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Understanding the biogeography and evolution of Miocene catarrhines relies on accurate specimen provenience. It has long been speculated that some catarrhine specimens among the early collections from Miocene sites in Kenya have incorrect provenience data. The provenience of one of these, the holotype of Equatorius africanus (NHM M16649), was previously revised based on x-ray fluorescence spectroscopy. Here we use nondestructive portable x-ray fluorescence spectroscopy to test the provenience of additional catarrhine specimens that, based on morphology and/or incomplete provenience information, were identified as possibly having been collected at a different site from that recorded at museum accession. We test the hypotheses that two specimens purportedly from the Early Miocene site of Rusinga (KNM-RU 1681 and KNM-RU 1999) are instead from Maboko, three specimens purportedly from the Middle Miocene site of Fort Ternan (KNM-FT 8, KNM-FT 41, and KNM-FT 3318) are instead from Songhor, and one specimen accessioned as being from Songhor (KNM-SO 5352) is from that site. Elemental data reveal that two of the specimens (KNM-FT 3318 and KNM-RU 1681) are likely to have been collected at sites other than their museum-accessioned provenience, while two others (KNM-RU 1999, and KNM-FT 41) were confirmed to have correct provenience. Results for both KNM-FT 8 and KNM-SO 5352, while somewhat equivocal, are best interpreted as supporting their accessioned provenience. Our results have implications for the distribution of certain catarrhine species during the Miocene in Kenya. Confirmation of the provenience of the specimens also facilitates taxonomic attribution, and resulted in additions to the morphological characterizations of some species. The protocol presented here has potential for wider application to assessing questions of provenience for fossils from other locations and periods., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

A dentate mandible and proximal femur of Mesopithecus pentelicusWagner, 1839 are described from the Shuitangba lignite mine in Zhaotong Prefecture, northeastern Yunnan Province, China. The remains were retrieved from sediments just below those that yielded a juvenile Lufengpithecus cranium and are dated at about ∼6.4 Ma. The mandible and proximal femur were found in close proximity and are probably of the same individual. The lower teeth are metrically and morphologically closely comparable with those of confirmed M. pentelicus from Europe, and on this basis, the specimen is assigned to this species. The anatomy of the proximal femur indicates that the Shuitangba Mesopithecus was a semiterrestrial quadruped that engaged in a range of mostly arboreal activities, including walking, climbing, and occasional leaping, with an abducted hip joint. The Shuitangba Mesopithecus is dentally typical for the genus but may have been more arboreal than previously described for M. pentelicus. M. pentelicus is well known from late Miocene (MN 11-12) sites in Europe and southwest Asia. Its estimated average rate of dispersal eastward was relatively slow, although it could have been episodically more rapid. The presence of a colobine, only slightly lower in the same section at Shuitangba that produced Lufengpithecus, is one of the only two well-documented instances of the near or actual co-occurrence of a monkey and ape in the Miocene of Eurasia. At Shuitangba, M. pentelicus occupied a freshwater-margin habitat with beavers, giant otters, swamp rabbits, and many aquatic birds. The presence of M. pentelicus in southwest China near the end of the Miocene further attests to the ecological versatility of a species long recognized as widespread and adaptable. The modern colobines of Asia, some or all of which are probable descendants of Mesopithecus, have gone on to inhabit some of the most highly seasonal and extreme habitats occupied by nonhuman primates., Competing Interests: Conflict of interest The authors declare no competing interests., (Copyright © 2020 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Here, we report on a new collection of mostly isolated molars of a colobine monkey from near Hasnot on the Potwar Plateau of Pakistan. The specimens are from three late Miocene localities, with ages constrained to between 7.9 and 7.1 Ma. Morphological and metrical comparisons of the new Hasnot molars with those of previously recognized Mesopithecus species and living Asian colobines lead to the conclusion that the Hasnot colobine is most probably Mesopithecus, but not Mesopithecus pentelicus. The most morphologically distinctive aspect of the Hasnot specimens is the lower third molars, which exhibit large and bulbous protoconids set off by deeply incised mesial buccal and median buccal clefts and large, broad distobuccally placed hypoconulids. Colobine specimens previously recovered from the Potwar Plateau have been assigned to Mesopithecus sivalensis, but because these specimens have not yet been fully described, a detailed comparison with the new Hasnot specimens is not yet possible. For these reasons, we assign the new Hasnot colobine fossils to cf. Mesopithecus sp. Mesopithecus was one of the most widespread and successful of late Miocene primates. As a colobine equipped with features of the molar teeth, limbs, and, presumably, gut enabling it to succeed in more highly seasonal woodland environments, Mesopithecus was able to rapidly disperse into and adapt to the conditions in South Asia brought about by profound climatic and concomitant environmental change during the latest Miocene., Competing Interests: Conflict of interest The authors state that they have no conflict of interest., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Kamoyapithecus hamiltoni is a potential early hominoid species described from fragmentary dentognathic specimens from the Oligocene site of Losodok (Turkana Basin, northwestern Kenya). Other catarrhine dental materials have been recovered at Losodok, but were not initially included in the Kamoyapithecus hypodigm. Here we present descriptions of the unpublished canine and incisor specimens from Losodok, and revisit the published specimens in light of recent changes in understanding of hominoid anterior dental evolution. The new fossils include a canine (KNM-LS 18354) that is morphologically compatible with being a female of Kamoyapithecus (KNM-LS 8). Randomization analyses of both Gorilla gorilla and middle Miocene Griphopithecus alpani demonstrate that the size difference between KNM-LS 8 and KNM-LS 18354 is also compatible with their being male and female canines of the same species. Significantly, a canine tip (KNM-LS 18352) attributed to Kamoyapithecus documents the distinctive burin tip morphology now recognized as characterizing Proconsul sensu stricto, which may indicate a close relationship between Kamoyapithecus and Proconsul. We also re-examined the enigmatic KNM-LS 1, a smaller lower canine assumed to derive from Losodok but for which historical provenience data are completely lacking. Elemental data derived from portable X-ray fluorescence spectroscopy show that KNM-LS 1 is almost certainly from Losodok rather than from nearby Miocene sites (i.e., Moruorot, Esha, Kalodirr). KNM-LS 1 displays a nyanzapithecine-like morphology and is shown by randomization analyses to be too small to be associated with the Kamoyapithecus canines. This demonstrates that there is a second hominoid taxon present at Losodok that records one of the earliest occurrences of the Nyanzapithecinae., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The Late Miocene sequence at Haritalyangar, Himachal Pradesh, India, has produced abundant remains of the hominid Sivapithecus and the sivaladapids Sivaladapis and Indraloris. Also recovered from these sediments is an isolated and worn upper molar that was made the holotype of Krishnapithecus krishnaii and assigned to the Pliopithecoidea. However, the heavy wear and absence of definitive pliopithecoid features on the tooth rendered the assignment to this superfamily unconvincing. Here, we describe two lower molars from Haritalyangar that bear unmistakable pliopithecoid features and that are plausibly assignable to the same species as the type specimen of K. krishnaii. They convincingly demonstrate for the first time the presence of the Pliopithecoidea in South Asia. The new molars also reveal that K. krishnaii was perhaps the largest known pliopithecoid and that it possessed highly derived postcanine dental morphology. Because of its highly derived nature, it is difficult to determine its relationships within Pliopithecoidea, but a sister taxon relationship with either the Dionysopithecidae or Pliopithecinae is equally plausible; it is only distantly related to the Crouzeliinae. It is sufficiently distinct, however, from all other pliopithecoids to warrant placement in a separate family., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

For more than 80 years, Proconsul has held a pivotal position in interpretations of catarrhine evolution and hominoid diversification in East Africa. The majority of what we 'know' about Proconsul, however, derives from abundant younger fossils found at the Kisingiri localities on Rusinga and Mfangano Islands rather than from the smaller samples found at Koru--the locality of the type species, Proconsul africanus--and other Tinderet deposits. One outcome of this is seen in recent attempts to expand the genus "Ugandapithecus" (considered here a junior subjective synonym of Proconsul), wherein much of the Tinderet sample was referred to that genus based primarily on differentiating it from the Kisingiri specimens rather than from the type species, P. africanus. This and other recent taxonomic revisions to Proconsul prompted us to undertake a systematic review of dentognathic specimens attributed to this taxon. Results of our study underscore and extend the substantive distinction of Tinderet and Ugandan Proconsul (i.e., Proconsul sensu stricto) from the Kisingiri fossils, the latter recognized here as a new genus. Specimens of the new genus are readily distinguished from Proconsul sensu stricto by morphology preserved in the P. africanus holotype, but also in I(1)s, lower incisors, upper and lower canines, and especially mandibular characteristics. A number of these differences are more advanced among Kisingiri specimens in the direction of crown hominoids. Proconsul sensu stricto is characterized by a suite of unique features that strongly unite the included species as a clade. There have been decades of contentious debate over the phylogenetic placement of Proconsul (sensu lato), due in part to there being a mixture of primitive and more advanced morphology within the single genus. By recognizing two distinct clades that, in large part, segregate these character states, we believe that better phylogenetic resolution can be achieved., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

The fossil ape Lufengpithecus is known from a number of late Miocene sites in Yunnan Province in southern China. Along with other fossil apes from South and Southeast Asia, it is widely considered to be a relative of the extant orangutan, Pongo pygmaeus. It is best represented at the type site of Shihuiba (Lufeng) by several partial to nearly complete but badly crushed adult crania. There is, however, an additional, minimally distorted cranium of a young juvenile from a nearly contemporaneous site in the Yuanmou Basin, which affords the opportunity to better assess the relationships between Lufengpithecus and Pongo. Comparison with similarly aged juvenile skulls of extant great apes reveals no features suggesting clear affinities to orangutans, and instead reveals a morphological pattern largely consistent with a stem member of the hominid (great ape and human) clade. The existence at this time of other hominids in South Asia (Sivapithecus) and Southeast Asia (Khoratpithecus) with clear craniofacial affinities to Pongo suggests both more diversity among Asian Late Miocene apes and more complex patterns of dispersal than previously supposed. Major differences in the associated mammal faunas from the southern China sites and those from South and Southeast Asia are consistent with these findings and suggest more than one dispersal route of apes into East Asia earlier in the Miocene.

Sexual size dimorphism in the postcanine dentition of the late Miocene hominoid Lufengpithecus lufengensis exceeds that in Pongo pygmaeus, demonstrating that the maximum degree of molar size dimorphism in apes is not represented among the extant Hominoidea. It has not been established, however, that the molars of Pongo are more dimorphic than those of any other living primate. In this study, we used resampling-based methods to compare molar dimorphism in Gorilla, Pongo, and Lufengpithecus to that in the papionin Mandrillus leucophaeus to test two hypotheses: (1) Pongo possesses the most size-dimorphic molars among living primates and (2) molar size dimorphism in Lufengpithecus is greater than that in the most dimorphic living primates. Our results show that M. leucophaeus exceeds great apes in its overall level of dimorphism and that L. lufengensis is more dimorphic than the extant species. Using these samples, we also evaluated molar dimorphism and taxonomic composition in two other Miocene ape samples--Ouranopithecus macedoniensis from Greece, specimens of which can be sexed based on associated canines and P(3)s, and the Sivapithecus sample from Haritalyangar, India. Ouranopithecus is more dimorphic than the extant taxa but is similar to Lufengpithecus, demonstrating that the level of molar dimorphism required for the Greek fossil sample under the single-species taxonomy is not unprecedented when the comparative framework is expanded to include extinct primates. In contrast, the Haritalyangar Sivapithecus sample, if itrepresents a single species, exhibits substantially greater molar dimorphism than does Lufengpithecus. Given these results, the taxonomic status of this sample remains equivocal.

The sample of the less common hominoid species at Paşalar, Kenyapithecus kizili, is characterized by a number of unusual attributes. All ten of the upper central incisors attributed to this species show a distinct, identical pattern of two linear enamel hypoplasias. The two hypoplasias occur on the same portion of the labial crown face, revealing that the two hypoplasia-causing events occurred at the same stage of development in all individuals. The morphology of the two hypoplasias and the amount of time between them, as determined by both their separation and counts of perikymata, are also the same on all teeth. In addition, all of the approximately 70 teeth assigned to K. kizili appear to come from young adults based on degrees of wear; there are no younger or older individuals (diagnostic morphology at most tooth positions would be evident even with heavy wear). Thus, all of the K. kizili individuals (minimum number of individuals is nine: seven males, two females) appear to have died at essentially the same age. It is concluded that the most plausible interpretation of all these features is that the incisor hypoplasias were caused by the same two events in all the K. kizili individuals and that these individuals therefore represent a single birth cohort. As such, and because they died at essentially the same age, they would also have died at the same time, which is consistent with the catastrophic nature of the Paşalar deposits. The number of coincidences needed to explain all of the attributes of the K. kizili sample if these animals were born in, and died in, different years seems highly improbable. Moreover, the lack of a typical age-class structure for the K. kizili sample, or any age-class structure at all beyond the one age class of young adult, strongly suggests that the species was not resident in the area that contributed to the Paşalar accumulation, and that K. kizili was not permanently sympatric with the other Paşalar hominoid, Griphopithecus alpani. Rather, the nine K. kizili individuals must have been transients in, or recent immigrants to, the area at the time of the events that led to the formation of the site. Recent observations on social associations in male chimpanzees offer at least a possible interpretive framework to explain this unprecedented occurrence in the primate fossil record.

A new species of fossil hominoid is described from the middle Miocene deposits at Paşalar, Turkey. It is the less common of the two Paşalar species discussed by Martin and Andrews (1993), making up approximately 10% of the individuals in the Paşalar hominoid sample according to analyses of the minimum number of individuals. To the diagnostic features of I(1) described by Alpagut et al. (1990) and Martin and Andrews (1993) can now be added further diagnostic features of all the anterior teeth, as well as both upper premolars and P(3). These include discrete, nonmetric features and metric differences at all the noted tooth positions. Attempts to distinguish the upper and lower molars of the two species have so far been unsuccessful, with the possible exception of M(3). The morphology of the new species is similar in most respects to that of Kenyapithecus wickeri from Fort Ternan, especially concerning maxillary morphology. They share robust and moderately deep maxillary alveolar processes, a restricted maxillary sinus with an elevated and uncomplicated floor, lacking the compartmentalization evident to varying degrees in many other taxa, and a zygomatic process that originates and turns laterally fairly high above the alveolar margin. There are also a number of distinctive similarities in the dentition, particularly for I(1), C(1), P(4) and P(3). The I(1) morphology in particular, with greatly hypertrophied lingual marginal ridges bounding a uniformly thickened basal crown area, is distinctive among Miocene hominoids. All of these similarities serve to reinforce the differences noted by others between the derived morphology of K. wickeri and the more primitive morphology of Equatorius africanus from Maboko and Kipsaramon. The new species differs from K. wickeri in morphological details of most of the anterior and premolar teeth that are known for both species, despite the general morphological similarity, and in the size of I(1) versus I(2). One striking feature of the new species is a relatively large incisive fossa, although it cannot be determined if this is associated with an open palatine fenestra, as in many early Miocene hominoids, or a minimally overlapping palate and nasoalveolar clivus, as in some middle and late Miocene hominoids.

During the past decade, studies of enamel development have provided a broad temporal and geographic perspective on evolutionary developmental biology in Miocene hominoids. Here we report some of the first data for molar crown development in one hominoid genus, Sivapithecus. The data are compared to a range of extant and extinct hominoids. Crown formation times (CFTs), daily rates of enamel secretion (DSR), Retzius line number and periodicity, and relative enamel thickness (RET) were calculated in a mandibular first molar of Sivapithecus parvada and a maxillary first molar of Sivapithecus indicus from the Siwalik sequence of Pakistan. A CFT of 2.40 years for the protoconid of S. parvada and 2.25 years for the protocone of S. indicus lie within the range of first molar (M1) formation times for the majority of Miocene hominoids (1.96-2.40 years, excluding Proconsul heseloni), and are similar to an M(1) from Gorilla (2.31 years) and M(1)s from Pan (2.22-2.39 years). This is unlike the longer CFTs in modern humans, which appear to be linked with their extended growth period. In contrast to extant great apes and humans, daily rates of enamel secretion are rapid in the Sivapithecus M1s during the early stages of growth, which seems to be a common pattern for most Miocene apes. The rapid accumulation of cuspal enamel in the Sivapithecus molars produced thicker enamel than either Pan or Gorilla in a comparable period of time. Future studies on larger samples of living and fossil hominoids are needed to clarify trends in crown development, which may be better understood in the context of life history strategies coupled with good data on body mass and brain size.

The earliest record of fossil apes outside Africa is in the latest early Miocene of Turkey and eastern Europe. There were at least 2, and perhaps 4, species of ape, which were found associated with subtropical mixed environments of forest and more open woodland. Postcranial morphology is similar to that of early Miocene primates and indicates mainly generalized arboreal quadrupedal behaviours similar to those of less specialized New World monkeys such as Cebus. Robust jaws and thick enamelled teeth indicate a hard fruit diet. The 2 best known species of fossil ape are known from the site of Paşalar in Turkey. They have almost identical molar and jaw morphology. Molar morphology is also similar to that of specimens from Germany and Slovakia, but there are significant differences in the anterior teeth of the 2 Paşalar species. The more common species, Griphopithecus alpani, shares mainly primitive characters with early and middle Miocene apes in Africa, and it is most similar phenetically to Equatorius africanus from Maboko Island and Kipsaramon. The second species is assigned to a new species of Kenyapithecus, an African genus from Fort Ternan in Kenya, on the basis of a number of shared derived characters of the anterior dentition, and it is considered likely that there is a phylogenetic link between them. The African sites all date from the middle Miocene, similar in age to the Turkish and European ones, and the earliest emigration of apes from Africa coincides with the closure of the Tethys Sea preceding the Langhian transgression. Environments indicated for the African sites are mixtures of seasonal woodlands with some forest vegetation. The postcrania of both African taxa again indicate generalized arboreal adaptation but lacking specialized arboreal function. This middle Miocene radiation of both African and non-African apes was preceded by a radiation of arboreal catarrhine primates in the early Miocene, among which were the earliest apes. The earliest Miocene apes in the genus Proconsul and Rangwapithecus were arboreal, and because of their association with the fruits of evergreen rain forest plants at Mfwangano Island, it would appear that they were forest adapted, i.e. were living in multi-storied evergreen forest. The same or similar species of the same genera from Rusinga Island, together with other genera such as Nyanzapithecus and the small ape Limnopithecus, were associated with plants and animals indicating seasonal woodland environments, probably with gallery forest forming corridors alongside rivers. While the stem ancestors of the Hominoidea were almost certainly forest adapted, the evidence of environments associated with apes in the later part of the early Miocene and the middle Miocene of East Africa indicates more seasonal woodlands, similar to those reconstructed for the middle Miocene of Paşalar in Turkey. This environmental shift was probably a requisite for the successful emigration of apes out of Africa and made possible later movement between the continents for much of the middle Miocene, including possible re-entry of at least one ape lineage back into Africa., (Copyright 2007 S. Karger AG, Basel.)

Several new postcranial elements of Sivapithecus from the Siwaliks of Pakistan are described. These include a distal femur from the U-level of the Dhok Pathan Formation, a navicular from the Chinji Formation, and seven manual and pedal phalanges from the Nagri Formation. The functional morphology of these elements adds new detail to the reconstruction of Sivapithecus positional behavior. Femoral cross-sectional geometry indicates that the shaft was adapted to support mediolaterally directed loading. Femoral condylar asymmetry and a broad but shallow trochlea are distinctly ape-like, revealing capabilities for both rotation and withstanding eccentric loading in the knee. The navicular is characterized by features relating to a broad mid-tarsus and broad distal articulations for the cuneiforms. It also lacks a navicular tubercle as in Pongo. These features suggest that the foot was capable of a powerful grip on large supports, with an inversion/supination capability that would permit foot placement in a variety of positions. The morphology of the new phalanges, including evidence for a relatively large pollex, similarly suggests powerful grasping, consistent with prior evidence from the hallux and tarsus. The functional features of the new specimens permit refinement of previous interpretations of Sivapithecus positional capabilities. They suggest a locomotor repertoire dominated by pronograde activities and also such antipronograde activities as vertical climbing and clambering, but not by antipronograde suspensory activities as practiced by extant apes., (Copyright 2002 Elsevier Science Ltd. All rights reserved.)

Forty-one isolated large hominoid teeth, as well as most of the mandibular and three maxillary teeth associated with a partial skeleton, were recovered from middle Miocene Muruyur sediments near Kipsaramon in the Tugen Hills, Baringo District, Kenya. The isolated teeth were collected as surface finds and the skeleton was excavated in situ at locality BPRP#122 dated between 15.58 Ma and 15.36 Ma. The majority of the teeth recovered at BPRP#122 are referable to a minimum of five individuals of the hominoid Equatorius africanus. Three of the teeth, however, are provisionally assigned to Nyanzapithecus sp. The new hominoids from Kipsaramon add to an increasing inventory of specimens that suggest greater large hominoid taxonomic diversity from the middle Miocene of Kenya than was previously recognized. It is suggested that there are two large-bodied hominoid species present at Mabako, only one of which is assignable to Equatorius., (Copyright 2002 Academic Press.)

The article reports that Middle Siwalik sediments exposed at Haritalyangar. India are best known for the diversity of fossil primates that no longer inhabit the Sub-Himalaya, including the late surviving large hominoids Sivapithecus and Indopithecus as well as primitive lemuriform primate. The fossils come from the middle level of the Haritalyangar stratigraphic section, which has a lithostratigraphic profile typical of the Middle Siwaliks of the Potwar Plateau in Pakistan.

Describes a partial hominoid skeleton from a Middle Miocene site located west of Lake Baringo in central Kenya that seems to be an early member of the great ape-human clade. The two species of Kenyapithecus that are currently recognized; Description of this new genus; Etymology.