Papio Cranium from the Hominin-Bearing Site of Malapa: Implications for the Evolution of Modern Baboon Cranial Morphology and South African Plio-Pleistocene Biochronology

A new partial cranium (UW 88-886) of the Plio-Pleistocene baboon Papio angusticeps from Malapa is identified, described and discussed. UW 88-886 represents the only non-hominin primate yet recovered from Malapa and is important both in the context of baboon evolution as well as South African hominin site biochronology. The new specimen may represent the first appearance of modern baboon anatomy and coincides almost perfectly with molecular divergence date estimates for the origin of the modern P. hamadryas radiation. The fact that the Malapa specimen is dated between ~2.026–2.36 million years ago (Ma) also has implications for the biochronology of other South African Plio-Pleistocene sites where P. angusticeps is found.     

Carnivora from the Plio-Pleistocene hominin site of Drimolen, Gauteng, South Africa

Drimolen is one of the newest and most productive hominin sites in South Africa, and is dated on faunal grounds between 2.0 Ma to 1.5 Ma. This paper provides the first overview of the Carnivora from Drimolen, updating the previously published preliminary faunal list, and describing all currently prepared craniodental and postcranial material. The Drimolen specimens are described in comparison with other modern and fossil South African carnivore material. The carnivores cover a range of taxa including hyaenids, felids, canids and herpestids. Most notable amongst these are the sabretooth Dinofelis aff. piveteaui craniodental and postcranial remains, which are described in detail, and a Chasmaporthetes nitidula cranium. The genus Chasmaporthetes is found at three other sites in the area - Sterkfontein, Swartkrans and Coopers D. There are two models for the geographic origin of Dinofelis piveteaui, in that it may have arisen in either eastern or southern Africa. These possibilities are discussed in the light of the new South African Dinofelis material, as the Drimolen material appears to represent a more primitive form with affinities with D. piveteaui. Fossil leopard material from Kromdraai B and Drimolen is also discussed, as the metapodia assigned to P. pardus from these two sites are very small, but lie within the variation of modern leopards. Such size differences in fossil postcrania may have implications for the niches that these animals may have occupied in the past.     

The large carnivores from ‘Ubeidiya (early Pleistocene, Israel): biochronological and biogeographical implications

The archaeological site of ‘Ubeidiya provides a unique, rich faunal assemblage, as well as evidence for the presence of early hominins in the Levant. Paleontological analysis of the faunal community is pertinent to understanding the biochronology and the timing of hominin dispersal events from Africa to Eurasia. This study presents the large carnivores from the recent seasons of excavations (1989-1999) and previously published material. Based on biochronological correlations of the carnivore guild and paloemagnetism, the age of the site is 1.5-1.2 Ma. This date is 100-200,000 years earlier than previous estimates. Of specific importance is the presence of the African origin saber tooth Megantereon cf. M. whitei and the Eurasian origin canids Canis moschbachensis and Lycaon lycoanoides.     

Morphometric variation in the papionin muzzle and the biochronology of the South African Plio‐Pleistocene karst cave deposits

Papionin monkeys are widespread, relatively common members of Plio‐Pleistocene faunal assemblages across Africa. For these reasons, papionin taxa have been used as biochronological indicators by which to infer the ages of the South African karst cave deposits. A recent morphometric study of South African fossil papionin muzzle shape concluded that its variation attests to a substantial and greater time depth for these sites than is generally estimated. This inference is significant, because accurate dating of the South African cave sites is critical to our knowledge of hominin evolution and mammalian biogeographic history. We here report the results of a comparative analysis of extant papionin monkeys by which variability of the South African fossil papionins may be assessed. The muzzles of 106 specimens representing six extant papionin genera were digitized and interlandmark distances were calculated. Results demonstrate that the overall amount of morphological variation present within the fossil assemblage fits comfortably within the range exhibited by the extant sample. We also performed a statistical experiment to assess the limitations imposed by small sample sizes, such as typically encountered in the fossil record. Results suggest that 15 specimens are sufficient to accurately represent the population mean for a given phenotype, but small sample sizes are insufficient to permit the accurate estimation of the population standard deviation, variance, and range. The suggestion that the muzzle morphology of fossil papionins attests to a considerable and previously unrecognized temporal depth of the South African karst cave sites is unwarranted. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

The stem species of our species: a place for the archaic human cranium from Ceprano, Italy

One of the present challenges in the study of human evolution is to recognize the hominin taxon that was ancestral to Homo sapiens. Some researchers regard H. heidelbergensis as the stem species involved in the evolutionary divergence leading to the emergence of H. sapiens in Africa, and to the evolution of the Neandertals in Europe. Nevertheless, the diagnosis and hypodigm of H. heidelbergensis still remain to be clarified. Here we evaluate the morphology of the incomplete cranium (calvarium) known as Ceprano whose age has been recently revised to the mid of the Middle Pleistocene, so as to test whether this specimen may be included in H. heidelbergensis. The analyses were performed according to a phenetic routine including geometric morphometrics and the evaluation of diagnostic discrete traits. The results strongly support the uniqueness of H. heidelbergensis on a wide geographical horizon, including both Eurasia and Africa. In this framework, the Ceprano calvarium--with its peculiar combination of archaic and derived traits--may represent, better than other penecontemporaneous specimens, an appropriate ancestral stock of this species, preceding the appearance of regional autapomorphic features.     

New Viverridae specimens from the Malapa Hominin Site and their utility as palaeoenvironmental indicators

The use of small-bodied animals, especially small Carnivora, as indicators of palaeoenvironment, is becoming more pronounced. These smaller taxa are more spatiotemporally constrained, owing to high territoriality, ecosystem sensitivity and physical limitations on their ability to travel large distances within their lifespan. This article describes seven new Viverridae specimens, together with a re-evaluation of a previously published specimen, from the hominin-bearing site of Malapa, located in the Cradle of Humankind World Heritage Site, South Africa. When combined with other palaeoenvironmental indicators, the addition of these eight specimens, consisting of Genetta sp., Genetta genetta, and Genetta cf. tigrina, allows for a more precise reconstruction of the environment surrounding Malapa as well as providing additional insights into possible depositional factors that could have resulted in the accumulation of the Malapa deposit.     

A revision of the genus Dinofelis (Mammalia, Felidae)

This paper consists of a taxonomic and systematic revision of the extinct felid genus Dinofelis (Felidae, Ma-chairodontinae) and an analysis of its ecomorphology and evolution. Dinofelis has a broad distribution, with material from all northern continents and Africa, the latter of which was the apparent centre of evolution of the genus. We describe new material of Dinofelis from a number of sites in eastern Africa and reconsider all previously described material. We name two new species and identify several other distinct species-level taxa but refrain from naming these due to a paucity of well-preserved material. At the same time, we synonymize the two named Asian species, D. cristata and D. abeli , of which the former has priority. There are few characters useful in systematic analysis, but we can suggest at least one migration from eastern to southern Africa. Ecomorphological analysis of both craniodental and postcranial characters suggests that Dinofelis in many respects converged on modern pantherine cats in morphology and behaviour, a trend culminating in the South African D. barlowi and the Asian D. cristata , which are the most pantherine-like of all machairodont felids. This trend is reversed in the evolution of the youngest species, D. piveteaui , which is also the most machairodont in its ecomorphology. The timing of the extinction of Dinofelis is difficult to determine. Outside Africa material is scarce at all times, while in Africa the apparent extinction of Dinofelis at about 1.4 Mya coincides with the end of the good, semi-continuous fossil record present in eastern Africa from about 4 Mya onwards. Dating of Kanam East (with D. piveteaui) to the Jaramillo Subchron (1.070-0.990 Mya) suggests possible survival considerably later. Thus, the extinction datum for Dinofelis cannot at present be firmly established.     

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.     

Taphonomic Analysis of the Faunal Assemblage Associated with the Hominins (Australopithecus sediba) from the Early Pleistocene Cave Deposits of Malapa,South Africa

Here we present the results of a taphonomic study of the faunal assemblage associated with the hominin fossils (Australopithecus sediba) from the Malapa site. Results include estimation of body part representation, mortality profiles, type of fragmentation, identification of breakage patterns, and microscopic analysis of bone surfaces. The diversity of the faunal spectrum, presence of animals with climbing proclivities, abundance of complete and/or articulated specimens, occurrence of antimeric sets of elements, and lack of carnivore-modified bones, indicate that animals accumulated via a natural death trap leading to an area of the cave system with no access to mammalian scavengers. The co-occurrence of well preserved fossils, carnivore coprolites, deciduous teeth of brown hyaena, and some highly fragmented and poorly preserved remains supports the hypothesis of a mixing of sediments coming from distinct chambers, which collected at the bottom of the cave system through the action of periodic water flow. This combination of taphonomic features explains the remarkable state of preservation of the hominin fossils as well as some of the associated faunal material.     

The Australopithecus assemblage from Sterkfontein Member 4 (South Africa) and the concept of variation in palaeontology

Interpreting morphological variation within the early hominin fossil record is particularly challenging. Apart from the fact that there is no absolute threshold for defining species boundaries in palaeontology, the degree of variation related to sexual dimorphism, temporal depth, geographic variation or ontogeny is difficult to appreciate in a fossil taxon mainly represented by fragmentary specimens, and such variation could easily be conflated with taxonomic diversity. One of the most emblematic examples in paleoanthropology is the Australopithecus assemblage from the Sterkfontein Caves in South Africa. Whereas some studies support the presence of multiple Australopithecus species at Sterkfontein, others explore alternative hypotheses to explain the morphological variation within the hominin assemblage. In this review, I briefly summarize the ongoing debates surrounding the interpretation of morphological variation at Sterkfontein Member 4 before exploring two promising avenues that would deserve specific attention in the future, that is, temporal depth and nonhuman primate diversity.     

Recent paleoanthropological excavations of in situ deposits at Makapansgat, South Africa--a first report

The Makapansgat Limeworks is a significant Pliocene site both for its sample of 35 hominin fossils as well as its wealth of fossil fauna. The lithological and paleontological successions reveal local environmental changes that are important for understanding the context of hominin evolution in southern Africa. Yet most of the site's fossils were found in dumps left behind by quarry operations, and the paleoecological interpretations rest upon debatable assumptions about the original fossil provenience. We have recently initiated systematic paleoanthropological excavations at Makapansgat to recover well provenanced fossils in order to: 1) assess whether faunal successions are discernable in the Makapansgat sequence; 2) assist environmental interpretations of the site; 3) and potentially recover the oldest hominins in South Africa, roughly coincident with Australopithecus afarensis in East Africa. This paper presents a summary of our current paleoenvironmental research at the Limeworks and preliminary results of ongoing in situ excavations.     

The Kromdraai early hominin-bearing site. A review of recent findings

The Kromdraai archaeological site is located in a fossiliferous paleokarst situated in the UNESCO World Heritage Site referred to as the “Cradle of Humankind” in the Gauteng Province of South Africa. Kromdraai is noteworthy because it features among the three southern African early hominin-bearing sites considered to represent distinct temporal periods within the same stratigraphic succession. Kromdraai also yielded a partial skull and dentition (TM 1517) in 1938 that was designated as the holotype of a new genus and species, Paranthropus robustus. Although the hominin fossil assemblage collected from Kromdraai between 1938 and 2014 is rather paltry, morphometric and cladistic analyses of this material suggested that it represented a somewhat less-derived form of P. robustus than the considerably larger assemblage from the nearby site of Swartkrans. However, the geochronological and biotic relationships among the P. robustus-bearing sites in South Africa are not resolved. Since 2014, the previously unknown, albeit densely fossiliferous Unit P produced 51 individually catalogued hominin fossils (36 craniodental and 15 postcranial) that currently represent 13% of the faunal assemblage from this unit with a minimum number of 10 juvenile and 9 adult individuals. P. robustus and early Homo coexisted at the time of the accumulation of Unit P at Kromdraai, with a relative abundance of 89% and 11%, respectively. P. robustus and early Homo are associated with a highly diverse fauna that includes several water-dependent species, and a large variety of bovid and carnivore taxa. Biochronological data from Unit P and an interval of reversed polarity measured in younger deposits above it are interpreted in the context of the regional chronological framework to infer that it represents a deposit that was likely accumulated prior to 2 Ma.     

Miocene cercopithecoidea from the Tugen Hills, Kenya

Miocene to Pleistocene fossiliferous sediments in the Tugen Hills span the time period from at least 15.5 Ma to 0.25 Ma, including time periods unknown or little known elsewhere in Africa. Consequently, the Tugen Hills deposits hold the potential to inform us about crucial phylogenetic events in African faunal evolution and about long-term environmental change. Among the specimens collected from this region are a number of discoveries already important to the understanding of primate evolution. Here, we describe additional cercopithecoid material from the Miocene deposits in the Tugen Hills sequence, including those from securely dated sites in the Muruyur Beds (16-13.4 Ma), the Mpesida Beds (7-6.2 Ma) and the Lukeino Formation (∼6.2-5.7 Ma). We also evaluate previously described material from the Ngorora Formation (13-8.8 Ma). Identified taxa include Victoriapithecidae gen. et sp. indet., cf. Parapapio lothagamensis, and at least two colobines. Specimens attributed to cf. Pp. lothagamensis would extend the species’ geographic range beyond its type locality. In addition, we describe specimens sharing derived characters with modern African colobines (Tribe: Colobina), a finding that is congruent with previous molecular estimates of colobine divergence dates. These colobine specimens represent some of the earliest known members of the modern African colobine radiation and, in contrast to previous hypotheses, suggest that early African colobines were mainly arboreal and that semi-terrestrial Late Miocene and Plio-Pleistocene colobine taxa were secondarily derived in their locomotor adaptations.     

Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression,Ethiopia

Hadar is well known as one of the most productive early hominin sites in the world. Between 1972 and 1994 a large sample of fossil cercopithecid specimens was collected from Hadar and the nearby sites of Geraru, Ahmado, and Leadu. At least five, and possibly six, species are present in the sample, including two chronological subspecies of Theropithecus oswaldi. T. o. cf. darti is known from the Middle Pliocene deposits in the Hadar area, along with Parapapio cf. jonesi, cf. Rhinocolobus turkanaensis, and a new species of Cercopithecoides, C. meaveae. There are also isolated molars from the Middle Pliocene of a large colobine which most likely represent cf. R. turkanaensis, but may also represent another large colobine known from the nearby site of Maka in the Middle Awash. T. o. oswaldi is represented from younger deposits of Late Pliocene and Early Pleistocene age, along with the large colobine Cercopithecoides kimeui. Throughout the sequence Theropithecus oswaldi is by far the most abundant cercopithecid, with the other taxa being comparatively rare. The Parapapio material from Hadar is important as the only securely identifiable material of the genus in the East African Pliocene. Furthermore, the Hadar material includes the only associated postcranial remains for the genus. If the tentative identification of Rhinocolobus is correct, then the Hadar sample is the only known occurrence outside of the Turkana Basin. Cercopithecoides meaveae is a new species, currently only known from the Hadar region, most importantly by the associated partial skeleton from Leadu. It appears to show adaptations for terrestrial locomotion. Finally, Cercopithecoides kimeui, a very large colobine previously known from Olduvai Gorge, Koobi Fora, and Rawi is recorded from the uppermost part of the Formation.     

Speleology and magnetobiostratigraphic chronology of the GD 2 locality of the Gondolin hominin-bearing paleocave deposits, North West Province, South Africa

Speleological, paleomagnetic, mineral magnetic, and biochronological analyses have been undertaken at the Gondolin hominin-bearing paleocave, North West Province, South Africa. Two fossiliferous but stratigraphically separate sequences, GD2 and GD1/3, which were once part of a large cavern system, have been identified. Although some comparative paleomagnetic samples were taken from the GD 1, 3, and 4 localities that are currently under investigation, the research presented here focuses on the fossil-rich, in situ deposits at locality GD 2, excavated by E.S. Vrba in 1979. The GD 2 deposits are dominated by normal-polarity calcified clastic deposits that are sandwiched between clastic-free flowstone speleothems. The lower flowstone has a sharp contact with the red siltstone deposits and is of reversed polarity. The capping flowstone shows a change from normal to reversed polarity, thereby preserving a polarity reversal. While the paleomagnetic work indicates that the GD 2 fossil material was deposited during a normal-polarity period, the shortness of the sequence made matching of the magnetostratigraphy to the geomagnetic polarity time scale (GPTS) impossible without the aid of biochronology. While lacking multiple time-sensitive taxa, the recovery of specimens attributable to Stage III Metridiochoerus andrewsi is consistent with a deposition date between 1.9 and 1.5 Ma. A comparison of the magnetostratigraphy with the GPTS therefore suggests that the fauna-bearing siltstone of GD 2 date to the Olduvai normal-polarity event, which occurred between 1.95 and 1.78 Ma, and that the reversal from normal to reversed polarity identified in the capping flowstone dates to 1.78 Ma. The main faunal layers therefore date to slightly older than 1.78 Ma. Deposits from the GD 1 locality are dominated by reversed directions of magnetization, which show that this deposit is not of the same age as the faunal layers from the GD 2 locality.     

Tracking changing environments using stable carbon isotopes in fossil tooth enamel: an example from the South African hominin sites

The environmental contexts of the karstic hominin sites in South Africa have been established largely by means of faunal associations; taken together these data suggest a trend from relatively closed and more mesic to open, drier environments from about 3 to 1.5 Ma. Vrba argued for a major shift within this trend ca. 2.4-2.6 Ma, an influential proposal that posited links between bovid (and hominin) radiation in Africa and the intensification of Northern Hemisphere Glaciation. Yet faunal approaches often rely on habitat and feeding preferences of modern taxa that may differ from those of their extinct predecessors. Here we explore ways of extending ¹³C/¹²C data from fossil mammals beyond denoting “presence” or “absence” of C₄ grasses using the evolution of open environments in South Africa as a case study. To do so we calculated the relative proportions of C₃-, mixed-, and C₄-feeding herbivores for all the hominin sites for which we have sufficient data based on ¹³C/¹²C analyses of fossil tooth enamel. The results confirm a general trend towards more open environments since 3 Ma, but they also emphasize a marked change to open grassy habitats in the latest Pliocene/early Pleistocene. Mean ¹³C/¹²C for large felids also mirrored this trend.     

Using radii-of-curvature for the reconstruction of extinct South African carnivoran masticatory behavior

Paleoanthropologists have hypothesized that, during the evolution of increased carnivory in our lineage, hominins transitioned through a scavenging niche created by certain carnivoran taxa (especially sabertooths) that may have lacked the morphology necessary to utilize all parts of carcasses, thus leaving an open niche of high-quality scavengable remains. In this article, we examine the postcanine dentition of modern and fossil carnivorans using quantifications of occlusal radii-of-curvature (ROC) and correlate this morphology with feeding behavior to deduce the carcass-processing capabilities of the Plio-Pleistocene carnivores of South Africa. ROC data do a good job of separating taxa by dietary category, revealing possible differences in the carcass-processing abilities of fossil and modern members of some extant species, and confirming that Chasmaporthetes was probably a hypercarnivore and not a durophage like the modern hyenas. Contrary to previous hypotheses, sabertooth felids do not appear to have been more hypercarnivorous than modern felids based on these data.     

在中国云南发现狒狒化石

现存狒狒类(Papionin)生活于非洲(如Papio和Theropithoan)、亚洲(如Macaca)和北非(M.sylvanas)。在上新世和更新世,Theropithecus经历了从非洲到亚洲的扩散过程,在印度发现了类似化石。这次在云南中甸金沙江附近发现的下更新世狒狒化石(Papio)证明,如同亚洲猕猴和现代人类祖先一样,非洲狒狒类(Papio和Theropithecus)在同一时期从非洲扩散到亚洲。所不同的是它们没有像猕猴和人类一样生存下来。这次化石的发现对于研究以下生物学问题提供了重要依据1)探讨旧大陆猴类在上新—更新世从非洲到亚洲的扩散过程;2)研究不同旧大陆猴类的进化和环境适应性;3)为现代人类祖先在非洲—亚大陆的扩散研究提供证据;4)由于化石产地包括有人类祖先和其他动物的化石,因此,狒狒在亚洲的生态适应研究将为探讨人类在同一时期的生态适应提供证据。