Geology and fauna of the middle Miocene hominoid site at Muruyur,Baringo district,Kenya

The middle Miocene sediments assigned to the Muruyur Beds have yielded abundant faunal remains which indicate an age somewhere near the early part of the middle Miocene, perhaps being earlier in time than Fort Ternan but probably coeval or slightly later than Maboko. Available radioisotopic age determinations suggest that the beds are between 13.5 and 14 m.y. old, which seems to be too young when compared with the biostratigraphic estimate. The importance of Muruyur Beds lies in their rich fossil content which includes hominoids of an age which is in general poorly represented in East Africa’s fossil record. This article places the fossil discoveries on record, and discusses their geological context.     

Miocene fossil cercopithecoids from Kenya

Fossil cercopithecoid material from Ngeringerowa, Ngorora, and Nakali, dated at between 8.5 and 10.5 m.y., is described. The specimens are the only cercopithecoid remains dated between 15 and 6 m.y. from sub-Saharan Africa. The mandible of asmall colobine from Ngeringerowa (similar in size to Colobus verus) is assigned to a new genus and species, Microcolobus tugenensis. Unlike other colobine genera, the symphysis of Microcolobus lacks an inferior transverse torus. A colobine lower M_{1 or 2} from Nakali is longer and narrower than molars of M. tugenensis, indicating that it may belong to a distinct taxon. A P₄ from Ngorora cannot be assigned confidently to subfamily, due to its unique metaconid morphology. The relationship between the new genus and other Miocene monkeys is considered.     

Brachypotherium (Perissodactyla,Rhinocerotidae) from the late Miocene of Samburu Hills,Kenya

Several isolated cheek teeth and mandibular specimens of Rhinocerotidae (Mammalia, Perissodactyla) from the upper Miocene Namurungule Formation in Samburu Hills, Kenya, are redescribed. Previously, these specimens had been identified as Chilotheridium pattersoni, Chilotheridium sp., Paradiceros mukirii, and Paradiceros sp. They are reidentified here as documenting the genus Brachypotherium based on their bucco-lingually broad molariform upper premolars with short crochet and flattened buccal walls on both upper and lower molars, the latter having a shallow external groove. Comparisons with other Brachypotherium species suggest that the present specimens belong to Brachypotherium sp. cf. B. minor. The presence of Brachypotherium in the Samburu Hills, at ca. 9.5 Ma, is concordant with the paleoenvironment (presence of lacustrine and river environments) known for this locality during the early late Miocene.     

Functional morphology and anatomy of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene hominoid from Kenya

This paper describes the morphology of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene primate species excavated from Nachola, Kenya in 1999-2002. The cervical vertebrae in Nacholapithecus are larger than those of Papio cynocephalus. They are more robust relative to more caudal vertebral bones. Since Nacholapithecus had large forelimbs, it is assumed that strong cervical vertebrae would have been required to resist muscle reaction forces during locomotion. On the other hand, the vertebral foramen of the lower cervical vertebrae in Nacholapithecus is almost the same size as or smaller than that of P. cynocephalus. Atlas specimens of Nacholapithecus resemble those of extant great apes with regard to the superior articular facet, and they have an anterior tubercle trait intermediate between that of extant apes and other primate species. Nacholapithecus has a relatively short and thick dens on the axis, similar to those of extant great apes and the axis body shape is intermediate between that of extant apes and other primates. Moreover, an intermediate trait between extant great apes and other primate species has been indicated with regard to the angle between the prezygapophyseal articular facets of the axis in Nacholapithecus. Although the atlas of Nacholapithecus is inferred as having a primitive morphology (i.e., possessing a lateral bridge), the shape of the atlas and axis leads to speculation that locomotion or posture in Nacholapithecus involved more orthograde behavior similar to that of extant apes, and, in so far as cervical vertebral morphology is concerned, it is thought that Nacholapithecus was incipiently specialized toward the characteristics of extant hominoids.     

Partial skeleton of Proconsul nyanzae from Mfangano Island,Kenya

A partial skeleton attributed to Proconsul nyanzae (KNM-MW 13142) is described. The fossils were found at a site on Mfangano Island, Kenya, which dates to 17.9 ± .1 million years ago. KNM-MW 13142 consists of six partial vertebrae (T12-S1), a nearly complete hipbone, most of the right femur and left femoral shaft, a fragmentary tibia and fibula, and a nearly complete talus and calcaneus. This skeleton provides the first pelvic fossil known for any East African Miocene hominoid. The new Proconsul specimen is compared to a large sample of extant anthropoids to determine its functional and phylogenetic affinities. In most aspects of its anatomy, KNM-MW 13142 closely resembles nonhominoid anthropoids. This individual had a long, flexible spine, narrow torso, and habitually pronograde posture, features characteristic of most extant monkeys. Evidence of spinal musculature suggests a generalized condition intermediate between that of cercopithecoids and hylobatids. The hindlimb of KNM-MW 13142 exhibits relatively mobile hip and ankle joints, with structural properties of the femur like those of hominoids. This mix of features implies a pattern of posture and locomotion that is unlike that of any extant primate. Many aspects of the Proconsul nyanzae locomotor skeleton may represent the primitive catarrhine condition. © 1993 Wiley-Liss, Inc.     

Morphology of Afropithecus turkanensis from Kenya

Forty-six specimens of a large Miocene hominoid, Afropithecus turkanensis, recently recovered from northern Kenya preserve many morphological details that are described. The specimens include cranial, mandibular, and postcranial parts. They are compared briefly with other Miocene hominoids. It is suggested that Afropithecus may have affinities with Heliopithecus, Kenyapithecus, and the large hominoid from Moroto and Napak, although it is noted that the comparative material is limited in the number of common anatomical parts preserved.     

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.     

Fossil leaves of Populus from the Middle Miocene of Yunnan,SW China

Today southern Yunnan, SW China, has a tropical or subtropical climate and seasonal rainforests. In the past, some temperate elements were also present. In this paper, a new species of Populus is reported from the Middle Miocene deposits in Zhenyuan. Its leaves are ovate or ovate-suborbicular, with serrate margins. They have a shallowly cordate to cuneate base without glands, short acuminate apex, and salicoid teeth with spherical glands. The veins are glabrous but unicellular hair bases occur on the lower epidermis of the lamina. Stomata are confined to the lower epidermis. The presence of Populus in the Middle Miocene of the region indicates an expansion of the genus into low-latitude Asia in the late Cenozoic and a more complicated history of vegetational change in southern Yunnan than has so far been assumed.     

A Rangwapithecus gordoni mandible from the early Miocene site of Songhor,Kenya

A mandible of Rangwapithecus gordoni from the early Miocene site of Songhor, Kenya, provides additional information about this relatively poorly known taxon. The R. gordoni sample is small, being composed of dental and a few gnathic parts. The fossil described here provides examples of previously unknown dental and mandibular anatomy, and confirms former reassignments of isolated anterior teeth based on less certain evidence. The phylogenetic status of Rangwapithecus, its distribution, and paleobiology are briefly reviewed. Rangwapithecus shows a suite of dental and gnathic features that warrants its generic distinction from Proconsul. Derived features shared with Nyanzapithecus and Turkanapithecus indicate that it is an early member of the subfamily Nyanzapithecinae. Its molar morphology suggests a considerable component of folivory in its diet. A review of the hypodigm shows Rangwapithecus to be restricted to Songhor. This distribution parallels that of Limnopithecus evansi, and is mirrored by Limnopithecus legetet and Micropithecus clarki suggesting that Songhor may have differed ecologically from other more or less contemporary sites in the region.     

Comparing levels of homoplasy in the primate skeleton

Hard-tissue morphological characters (bones and teeth) are a primary source of information about the evolutionary history of primates. These tissues are commonly found as isolated elements in the fossil record and studied as three separate partitions: the dentition, the cranium, and the postcranium. The relative phylogenetic utility of characters from each partition is often called into question with respect to varying amounts of homoplasy. In this paper, the consistency index (CI) was used to measure levels of homoplasy in each data partition for a sample of fossil and living primates. Sources of bias in the collection and treatment of data and in the internal structure of the data set are addressed. These biases include number of taxa, number of characters, ordering of characters, amounts of polymorphically scored or missing data, and character-state distribution. The results of this study suggest that the levels of homoplasy are very similar, though the postcranial data may be slightly less homoplastic than either the dental or cranial data.     

A ctenodactylid rodent (Mammalia: Rodentia) from the Middle Miocene of Chios Island (Greece)

Field work carried out in 1991 and 1993 at the Middle Miocene locality of Thymiana (Chios Island, Greece), produced much rodent material including a number of ctenodactylid dental specimens. They represent a single taxon whose upper and lower cheek tooth morphology clearly differs from Prosayimys flynni and all Sayimys species known so far, except for Sayimys intermedius. The ctenodactylid teeth from the Middle Miocene of Chios are identified as pertaining to the latter species, despite minor differences from the Pakistani and Saudi Arabian representatives of S. intermedius.     

Comparative and functional anatomy of phalanges in<Emphasis Type="Italic"> Nacholapithecus kerioi</Emphasis>, a Middle Miocene hominoid from northern Kenya

We describe phalanges of the KNM-BG 35250 Nacholapithecus kerioi skeleton from the Middle Miocene of Kenya. Phalanges of N. kerioi display similarities to those of Proconsul heseloni despite their enhanced robusticity. They do not show highly specialized features as in living suspensory primates. However, N. kerioi manifests several distinctive features that are observed in neither living arboreal quadrupeds nor P. heseloni or P. nyanzae. The most remarkable of them is its phalangeal elongation. N. kerioi phalanges (particularly pedal) are as long as those of Pan despite its much smaller body size. While lengthened digits enable a secure grip of supports and are especially adaptive for grasping large vertical trunks, the skeletal and soft tissues are subjected to greater stress. Probably, strong selective pressures favored powerful hallucal/pollical assisted grips. Although this functional adaptation does not exclude the possible use of the terrestrial environment, arboreal behavioral modes must have been crucial in its positional repertoire. N. kerioi is distinguished from P. heseloni in the greater size of its manual phalanges over its pedal phalanges. These derived features of N. kerioi suggest positional modes supporting more weight on the forelimb, and which occur more frequently on vertical supports. If Proconsul is referred to as an "above-branch arboreal quadruped" with a deliberate and effective climbing capability, N. kerioi may be thought of as an "orthograde climber". While living apes are powerful orthograde climbers, they are also more or less suspensory specialists. Suspensory behavior (plus climbing) and pronograde quadrupedalism (plus climbing) are the two main arboreal behavioral adaptations in living anthropoids. Thus, N. kerioi is an unusual fossil primate in that it cannot be incorporated into this dichotomy. It is plausible that a N. kerioi-like orthograde climber with large forelimbs and cheiridia was a precursor of suspensory living apes, and N. kerioi may demonstrate what an initial hominoid of this grade might have looked like.     

New hominids from East Turkana,Kenya

Thirty-five new fossil hominid specimens are described. They were recovered from the Plio-Pleistocene sediments to the east of Lake Turkana (formerly Lake Rudolf). They include cranial and mandibular parts, teeth, and postcranial bones of upper and lower limbs. Parts of a single skeleton are also described. All of the specimens are described in anatomical detail and selected measurements are given. Some of the specimens are illustrated. It is proposed that they should be attributed to the family Hominidae, with genus and species undetermined until detailed comparative studies have been undertaken.     

Additional specimens of Diceros (Perissodactyla,Rhinocerotidae) from the Upper Miocene Nakali Formation in Nakali,central Kenya

An upper incisor and upper and lower cheek teeth of Rhinocerotidae from the Upper Miocene of Nakali in central Kenya are described. Those specimens are identified as Diceros sp. The present study confirms the presence of Diceros in sub-Saharan East Africa during Vallesian as noted by several studies. The present result and the fossil records of Diceros in Africa and Eurasia suggest that Diceros might have migrated to Eurasia from Africa by Vallesian, although more fossil records and detailed phylogenetic analysis of Diceros are needed to discuss this hypothesis.     

Further hominids from the Plio-Pleistocene of Koobi Fora, Kenya

Fifty-eight new fossil hominids from Plio-Pleistocene sediments east of Lake Turkana, Kenya, are described. They include cranial, mandibular, dental, and postcranial parts. Some are illustrated.