any case, stratigraphic evidence, presented here, shows that there is little difference in time between the deposition of the bone-bearing breccias.     

Brief Communication: Beyond the South African cave paradigm—Australopithecus africanus from Plio–Pleistocene paleosol deposits at Taung

Following the discovery of the “Taung Child” (Australopithecus africanus) in 1924 in the Buxton‐Norlim Limeworks near Taung, the fossil‐bearing deposits associated with the Dart and Hrdli?ka pinnacles have been interpreted as the mined remnants of cave sediments that formed within the Plio–Pleistocene Thabaseek Tufa: either as a younger cave‐fill or as contemporaneous carapace caves. When combined with the Plio–Pleistocene dolomitic cave deposits from the “Cradle of Humankind,” a rather restricted view emerges that South African early hominins derived from cave deposits, whereas those of east and central Africa are derived from fluvio‐lacustrine and paleosol deposits. We undertook a sedimentological and paleomagnetic analysis of the pink‐colored deposit (PCS) from which the “Taung Child” is purported to have derived and demonstrate that it is a calcrete, a carbonate‐rich pedogenic sediment, which formed on the paleo‐land surface. The deposit extends 100 s of meters laterally beyond the Dart and Hrdli?ka Pinnacles where it is interbedded with the Thabaseek Tufa, indicating multiple episodes of calcrete development and tufa growth. The presence of in situ rhizoconcretions and insect trace fossils (Celliforma sp. and Coprinisphaera sp.) and the distinctive carbonate microfabric confirm that the pink deposit is a pedogenic calcrete, not a calcified cave sediment. Paleomagnetic and stratigraphic evidence indicates that a second, reversed polarity, fossil‐bearing deposit (YRSS) is a younger fissure‐fill formed within a solutional cavity of the normal polarity tufa and pink calcrete (PCS). These observations have implications for the dating, environment, and taphonomy of the site, and increase the likelihood of future fossil discoveries within the Buxton‐Norlim Limeworks. Am J Phys Anthropol 151:316–324, 2013. © 2013 Wiley Periodicals, Inc.     

The Jinniushan hominin pedal skeleton from the late Middle Pleistocene of China

Bipedalism has long been recognized as the seminal adaptation of the hominin radiation and thus used to distinguish hominins from great ape fossils. Notwithstanding preconceptions and varied interpretations, the distinctive features of the modern human foot and accompanying striding gait, appear to be recent innovations that are largely absent in the earliest facultative bipeds. These distinctive features are mainly components of fixed longitudinal and transverse pedal arches, and of a uniquely derived hallucal metatarsophalangeal joint. They enhance ankle joint plantar flexor function and accommodate localized peak plantar pressures at the medial ball during terminal stance. To date, the paleontological record has yielded very little of the hominin foot, especially of the Middle Pleistocene hominins. New specimens from this time interval should help provide insights into the timing and pattern of what appears to be a mosaic pattern of evolution of the modern human foot features. Here we describe the fossil hominin foot skeleton recovered from the Jinniushan site, Liaoning Province, People's Republic of China. It affords a singular glimpse of the pedal morphology of a late Middle Pleistocene hominin (c.f. Homo heidlebergensis). Dated to 200 ka or older, this foot offers the earliest evidence for increased stability of the medial longitudinal arch, while retaining a number of primitive features apparently characteristic of robust premodern hominins, including lower arches and a less stable hallucal metatarsophalangeal joint (medial ball) than in modern humans. These features reflect different foot capabilities and suggest the bipedal stride of the Jinniushan hominin differed subtlety from that of modern humans.     

Theropithecus and 'Out of Africa' dispersal in the Plio-Pleistocene

Theropithecus oswaldi was one of the most widely distributed Plio-Pleistocene primates, found in southern, East, and North Africa, as well as in Spain, India, and possibly Italy. Such a large geographic range for a single primate species is highly unusual. Here, the nature and timing of its dispersal is examined using the Stepping Out cellular automata model. A hypothetical dispersal of T. darti is also modelled to assess whether the late Pliocene might have been a more favorable period for Afro-Eurasian dispersal than the early Pleistocene. Stepping Out draws on climatic and biome reconstruction to provide the paleovegetative and climatic background necessary for the simulations, and model parameters for T. oswaldi and T. darti were set a priori on the basis of their fossil records and paleobiologies. The simulations indicate that T. darti could have readily left Africa in the Pliocene, and that it swiftly reaches Asia. A European T. darti colonization was less certain and less rapid. The simulated T. oswaldi dispersal out of Africa was slower, but nonetheless T. oswaldi arrived at Mirzapur within the time period indicated by the fossil record. Using the a priori parameters, T. oswaldi did not arrive at the European sites of Cueva Victoria and Pirro Nord. It cannot be discounted, therefore, that some of the European fossils are a result of an earlier T. darti dispersal. The simulations also showed that in order for Theropithecus to reach Europe, it needed to be tolerant of a relatively wide range of habitats. In addition, our finding that Asian colonization was more rapid and more probable parallels the information from the hominin fossil record, in which the fossils from Asia predate those from Europe by several hundred thousand years.