Portable x-ray fluorescence spectroscopy geochemical sourcing of Miocene primate fossils from Kenya.

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.

Earliest giant panda false thumb suggests conflicting demands for locomotion and feeding.

Of the many peculiarities that enable the giant panda (Ailuropoda melanoleuca), a member of the order Carnivora, to adapt to life as a dedicated bamboo feeder, its extra "thumb" is arguably the most celebrated yet enigmatic. In addition to the normal five digits in the hands of most mammals, the giant panda has a greatly enlarged wrist bone, the radial sesamoid, that acts as a sixth digit, an opposable "thumb" for manipulating bamboo. We report the earliest enlarged radial sesamoid, already a functional opposable "thumb," in the ancestral panda Ailurarctos from the late Miocene site of Shuitangba in Yunnan Province, China. However, since the late Miocene, the "thumb" has not enlarged further because it must be balanced with the constraints of weight bearing while walking in a plantigrade posture. This morphological adaptation in panda evolution thus reflects a dual function of the radial sesamoid for both bamboo manipulation and weight distribution. The latter constraint could be the main reason why the panda's false thumb never evolved into a full digit. This crude "thumb" suggests that the origin of the panda's dedicated bamboo diet goes back to as early as 6-7 Ma.

Mesopithecus pentelicus from Zhaotong, China, the easternmost representative of a widespread Miocene cercopithecoid species.

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.

Age at first molar emergence in Pan troglodytes verus and variation in the timing of molar emergence among free-living chimpanzees.

Age at lower first molar (M1) 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 M1 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 M1, both collected in Liberia during the mid-1950s, and estimated ages at M1 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 M1 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 M1 emergence.

New fossils of Mesopithecus from Hasnot, Pakistan.

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.

Evidence for increased hominid diversity in the Early to Middle Pleistocene of Indonesia.

Since the first discovery of Pithecanthropus (Homo) erectus by E. Dubois at Trinil in 1891, over 200 hominid dentognathic remains have been collected from the Early to Middle Pleistocene deposits of Java, Indonesia, forming the largest palaeoanthropological collection in South East Asia. Most of these fossils are currently attributed to H. erectus. However, because of the substantial morphological and metric variation in the Indonesian assemblage, some robust specimens, such as the partial mandibles Sangiran 5 and Sangiran 6a, were formerly variably allocated to other taxa (Meganthropus palaeojavanicus, Pithecanthropus dubius, Pongo sp.). To resolve the taxonomic uncertainty surrounding these and other contentious Indonesian hominid specimens, we used occlusal fingerprint analysis (OFA) to reconstruct their chewing kinematics; we also used various morphometric approaches based on microtomography to examine the internal dental structures. Our results confirm the presence of Meganthropus as a Pleistocene Indonesian hominid distinct from Pongo, Gigantopithecus and Homo, and further reveal that Dubois's H. erectus paratype molars from 1891 are not hominin (human lineage), but instead are more likely to belong to Meganthropus.

Hominoid anterior teeth from the late Oligocene site of Losodok, Kenya.

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.

A highly derived pliopithecoid from the Late Miocene of Haritalyangar, India.

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.

A systematic revision of Proconsul with the description of a new genus of early Miocene hominoid.

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.

A partial hominoid innominate from the Miocene of Pakistan: description and preliminary analyses.

We describe a partial innominate, YGSP 41216, from a 12.3 Ma locality in the Siwalik Group of the Potwar Plateau in Pakistan, assigned to the Middle Miocene ape species Sivapithecus indicus. We investigate the implications of its morphology for reconstructing positional behavior of this ape. Postcranial anatomy of extant catarrhines falls into two distinct groups, particularly for torso shape. To an extent this reflects different although variable and overlapping positional repertoires: pronograde quadrupedalism for cercopithecoids and orthogrady for hominoids. The YGSP innominate (hipbone) is from a primate with a narrow torso, resembling most extant monkeys and differing from the broader torsos of extant apes. Other postcranial material of S. indicus and its younger and similar congener Sivapithecus sivalensis also supports reconstruction of a hominoid with a positional repertoire more similar to the pronograde quadrupedal patterns of most monkeys than to the orthograde patterns of apes. However, Sivapithecus postcranial morphology differs in many details from any extant species. We reconstruct a slow-moving, deliberate, arboreal animal, primarily traveling above supports but also frequently engaging in antipronograde behaviors. There are no obvious synapomorphic postcranial features shared exclusively with any extant crown hominid, including Pongo.

Juvenile hominoid cranium from the late Miocene of southern China and hominoid diversity in Asia.

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.

Dental development and life history in living African and Asian apes.

Life-history inference is an important aim of paleoprimatology, but life histories cannot be discerned directly from the fossil record. Among extant primates, the timing of many life-history attributes is correlated with the age at emergence of the first permanent molar (M1), which can therefore serve as a means to directly compare the life histories of fossil and extant species. To date, M1 emergence ages exist for only a small fraction of extant primate species and consist primarily of data from captive individuals, which may show accelerated dental eruption compared with free-living individuals. Data on M1 emergence ages in wild great apes exist for only a single chimpanzee individual, with data for gorillas and orangutans being anecdotal. This paucity of information limits our ability to make life-history inferences using the M1 emergence ages of extinct ape and hominin species. Here we report reliable ages at M1 emergence for the orangutan, Pongo pygmaeus (4.6 y), and the gorilla, Gorilla gorilla (3.8 y), obtained from the dental histology of wild-shot individuals in museum collections. These ages and the one reported age at M1 emergence in a free-living chimpanzee of approximately 4.0 y are highly concordant with the comparative life histories of these great apes. They are also consistent with the average age at M1 emergence in relation to the timing of life-history events in modern humans, thus confirming the utility of M1 emergence ages for life-history inference and providing a basis for making reliable life-history inferences for extinct apes and hominins.

Root growth during molar eruption in extant great apes.

While there is gradually accumulating knowledge about molar crown formation and the timing of molar eruption in extant great apes, very little is known about root formation during the eruption process. We measured mandibular first and second molar root lengths in extant great ape osteological specimens that died while either the first or second molars were in the process of erupting. For most specimens, teeth were removed so that root lengths could be measured directly. When this was not possible, roots were measured radiographically. We were particularly interested in the variation in the lengths of first molar roots near the point of gingival emergence, so specimens were divided into early, middle and late phases of eruption based on the number of cusps that showed protein staining, with one or two cusps stained equated with immediate post-gingival emergence. For first molars at this stage, Gorilla has the longest roots, followed by Pongo and Pan. Variation in first molar mesial root lengths at this stage in Gorilla and Pan, which comprise the largest samples, is relatively low and represents no more than a few months of growth in both taxa. Knowledge of root length at first molar emergence permits an assessment of the contribution of root growth toward differences between great apes and humans in the age at first molar emergence. Root growth makes up a greater percentage of the time between birth and first molar emergence in humans than it does in any of the great apes.

Beyond Gorilla and Pongo: alternative models for evaluating variation and sexual dimorphism in fossil hominoid samples.

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.

A new hominoid species from the middle Miocene site of Pasalar, Turkey.

A new species of fossil hominoid is described from the middle Miocene deposits at Pasalar, Turkey. It is the less common of the two Pasalar species discussed by Martin and Andrews (1993), making up approximately 10% of the individuals in the Pasalar 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.

Identification of a single birth cohort in Kenyapithecus kizili and the nature of sympatry between K. kizili and Griphopithecus alpani at Pasalar.

The sample of the less common hominoid species at Pasalar, 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 Pasalar 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 Pasalar accumulation, and that K. kizili was not permanently sympatric with the other Pasalar 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.

Middle Miocene dispersals of apes.

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 Pasalar 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 Pasalar 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 Pasalar 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.

Molar crown formation in the Late Miocene Asian hominoids, Sivapithecus parvada and Sivapithecus indicus.

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.