Systematics of early and middle Miocene Old World monkeys.

New information about the early cercopithecoids Prohylobates tandyi (Wadi Moghra, Egypt) and Prohylobates sp. indet. (Buluk and Nabwal, Kenya) is presented. Comparisons are made among all major collections of Early and Middle Miocene catarrhine monkeys, and a systematic revision of the early Old World monkeys is provided. Previous work involving the systematics of early Old World monkeys (Victoriapithecidae; Cercopithecoidea) has been hampered by a number of factors, including the poor preservation of Prohylobates material from North Africa and lack of comparable anatomical parts across collections. However, it is now shown that basal cercopithecoid species from both northern and eastern Africa can be distinguished from one another on the basis of degree of lower molar bilophodonty, relative lower molar size, occlusal details, symphyseal construction, and mandibular shape. Results of particular interest include: 1) the first identification of features that unambiguously define Prohylobates relative to Victoriapithecus; 2) confirmation that P. tandyi is incompletely bilophodont; and 3) recognition of additional victoriapithecid species.

Implications of new early Homo fossils from Ileret, east of Lake Turkana, Kenya.

Sites in eastern Africa have shed light on the emergence and early evolution of the genus Homo. The best known early hominin species, H. habilis and H. erectus, have often been interpreted as time-successive segments of a single anagenetic evolutionary lineage. The case for this was strengthened by the discovery of small early Pleistocene hominin crania from Dmanisi in Georgia that apparently provide evidence of morphological continuity between the two taxa. Here we describe two new cranial fossils from the Koobi Fora Formation, east of Lake Turkana in Kenya, that have bearing on the relationship between species of early Homo. A partial maxilla assigned to H. habilis reliably demonstrates that this species survived until later than previously recognized, making an anagenetic relationship with H. erectus unlikely. The discovery of a particularly small calvaria of H. erectus indicates that this taxon overlapped in size with H. habilis, and may have shown marked sexual dimorphism. The new fossils confirm the distinctiveness of H. habilis and H. erectus, independently of overall cranial size, and suggest that these two early taxa were living broadly sympatrically in the same lake basin for almost half a million years.

Growth processes in teeth distinguish modern humans from Homo erectus and earlier hominins.

A modern human-like sequence of dental development, as a proxy for the pace of life history, is regarded as one of the diagnostic hallmarks of our own genus Homo. Brain size, age at first reproduction, lifespan and other life-history traits correlate tightly with dental development. Here we report differences in enamel growth that show the earliest fossils attributed to Homo do not resemble modern humans in their development. We used daily incremental markings in enamel to calculate rates of enamel formation in 13 fossil hominins and identified differences in this key determinant of tooth formation time. Neither australopiths nor fossils currently attributed to early Homo shared the slow trajectory of enamel growth typical of modern humans; rather, both resembled modern and fossil African apes. We then reconstructed tooth formation times in australopiths, in the approximately 1.5-Myr-old Homo erectus skeleton from Nariokotome, Kenya, and in another Homo erectus specimen, Sangiran S7-37 from Java. These times were shorter than those in modern humans. It therefore seems likely that truly modern dental development emerged relatively late in human evolution.

Morphology of Australopithecus anamensis from Kanapoi and Allia Bay, Kenya.

The hominid species Australopithecus anamensis was originally described in 1995, with new specimens and more secure dates given in 1998. This paper lists all fossils attributed to A. anamensis, and provides anatomical descriptions of those not yet described in detail with photographs of all but undiagnostic fragments. We also provide comparative analysis of these specimens. The A. anamensis holotype mandible was found at Kanapoi, as were most of the paratypes. The Allia Bay sample is less well represented, and does not preserve many anatomical elements diagnostic of this species. Still, the Allia Bay sample most closely resembles that from Kanapoi, and we suggest that for the time being it be retained as A. anamensis. A. anamensis most closely resembles A. afarensis, but can be distinguished from it in many features. Most of these features are inferred to be primitive for the genus. Based on the limited postcranial evidence available, A. anamensis appears to have been habitually bipedal, although it retained some primitive features of its upper limbs. A. anamensis differs from A. afarensis in having narrower, more parallel jaws with a very slightly more ape-like canine/premolar complex than is found in A. afarensis, although not as ape-like as in Ardipithecus ramidus. It had slightly larger lower lateral incisors, a unique upper canine morphology, and a different structure of the lateral nasal aperture than A. afarensis. A. anamensis had at least as great a range of body size, and perhaps slightly greater canine dimorphism, although this is difficult to determine. At present, there appears to be no autapomorphies precluding A. anamensis from ancestry of A. afarensis.

New hominin genus from eastern Africa shows diverse middle Pliocene lineages.

Most interpretations of early hominin phylogeny recognize a single early to middle Pliocene ancestral lineage, best represented by Australopithecus afarensis, which gave rise to a radiation of taxa in the late Pliocene. Here we report on new fossils discovered west of Lake Turkana, Kenya, which differ markedly from those of contemporary A. afarensis, indicating that hominin taxonomic diversity extended back, well into the middle Pliocene. A 3.5 Myr-old cranium, showing a unique combination of derived facial and primitive neurocranial features, is assigned to a new genus of hominin. These findings point to an early diet-driven adaptive radiation, provide new insight on the association of hominin craniodental features, and have implications for our understanding of Plio-Pleistocene hominin phylogeny.

South Turkwel: a new pliocene hominid site in Kenya.

New fossils discovered south of the Turkwel River in northern Kenya include an associated metacarpal, capitate, hamate, lunate, pedal phalanx, mandibular fragment, and teeth. These fossils probably date to around 3.5 m.y.a. Faunal information suggests that the environment at South Turkwel was predominantly bushland. The mandibular and dental remains are fragmentary, but the postcranial fossils are informative. Comparisons with Australopithecus, modern human, chimpanzee and gorilla hand bones suggest that the Turkwel hominid was most like Australopithecus afarensis and A. africanus. Carpometacarpal articulations are intermediate between those of modern humans and African apes, suggesting enhanced gripping capabilities compared with extant apes. The hamulus was strikingly large, similar in proportion only to Neandertals and some gorillas, suggesting the presence of powerful forearms and hands. There are no indicators of adaptations to knuckle-walking or suspensory locomotion in the hand, and the pedal phalanx suggests that this hominid was habitually bipedal.

Comparative dental development and microstructure of Proconsul teeth from Rusinga Island, Kenya.

Eighteen histological sections were prepared from eleven teeth attributed to Proconsul heseloni and two molar teeth attributed to Proconsul nyanzae. Measurements of spacings and counts of daily incremental markings in both enamel and dentine were possible in the majority of these tooth sections. Measurements of the spacings and angles to the enamel dentine junction (EDJ) of regular striae of Retzius and of equivalent markings in dentine were also made. In addition to these measurements, counts of perikymata were made on replicas of all other Proconsul teeth housed in the National Museum of Kenya, Nairobi, that preserved good perikymata on any aspect of their tooth surface. The sequence of crown formation in Proconsul and the crown formation times of the enamel and dentine were estimated from these data. In addition, the rates of root extension were estimated using the formula derived for this purpose by Shellis (Archs. oral Biol. 29, 697-705, 1984) and estimates of the total period of root formation subsequently made for premolar and molar teeth based on measurements of root length. A composite chart of dental development for P. heseloni is presented which suggests M3 root completion was between six and seven years of age. In general Proconsul molar teeth have high stria angles to the EDJ, a high ratio of enamel formed with respect of dentine formed at the same time, median values of rates of enamel formation close to the EDJ in excess of 4 microns per day and the occasional presence of "S-shaped" striae in the lateral enamel. There is no evidence to suggest that Proconsul from Rusinga Island, Kenya, had relatively thin enamel on molar or premolar teeth. When all of these data are considered in a comparative context, Proconsul emerges overall as hominoid-like in its enamel and dentine microstructure and as most similar to Pongo but with some features shared with Pan and Homo. Similar data for other Miocene primates will have considerable bearing on how these data are interpreted. These new data on dental microanatomy and on dental development in Proconsul make a further contribution to our understanding of the total morphological picture of this early Miocene primate.

New specimens and confirmation of an early age for Australopithecus anamensis.

The discovery of Australopithecus anamensis fossils from strata lying between tephra dated at 4.17 and 4.12 million years ago, and from slightly higher strata not well constrained in age by overlying dated units, provoked the claim that more than one species might be represented: it was suggested that the stratigraphically higher fossils, which include the important tibia, humerus and a large, presumed male, mandible (KNM-KP 29287), might belong to a later, more derived hominid. We have recovered new fossils from Kanapoi and Allia Bay, Kenya, during field work in 1995-1997 that confirm the primitive status of Australopithecus anamensis, the earliest species of Australopithecus. Isotope dating confirms A. anamensis' intermediate age as being between those of Ardipithecus ramidus and Australopithecus afarensis. New specimens of maxilla, mandible and capitate show that this species is demonstrably more primitive than A. afarensis. A lower first deciduous molar (dm 1) is intermediate in morphology between that reported for Ardipithecus ramidus and A. afarensis. Single-crystal 40Ar-39Ar age determinations on the Kanapoi Tuff show that, except for a large mandible, all of the hominid fossils from Kanapoi are from sediments deposited between 4.17+/-0.03 and 4.07+/-0.02 million years ago.

Early hominid fossils from Africa.

A new species of Australopithecus, the ancestor of Homo, pushes back the origins of bipedalism to some four million years ago.

Dietary and environmental reconstruction with stable isotope analyses of herbivore tooth enamel from the Miocene locality of Fort Ternan, Kenya.

Tooth enamel of nine Middle Miocene mammalian herbivores from Fort Ternan, Kenya, was analyzed for delta 13C and delta 18O. The delta 18O values of the tooth enamel compared with pedogenic and diagenetic carbonate confirm the use of stable isotope analysis of fossil tooth enamel as a paleoenvironmental indicator. Furthermore, the delta 18O of tooth enamel indicates differences in water sources between some of the mammals. The delta 13C values of tooth enamel ranged from -8.6(-)-13.0/1000 which is compatible with a pure C3 diet, though the possibility of a small C4 fraction in the diet of a few of the specimens sampled is not precluded. The carbon isotopic data do not support environmental reconstructions of a Serengeti-typed wooded grassland with a significant proportion of C4 grasses. This study does not preclude the presence of C3 grasses at Fort Ternan; it is possible that C3 grasses could have had a wider geographic range if atmospheric CO2 levels were higher than the present values.

New four-million-year-old hominid species from Kanapoi and Allia Bay, Kenya.

Nine hominid dental, cranial and postcranial specimens from Kanapoi, Kenya, and 12 specimens from Allia Bay, Kenya, are described here as a new species of Australopithecus dating from between about 3.9 million and 4.2 million years ago. The mosaic of primitive and derived features shows this species to be a possible ancestor to Australopithecus afarensis and suggests that Ardipithecus ramidus is a sister species to this and all later hominids. A tibia establishes that hominids were bipedal at least half a million years before the previous earliest evidence showed.

Four-million-year-old hominids from East Lake Turkana, Kenya.

A piece of mandible and several isolated teeth are reported from fluviatile sediments older than 4 million years at East Lake Turkana. They most closely resemble hominids from Laetoli, Tanzania and Hadar, Ethiopia which have been assigned to Australopithecus afarensis.

Similarities in Aegyptopithecus and Afropithecus facial morphology.

Recently discovered cranial fossils from the Oligocene deposits of the Fayum depression in Egypt provide many details of the facial morphology of Aegyptopithecus zeuxis. Similar features are found in the Miocene hominoid Afropithecus turkanensis. Their presence is the first good evidence of a strong phenetic link between the Oligocene and Miocene hominoids of Africa. A comparison of trait lists emphasizes the similarities of the two fossil species, and leads us to conclude that the two fossil genera share many primitive facial features. In addition, we studied facial morphology using finite-element scaling analysis and found that the two genera show similarities in morphological integration, or the way in which biological landmarks relate to one another in three dimensions to define the form of the organism. Size differences between the two genera are much greater than the relatively minor shape differences. Analysis of variability in landmark location among the four Aegyptopithecus specimens indicates that variability within the sample is not different from that found within two samples of modern macaques. We propose that the shape differences found among the four Aegyptopithecus specimens simply reflect individual variation in facial characteristics, and that the similarities in facial morphology between Aegyptopithecus and Afropithecus probably represent a complex of primitive facial features retained over millions of years.

Morphology of Turkanapithecus kalakolensis from Kenya.

The type specimen of Turkanapithecus kalakolensis recently recovered from northern Kenya preserves a partial cranium and mandible. This specimen, together with postcranial elements that are probably associated with it, and two additional specimens are described. The collection is compared with other Miocene hominoids.

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.

Pliocene and pleistocene hominid-bearing sites from west of lake turkana, kenya.

Pliocene and Pleistocene fossil localities near the western shoreline of Lake Turkana, ranging in age between 1 million and 3.5 million years in age, have produced important new hominid specimens including most of a Homo erectus skeleton and a relatively complete early robust australopithecine cranium. The lacustrine, fluviatile, and terrestrial strata are designated the Nachukui Formation, which is subdivided into eight members. The distribution of sedimentary facies within the Nachukui Formation suggests that, as today, the Labur and Murua Rith ranges formed the western margin of the basin and were drained by eastward-flowing rivers that fed into the forerunner of the present lake or a major river system. There is also stratigraphic evidence for tectonic movement during the deposition of these sediments. Twenty-three of the tuffs observed in the succession occur also in the Koobi Fora Formation east of the lake and in the Shungura Formation of the lower Omo Valley and permit precise correlation among these three localities. Fortyseven fossiliferous sites from West Turkana have yielded more than 1000 specimens of 93 mammalian species. The mammalian fossils represent nine sequential assemblages that augment information about faunal and environmental change from elsewhere in the basin.