New craniodental fossils of papionin monkeys from Cooper's D,South Africa

Papionin monkey fossils are common in the Plio‐Pleistocene aged karst cave deposits northwest of Johannesburg in South Africa. These deposits have yielded important primate and other vertebrate fauna since their discovery in the early part of the 20th century. In this article, we describe new primate cranial and dental specimens from excavations at the site of Cooper's D in the Sterkfontein Valley that date to around 1.5 million years ago. Unlike other localities in southern Africa, most of the new fossils are referred to Theropithecus oswaldi oswaldi, an extinct gramnivorous monkey related to the living gelada. Diagnostic features of T. o. oswaldi crania and teeth include large, thickly enameled molars with tall, columnar cusps, and high molar relief, an upright mandibular ramus, postorbital constriction, and anterior fusion of temporal lines. Also present in the new sample are teeth referred to Papio sp., which show low crowned bunodont molars, and a number of indeterminate papionin teeth and skull fragments. The presence of T. o. oswaldi at Cooper's D extends the list of known localities where the taxon is found, and may indicate the presence of an open, grassland environment in the area during the early Pleistocene. The abundance of theropith fossils at Cooper's suggests that Papio was not consistently the most common papionin in southern Africa over the past three million years. Am J Phys Anthropol 151:613–629, 2013. © 2013 Wiley Periodicals, Inc.     

Brief communication: Plio‐Pleistocene eagle predation on fossil cercopithecids from the Humpata Plateau,southern Angola

Recent studies suggest a large raptor such as the crowned eagle (Stephanaoetus coronatus) was responsible for collecting at least a portion of the primate fauna from the South African fossil site of Taung, including its lone hominin specimen. This taphonomic signature at Taung is currently regarded as a unique and, most likely, isolated case in primate and human evolution. However, the activities of large, carnivorous birds should also be detectable at other primate fossil localities in Africa if raptors have been a strong selective force throughout primate evolution. Over the last 60 years, a collection of extinct cercopithecids has been assembled from several cave breccias on the Humpata Plateau in southern Angola. The material, dated near the Plio‐Pleistocene boundary, includes an assortment of craniodental and postcranial remains variably assigned to Papio (Dinopithecus) cf. quadratirostris, Parapapio, Cercopithecoides, and Theropithecus. We compare the Angolan and Taung material to remains of extant primates killed by crowned eagles in the Ivory Coast's Tai National Park. Our analysis indicates that the size distribution and composition of fauna from the localities is quite similar and that there are striking consistencies in damage to the crania from each site. The absence of large bodied (>20 kg) primates and other mammalian taxa at the Taung hominin locality and Tai, and their rarity in Angola, combined with the strong likelihood that raptor nests were positioned near fissure openings at both fossil localities, provides additional support for eagle involvement. On the basis of this evidence, we conclude that at least some of the Angolan cercopithecids were most likely raptor prey and hypothesize that raptor predation has been a strong and perhaps underappreciated selective force during the course of primate evolution. Am J Phys Anthropol 2009. © 2009 Wiley‐Liss, Inc.     

The mid-face of lower Pleistocene hominins and its bearing on the attribution of SK 847 and StW 53

SK 847 and StW 53 have often been cited as evidence for early Homo in South Africa. To examine whether midfacial morphology is in agreement with these attributions, we analyze Euclidean distances calculated from 3-D coordinates on the maxillae of SK 847 and StW 53, as well as Australopithecus africanus (Sts 5, Sts 71), Paranthropus robustus (SK 46, SK 48, SK 52, SK 83), early Homo (KNM-ER 1813, KNM-ER 1805, KNM-ER 3733, KNM-WT 15000), P. boisei (KNM-ER 406, KNM-WT 17000, KNM-WT 17400), Gorilla gorilla (n = 116), Homo sapiens (n = 342), Pan paniscus (n = 21) and P. troglodytes (n = 65). Multivariate analyses separate extant hominoids suggesting we have captured taxonomic affinity. With the exception of SK 847 and SK 52, South African fossils tend to cluster together. P. robustus differs substantially from East African robust megadonts. SK 847 and StW 53 resemble the East African Homo specimens that are the most australopith-like, such as KNM-WT 15000 and KNM-ER 1813. The resemblance between StW 53 and Homo is driven partly by similarities in maxillary size. When distances are scaled, StW 53 aligns with A. africanus, while SK 847 clusters primarily with early Homo.     

Modeling the Biogeography of Fossil Baboons

We use a model of modern baboon socio-ecology to explore the behavioral ecology and biogeography of the extinct Plio-Pleistocene baboons (genera Parapapio, Gorgopithecus, Dinopithecus, and Papio). The model is based on the way climate affects the baboons’ time budgets, and focuses on intersite variability in behavior rather than on species-typical patterns of behavior, as most previous approaches have done. We use climate estimates for individual fossil sites based on matching modern habitats using faunal profiles and estimates of individual species’ body masses given in the literature. The model allows us to examine the minimum and maximum sizes of groups that individual species would have been able to support at particular localities, and hence the biogeography of a species on a continental scale. In doing so, the model allows us to examine which variables are most responsible for limiting a species’ ecological and biogeographic flexibility, and through this to explore a species’ capacity for coping with climate change. Feeding time is identified as the main constraint. In general, large-bodied species would have had more difficulty surviving in as wide a range of habitats as smaller-bodied species, and this may explain the limited geographical distribution of large-bodied baboons such as Gorgopithecus and Dinopithecus.     

Partial skeleton of Theropithecus brumpti (Primates, Cercopithecidae) from the Chemeron Formation of the Tugen Hills, Kenya

Here we describe a complete skull and partial skeleton of a large cercopithecoid monkey (KNM-TH 46700) discovered in the Chemeron Formation of the Tugen Hills at BPRP Site #152 (2.63 Ma). Associated with the skeleton was a mandible of an infant cercopithecoid (KNM-TH 48364), also described here. KNM-TH 46700 represents an aged adult female of Theropithecus brumpti, a successful Pliocene papionin taxon better known from the Omo Shungura Formation in Ethiopia and sites east and west of Lake Turkana, Kenya. While the morphology of male T. brumpti is well-documented, including a partial skeleton with both cranial and postcranial material, the female T. brumpti morphotype is not well-known. This skeleton represents some of the first associated evidence of cranial and postcranial female T. brumpti remains. In addition to the complete skull, postcranial material includes elements of the axial skeleton and lower limb. While aspects of the skeleton conform to those of specimens previously assigned to T. brumpti, other features on the femur and tibia appear to differ from those previously described for this species. It is unclear whether these differences represent general variation within the T. brumpti population, variation between the sexes in T. brumpti, or the incorrect assignment of previous isolated hindlimb specimens. In total, the observable morphological features of the hindlimb suggest that KNM-TH 46700 was a terrestrial quadruped similar to modern savannah baboons (Papio). From the available evidence, it is difficult to assess whether or not KNM-TH 46700 frequently engaged in the specialized squatting and shuffling behavior observed in extant geladas (Theropithecus gelada).     

Occurrence of human-pathogenic Enterocytozoon bieneusi,Giardia duodenalis and Cryptosporidium genotypes in laboratory macaques in Guangxi,China

Captive nonhuman primates have been identified as common hosts of Enterocytozoon bieneusi, Giardia duodenalis, Cryptosporidium hominis, and Cyclospora spp., thus are potential reservoirs of some enteric parasites in humans. However, few studies have examined the source and human-infective potential of enteric parasites in laboratory nonhuman primates. In the present work, 205 fecal specimens were collected from three groups of captive Macaca fascicularis kept in different densities in a laboratory animal facility in Guangxi, China, and examined by PCR for E. bieneusi, G. duodenalis, Cryptosporidium spp., and Cyclospora spp. The infection rates of E. bieneusi and G. duodenalis were 11.3% and 1.2% in Group 1 (young animals kept individually; n = 168), 72.2% and 11.1% in Group 2 (young animals kept in groups; n = 18), and 31.6% and 5.3% in Group 3 (adults kept in groups; n = 19), respectively. Sequence analysis of PCR products showed the presence of five E. bieneusi genotypes, with genotype D (in 16/36 genotyped specimens) and a new genotype (in 15/36 genotyped specimens) as the dominant genotypes. All five E. bieneusi genotypes belonged to the zoonotic group (Group 1). The G. duodenalis genotypes (assemblages AII and B) in five specimens and C. hominis subtype (IdA14) in one specimen were also known human-pathogens, although the Cyclospora seen in one animal appeared to be unique to macaque monkeys. The higher infection rate in younger animals reared in groups and common occurrence of zoonotic genotypes indicated that human-pathogenic E. bieneusi could be transmitted efficiently in captive nonhuman primates, and group-housing was a risk factor for transmission of zoonotic pathogens in young nonhuman primates in research facilities.     

Does the tropical agricultural matrix bear potential for primate conservation? A baseline study from Western Uganda

As for most large mammals, conservation research on primates usually focuses on protected areas, and yet not much is known about primate communities in land-use systems in the absence of hunting. Using line transects, we estimated population densities for four primate species in a mixed agroforest landscape up to a distance of 4000 m from a forest reserve, in a region where no primate hunting takes place. We then modelled encounter rates and cluster size in relation to landscape parameters by means of bivariate analysis and Generalised Linear Models (GLMs). Black-and-white colobus (Colobus guereza) were most common, with confidence intervals for density estimates of 31.1–62.8 individuals/km². Red-tailed monkeys (Cercopithecus ascanius) and blue monkeys (Cercopithecus mitis) occurred at 15.2–37.9 ind/km² and 13.9–36.7 ind/km² respectively, and chimpanzees (Pan troglodytes) at 1.0–2.8 ind/km². Chimpanzee nest numbers and distribution appeared to be significantly constrained by transect forest coverage, forest coverage within 500 m and distance from the main forest, whereas monkey sightings were generally less restricted by landscape variables. The considerable population density of monkeys suggests that, in the absence of hunting, mixed agroforest systems may play a relevant role in primate conservation and highlight that it is useful to consider primate ecology in land sharing approaches to conservation.     

A reassessment of the taxonomic status and phyletic relationships of Papio baringensis and Papio quadratirostris (Primates: Cercopithecidae)

Comparison of the skulls of Papio baringensis Leakey 1969 from the Chemeron Formation and P. quadratirostris Iwamoto 1982 from the Usno Formation with those of Theropithecus gelada, T. brumpti, T. darti, T. oswaldi, and several species of Papio indicate that the species from Chemeron and Usno exhibit all, or most, respectively, of the shared derived characters of Theropithecus. We propose that they be removed from Papio and placed in Theropithecus as T. baringensis and T. quadratirostris. Comparison of the specimens of T. baringensis and T. quadratirostris with those of T. brumpti indicate that the former two species have some of the derived characters of T. brumpti but are primitive in others. We propose that the three species form the following phyletic lineage: T. baringensis-T. quadratirostris-T. brumpti. With these referrals, there are now six species of the genus Theropithecus. Based on geologic grounds, the specimens from Chemeron are about 4.0 million years (m.y.), that from Usno between 3.3 and 3.4 m.y., and those of T. brumpti between 2.0 and 2.8 m.y. in age. We also show that the most complete specimen of Papio sp. nov. from Olduval Gorge belongs to T. oswaldi. With the removal of these specimens from Papio, the East African fossil record of this genus, apart from isolated teeth, comprises only 21 specimens, while that of Theropithecus comprises at least 300 specimens.     

Brief communication: New primate remains from the Miocene of Namibia,Southern Africa

Miocene primates from southern Africa are extremely rare. For this reason we wish to place on record several interesting new fossil primate specimens recently recovered from the Miocene sites of Berg Aukas and Harasib in the Otavi Mountain region of northern Namibia. The new finds consist of a virtually complete atlas vertebra from Berg Aukas attributable to the hominoid Otavipithecus namibiensis and two teeth and four postcranial fragments from Harasib referrable to Cercopithecoidea. The atlas vertebra exhibits anatomical characteristics intermediate between those of modern cercopithecoids and hominoids which may be indicative of a transition from pronograde to orthograde postures. The cercopithecoid remains show that the earliest Old World monkeys known from southern Africa were small, approximately the size of vervet monkeys. These new specimens are important because they provide the first evidence relating to possible positional behaviors of Otavipithecus and the earliest fossil record of cercopithecoids from southern Africa. © 1996 Wiley-Liss, Inc.     

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.     

The enigmatic molar from Gondolin, South Africa: Implications for Paranthropus paleobiology

The specific attribution of the large hominin M₂ (GDA-2) from Gondolin has significant implications for the paleobiology of Paranthropus. If it is a specimen of Paranthropus robustus it impacts that species' size range, and if it belongs to Paranthropus boisei it has important biogeographic implications. We evaluate crown size, cusp proportions and the likelihood of encountering a large-bodied mammal species in both East and South Africa in the Early Pleistocene. The tooth falls well outside the P. robustus sample range, and comfortably within that for penecontemporaneous P. boisei. Analyses of sample range, distribution and variability suggest that it is possible, albeit unlikely to find a M₂ of this size in the current P. robustus sample. However, taphonomic agents - carnivore (particularly leopard) feeding behaviors - have likely skewed the size distribution of the Swartkrans and Drimolen P. robustus assemblage. In particular, assemblages of large-bodied mammals accumulated by leopards typically display high proportions of juveniles and smaller adults. The skew in the P. robustus sample is consistent with this type of assemblage. Morphological evidence in the form of cusp proportions is congruent with GDA-2 representing P. robustus rather than P. boisei. The comparatively small number of large-bodied mammal species common to both South and East Africa in the Early Pleistocene suggests a low probability of encountering an herbivorous australopith in both. Our results are most consistent with the interpretation of the Gondolin molar as a very large specimen of P. robustus. This, in turn, suggests that large, presumptive male, specimens are rare, and that the levels of size variation (sexual dimorphism) previously ascribed to this species are likely to be gross underestimates.     

Fossil papio cranium from !Ncumtsa (Koanaka) Hills, western Ngamiland, Botswana

Three fossils, a cranium of Papio, a cercopithecid frontal bone, and a mandible of juvenile Papio, have been recovered from cave deposits in the !Ncumtsa (Koanaka) Hills of western Ngamiland, Botswana. These specimens are significant because well‐preserved crania of Papio are extremely rare in the fossil record outside of South Africa and because this is the first report of fossil primate cranial remains from Botswana. Thermoluminescence dating of surrounding cave matrix indicates an age of ≥317 ± 114 ka, within the Middle Pleistocene, although it may be older. Based on univariate and multivariate analyses, the adult !Ncumtsa specimen falls within the range of variation seen in extant forms of Papio, yet is distinct from any living species/subspecies and represents a new taxon, named here as a new subspecies of Papio hamadryas—Papio hamadryas botswanae. Am J Phys Anthropol, 2012. © 2012 Wiley Periodicals, Inc.     

Tsuga seed cones from the late Paleogene of southwestern China and their biogeographical and paleoenvironmental implications

Six Tsuga ovuliferous/seed cone impression fossils were discovered from the late Eocene (34.6 ± 0.8 Ma) Lawula Formation in Mangkang County, eastern Tibet and the early Oligocene (32 ± 1 Ma) lacustrine deposits in Lühe Basin, Nanhua County, Yunnan Province. These two fossil sites are both located in southwestern China, ∼800 km apart from each other. These fossils represent the oldest records of this genus in southwestern China, even earliest reliable macrofossil records of this genus in the world. These well-preserved seed cones provide sufficient materials for the establishment of Tsuga asiatica Wu et Zhou n. sp. to accommodate five specimens, leaving one to be assigned to T. cf. dumosa Eichler (cf. Wu et Zhou). Both qualitative and quantitative comparisons with other cone fossils and cones of all living species of the genus suggested that T. asiatica shares more similarities with one of the basal species of the genus T. heterophylla. The discovery of late Paleogene macrofossil records of Tsuga in southwestern China supports the previous hypothesis of the early disposal routes of this coniferous genus predicted by phylogenetic analysis. The elevation ranges and the climate requirements of living species that are closely related to our fossils suggest that the southeastern edge of the Qinghai-Tibetan Plateau should be much warmer, and wetter in late Paleogene than nowadays.     

Does trabecular bone structure within the metacarpal heads of primates vary with hand posture?

Reconstructing function from hominin fossils is complicated by disagreements over how to interpret primitively inherited, ape-like morphology. This has led to considerable research on aspects of skeletal morphology that may be sensitive to activity levels during life. We quantify trabecular bone morphology in three volumes of interest (dorsal, central, and palmar) in the third metacarpal heads of extant primates that differ in hand function: Pan troglodytes, Pongo pygmaeus, Papio anubis, and Homo sapiens. Results show that bone volume within third metacarpal heads generally matches expectations based on differences in function, providing quantitative support to previous studies. Pongo shows significantly low bone volume in the dorsal region of the metacarpal head. Humans show a similar pattern, as manipulative tasks mostly involve flexed and neutral metacarpo-phalangeal joint postures. In contrast, Pan and Papio have relatively high bone volume in dorsal and palmar regions, which are loaded during knuckle-walking/digitigrady and climbing, respectively. Regional variation in degree of anisotropy did not match predictions. Although trabecular morphology may improve behavioral inferences from fossils, more sophisticated quantitative strategies are needed to explore trabecular spatial distributions and their relationships to hand function.     

The alpha taxonomy of Australopithecus at Sterkfontein: The postcranial evidence

The possibility that the fossils attributed to Australopithecus africanus represent more than a single species is of significance because of the pivotal role that A. africanus has played in discussions about hominin evolution. The A. africanus hypodigm that is currently widely recognized evinces considerable variation in a number of craniodental characters, and this has led to speculation that more than one australopith taxon may be represented among the specimens from Sterkfontein. Although crania, mandibles and teeth have dominated these taxonomic discussions, the Sterkfontein postcranial remains also have been invoked. While several workers have proposed that some of the craniodental remains from Sterkfontein can be partitioned into two groups, there is a notable lack of agreement among them as to their actual sorting. Most of the craniodental observations that have been put forward in support of arguments for taxonomic heterogeneity of the Sterkfontein australopith assemblage have been subjective and anecdotal in nature. So too, the postcranial evidence that has been cited in support of more than one australopith species at Sterkfontein has been largely subjective, and limited to a small number of elements. The results of quantitative statistical analyses of the craniodental and postcranial fossils that have been undertaken to date are not necessarily consistent with the hypothesis of taxonomic heterogeneity.     

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.     

The systematic position of Hoplitomerycidae (Ruminantia) revisited

Hoplitomeryx Leinders was originally described only on cranial characters. The type specimens were found during the 1970's in karstic fissure fillings, most likely of Messinian age, in Gargano (Apulia, southeastern Italy), between Poggio Imperiale (41°49′30′ N, 15°21′58′ E) and Apricena (41°47′06′ N, 15°26′41′ E). During the 1990's, Hoplitomeryx remains were also discovered in the lower Tortonian layered calcarenites near Scontrone (Abruzzo, central Italy; 41°45′15.55′ N, 14°02′13.23′ E). The skull fragments, teeth, and jawbones from both localities have been examined. The dental characters had never been described before, and also some maxillaries and jawbones were not part of the original sample that was analyzed to establish the genus. Because they possess two lacrimal orifices and closed metatarsal gulley, hoplitomerycids have been linked more closely with Cervids, and accommodated in Cervoidea. A cladistic analysis of a character-taxon matrix of 121 morphological features (48 cranial, 51 dental and 22 postcranial characters) is performed. The analysis shows that hoplitomerycids stem either between antilocaprids and bovids, or antilocaprids and giraffids. They are not linked directly with cervids. Hoplitomerycids likely stemmed from a primitive ruminant stock, perhaps around 29 Ma when a land bridge connected the Abruzzo-Apulia platform with the Balkans across the Adriatic Sea. In the new land, hoplitomerycids developed a mosaic of apomorphic and homoplastic (convergent) character states that recall those found in other higher ruminants.