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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. [ABSTRACT FROM AUTHOR]

Abstract: 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. [Copyright &y& Elsevier]

Abstract: 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 I1 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 P3. 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 M3. 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 I1, C1, P4 and P3. The I1 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 I1 versus I2. 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. [Copyright &y& Elsevier]

Among primates, age at first molar emergence is correlated with a variety of life history traits. Age at first molar emergence can therefore be used to broadly infer the life histories of fossil primate species. One method of determining age at first molar emergence is to determine the age at death of fossil individuals that were in the process of erupting their first molars. This was done for an infant partial mandible of Afropithecus turkanensis (KNM-MO 26) from the ∼17.5 Ma site of Moruorot in Kenya. A range of estimates of age at death was calculated for this individual using the permanent lateral incisor germ preserved in its crypt, by combining the number and periodicity of lateral enamel perikymata with estimates of the duration of cuspal enamel formation and the duration of the postnatal delay in the inception of crown mineralization. Perikymata periodicity was determined using daily cross striations between adjacent Retzius lines in thin sections of two A. turkanensis molars from the nearby site of Kalodirr. Based on the position of the KNM-MO 26 M1in relation to the mandibular alveolar margin, it had not yet undergone gingival emergence. The projected time to gingival emergence was estimated based on radiographic studies of M1eruption in extant baboons and chimpanzees.The estimates of age at M1emergence in KNM-MO 26 range from 28.2 to 43.5 months, using minimum and average values from extant great apes and humans for the estimated growth parameters. Even the absolute minimum value is well outside the ranges of extant large Old World monkeys for which there are data (12.5 to <25 months), but is within the range of chimpanzees (25.7 to 48.0 months). It is inferred, therefore, that A. turkanensis had a life history profile broadly like that of Pan. This is additional evidence to that provided by Sivapithecus parvada (Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptations, 1997, 173) that the prolonged life histories characteristic of extant apes were achieved early in the evolutionary history of the group. However, it is unclear at present whether life-history prolongation in apes represents the primitive catarrhine pace of life history extended through phyletic increase in body mass, or whether it is derived with respect to a primitive, size-adjusted life history that was broadly intermediate between those of extant hominoids and cercopithecoids. Life history evolution in primates as a whole may have occurred largely through a series of grade-shifts, with the establishment of fundamental life-history profiles early in the histories of major higher taxa. These may have included shifts that were largely body mass dependent, as well as those that occurred in the absence of significant changes in body mass. [Copyright &y& Elsevier]

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 toEquatorius. [Copyright &y& Elsevier]

Age at lower first molar (M 1) emergence is a commonly used proxy for inferring life-history scheduling in fossil primates, but its utility is dependent on knowing to what extent extant populations vary in this datum and how this variation correlates with the scheduling of life-history variables. Here, we address the first of these issues among extant chimpanzees. While age at M 1 emergence has been documented in several live individuals from the Kanyawara population of Pan troglodytes schweinfurthii in Uganda, it has been estimated for only one individual of Pan troglodytes verus , based on a deceased animal from the Taï Forest in Côte d'Ivoire. To further explore interpopulation variation in this variable in chimpanzees, and using dental histology, we calculated ages at death for two wild-shot individuals of P. t. verus with erupting M 1 , both collected in Liberia during the mid-1950s, and estimated ages at M 1 emergence from the ages at death. The overall range for these two individuals is ∼4.2–4.6 yr, compared with an age of ∼3.7 yr for the individual from the Taï Forest, and <2.5–3.3 yr for the several individuals of P. t. schweinfurthii. While the absolute range of ∼2 yr in these samples combined is little greater than in captive chimpanzees, the disparity between the samples of P. t. schweinfurthii and P. t. verus is striking, although it cannot be determined if this disparity represents a subspecies difference or simply population differences expressed in two different subspecies. While life-history data are unavailable for the population to which the Liberian chimpanzees belonged, the difference in M 1 emergence ages between these individuals and those from Kanyawara still suggests caution when attempting even broad life-history inference in fossil apes and hominins based on age at M 1 emergence. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Abstract 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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 ofSivapithecus 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 &y& Elsevier]

A dentate mandible and proximal femur of Mesopithecus pentelicus Wagner, 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. [ABSTRACT FROM AUTHOR]

Abstract: 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 M1 from Gorilla (2.31 years) and M1s 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. [Copyright &y& Elsevier]