Fort Ternan Research Articles

Palaeoecology of Miocene sites in Western Kenya

The palaeoecology of the Miocene sites of Western Kenya is reviewed. Because fossil floras are usually not preserved the evidence on palaeoecology is mostly obtained from the fossil faunas. These are analysed by five methods: analysis of indicator species, habitat spectra, taxonomic habitat indices, ecological diversity analyses, and socio-ecology. The results indicate that the Songhor fauna was probably derived from forest habitat close to the place of deposition. In the Koru area the faunas from the Legetet Formation and the Chamtwara Member of the Kapurtay Agglomerates were similarly derived from forest habitats, but the Koru Formation fauna indicates more open conditions. The Rusinga and Karungu faunas have not been analysed in detail, and as they are associated with flood plain sedimentary environments it is probable that they are biased and not representative of any on living community, although there is some evidence also for forest conditions. This is supported by evidence from the fossil flora of Rusinga. Two Middle Miocene faunas from Maboko and Fort Ternan both indicate the presence of woodland or woodland savanna habitats, suggesting a major ecological shift from the faunal habitat of the Early Miocene.

The environment ofRamapithecusin Africa

The faunas of three fossil sites in Africa that contain time successive groups of fossil Hominoidea have been analyzed to determine their paleoenvironments. The basis for the analysis is an assessment of the ecological diversity of the fauna, which is expressed in terms of four categories: taxonomic composition, body size, feeding habits and locomotor zonal adaptation. This method has shown that the community structures of the three fossil faunas are significantly different, and comparisons with the community structure of modern habitats suggest that the environment of the early Miocene fauna of Songhor, and the primitive apes associated with it, was probably a type of lowland forest; the habitat ofRamapithecusand the Fort Ternan middle Miocene fauna compares best with modern woodland-bushland habitats; and the habitat ofHomo habilisat Olduvai appears to have been intermediate between grassland and woodland-bushland. If man evolved from one of the early forest living apes, as seems likely on present evidence, an adaptive shift from forest to non-forest habitats must have occurred at some stage in his evolution. The evidence from Fort Ternan shows that in AfricaRamapithecusmade this adaptive shift, and it is also now becoming clear that several genera of Eurasian apes, includingRamapithecus, made a similar environmental change at the same time.

Fossil giraffes from the Miocene of Africa and a revision of the Phylogeny of the Giraffoidea

Specimens of Climacoceras africanus are described from Maboko, Kenya. The new species Climacoceras gentryi is established on the basis of ossicones, mandibles, and upper and lower dentitions from Fort Ternan and Baringo, Kenya. By interpretation of its lower canines Climacoceras is identified as a giraffoid and is placed in the new family Climacoceridae. Canthumeryx sirtensis is identified from Muruarot and Rusinga, Kenya. A dentition and associated partial skeleton of this species are described. The teeth agree closely with specimens of the same species from Gebel Zelten, Libya. Zarafa zelteni from Gebel Zelten is synonymized with Canthumeryx sirtensis . Again on the basis of its lower canines Canthumeryx is identified as a giraffoid and is placed in the new family Canthumerycidae. Specimens of Palaeotragus primaevus are described from Baringo, Kenya. This material includes a cranium with the ossicones, skull roof, occipital and basicranial regions preserved. Palaeotragus primaevus specimens from Fort Ternan are used in this description and some of these are redescribed. The relations of the giraffoids are assessed by methods of phylogenetic systematics. Palaeomeryx, Prolibytherium and Propalaeoryx are excluded from the Giraffoidea as their lower canines are not known. The Palaeotraginae is shown to be an invalid polyphyletic grouping and the genus Palaeotragus is also shown to be polyphyletic. Palaeotragus microdon is probably synonymous with Palaeotragus rouenii and the three species Palaeotragus rouenii (P. microdn), Palaeotragus coelophrys and Palaeotragus quadricornis are retained in the genus Palaeotragus . It is suggested that 'Palaeotragus’ expectans and ‘Palaeotragus’ decipiens are closely related to Samotherium . Palaeotragus primaevus is probably synonymous with Palaeotragus tungurensis and this species is closely related to the giraffines. With slight changes the subfamilies Sivatheriinae and Giraffinae are valid monophyletic groups. Hydaspitherium is synonymized with Bramatherium and the Sivatheriinae includes the genera Giraffokeyx, Birgerbohlinia , Bramatherium and Sivatherium while the Giraffinae includes the genera Honanotherium, Bohlinia and and Giraffa and the species ' Palaeotragus ' tungurensis (P. primaevus). Okapia is identified as the sistergroup of the other giraffids. Triceromeryx is the sister-group of the Giraffidae. Canthumeryx is the sister-group of Triceromeryx plus the Giraffidae while Climacoceras is the sister-group of the other giraffoids.

The origin of the Hominidae: Africa or Asia?

The authors review the evidence for the evolution of the apes and their geographical expansion out of Africa during the Miocene. The ecological background is discussed at each period. From this it is concluded that Dryopithecines in either Africa or Eurasia could have given rise to early Hominidae and that the ecological conditions in both continents were suitable for the appearance of Hominid adaptations. The evidence for early Hominids is at present more impressive in Eurasia than in Africa, though the African find from Fort Ternan, Kenya, is the earliest. On present evidence, neither continent can be precluded as the place of origin of the Hominidae and it appears to be a possibility that Hominidae evolved in both continents with intermittent gene exchange.

The nature of the transition in the dental mechanism from pongids to hominids

Many recent discoveries of Ramapithecus, and of probably ground-living dryopithecines, Dryopithecus (subgenus Sivapithecus), clarify the nature of the transition of the dental mechanism from that of pongids to the hominid stage with reduced canines and flattened cheek teeth with thick enamel. Faunal correlation with potassium/argon dated sites indicates that Sivapithecus and Ramapithecus appeared in the Old World about the same time, approximately 13 million years ago. The thickened molar enamel of these hominoids suggests a terrestrial adaptation in both groups, probably resulting from climatic changes. This adaptation was not necessarily a unique event in the ancestry of the two genera, for species of the two seem to have been different sizes when the change was made. New Ramapithecus finds come from Pyrgos, near Athens, from Çandir in Anatolia, and from Rudabánya, Hungary. At the latter site various specimens preserve all upper and lower teeth in place, while the Çandir and Pyrgos mandibles give important new information about symphyseal structure and orientation, as well as about arcade arrangement. The Rudabánya finds confirm, as do the others, marked facial foreshortening, relatively orthal incisors, anteriorly abbreviated mandible and canine reduction in Ramapithecus. The dental mechanics of Ramapithecus suggested from earlier described finds recovered in the Siwalik deposits of India and Pakistan, as well as at Fort Ternan, Kenya are clarified by the finds from Athens, Anatolia, and Hungary. Like Australopithecus, Ramapithecus mandibles have well-developed double transverse, shallow but transversely thick horizontal rami and anteriorly shifted, vertically oriented, deep ascending branches. These addes resemblances increase the probability that Ramapithecus is in or near the ancestry of Australopithecus and other hominids.

New Fossil Orycteropodidae (Mammalia, Tubulidentata) From East Africa. Orycteropus Minutus Sp. Nov. and Orycteropus Chemeldoi Sp. Nov

Two new species of Orycteropus are described from Miocene deposits of Kenya. O. minutus, sp.nov. the smallest known orycteropodid is recorded from the lower Miocene of Songhor, Rusinga and Mfwangano. O. chemeldoi sp.nov. is known from the mid-Miocene of Ngorora and Fort Ternan. Additional material of O. africanus (MacInnes) is described and parts of three specimens of O. afer crassidens (MacInnes) are described from the lower Pleistocene of East Rudolf. A review of fossil Orycteropodidae is given, and maps and diagrams presented which show all the recorded localities which have yielded Orycteropus. The Eurasian species are considered to form a closely related group which lived there from about 9 million until about 5 million years ago, in which case they may represent a good key group for stratigraphic purposes.

Problems in the interpretation of Ramapithecus: with special reference to anterior tooth reduction.

AbstractThe dental proportions of Ramapithecus specimen FT 1271‐2 (from Fort Ternan, Kenya) have been compared with undoubted fossil pongids from the Miocene of East Africa. Compared to its Miocene pongid contemporaries, Ramapithecus exhibits distinct anterior tooth reduction both in its incisor and canine dimensions. This distinction is most clearly seen in comparisons of Ramapithecus with pongids of similar cheek tooth size, i.e., Dryopithecus africanus and Pan paniscus. The differences in dental proportions between the phylogenetic lines of D. africanus to Pan and Ramapithecus to Homo are discussed in terms of various dietary hypotheses. The similarities in dental proportions of Gorilla and Ramapithecus illustrate their non‐frugivorous dietary preferences, but have little or no value as far as the taxonomic assessment of Ramapithecus is concerned.

Succession of Cainozoic Vertebrate Assemblages from the Northern Kenya Rift Valley

A succession of twelve different fossiliferous units in the Baringo area ranges in time from about 14 m.y. to the Holocene. The early part of the sequence provides for the first time faunal and stratigraphic evidence for the period between 5 m.y. and the Fort Ternan sediments dated about 14 m.y.

Ramapithecus wickeri Mandible from Fort Ternan, Kenya

THE portion of a hominoid mandible (KNM*–FT 45) found in 1962 at Fort Ternan in Kenya has been referred provisionally by Leakey1 to Dryopithecus (sensu strictu). Simons2 has agreed essentially with this. Other primates found at Fort Ternan include the maxilla fragments of Ramapithecus wickeri (Leakey) 1962 (ref. 3), several individuals of Pliopithecus, and several isolated teeth attributed to Proconsul and Oreopithecus. By size alone the mandibular specimen (KNM–FT 45) can be distinguished from all these except R. wickeri. A detailed examination of occlusal relationships between the upper teeth in the maxilla of R. wickeri (KNM–FT 46)† and the two premolars in KNM–FT 45 suggests that the two specimens belong to the same individual; and even if this is not so they definitely belong to the same species. Simons4 and Pilbeam5 have argued that Leakey's taxon Kenyapithecus wickeri is a junior synonym of Ramapithecus punjabicus (Pilgrim) 1910 (ref. 6), but the evidence indicates at least a species difference, and this is the position I take here.

Fort Ternan Hominid.

SIMONS'S paper1 on the Late Miocene Hominid from Fort Ternan, Kenya, states, inter alia, at the end of the second paragraph, “It is the intention of this paper to indicate the continuing validity of this synonymy”, that is, that the genus Kenyapithecus Leakey is no more than a synonym of Ramapithecus Lewis.

Late Miocene hominid from Fort Ternan, Kenya.

Faunal and temporal relationships of the early hominid sites at Fort Ternan and in North India further support the morphological evidence that “Kenyapithecus” from Fort Ternan ought to continue in the synonymy of Ramapithecus.

Upper Miocene Primates from Kenya

Specimens of Kenyapithecus wickeri representing the family Hominidae have already been described from the Upper Miocene site at Fort Ternan, Kenya. The site has also yielded fossil remains of a member of the Hylobatidae resembling Limnopithecus; teeth of a primate closely resembling Oreopithecus; teeth of a species of Proconsul; part of a jaw which suggests the genus Dryopithecus and some teeth of the family Cercopithecoidea.

Bone smashing by late Miocene Hominidae.

Bones found in the Upper Miocene Fossil Beds of Fort Ternan, Kenya, show evidence of having been broken up by some kind of blunt instrument.

Historical Zoogeography of Antelopes

STUDIES of the Bovidae of the later Miocene of Fort Ternan, Kenya1, and of the lower Pliocene of Samos are producing interesting zoogeographical findings. In this communication, names above the species level are from Simpson2.

An early Miocene member of Hominidae.

Hitherto, the earliest fossils recognized as belonging to the Hominidae, from Fort Ternan and the Siwaliks, have been dated to the Mio-Pliocene. Discoveries in early Miocene deposits at Songhor and Rusinga, Kenya, have made it possible to carry the date of the separation of the true Hominidae from the Pongidae back to the Lower—or possibly Early Middle—Miocene

Note on the Fort Ternan beds of Kenya

Precise dating of the Fort Ternan beds has not been possible as yet, but a late Miocene age is suggested.

Fossil Rodents from Fort Ternan, Kenya

THE rodent fauna from Fort Ternan which Dr. L. S. B. Leakey has sent to me for examination consists for the moment of only a few specimens comprising the following: Seventeen mandibles, a fragment of a skull in which the teeth are unfortunately very worn, and 26 incisors. The mandibles represent creatures of small size, but on the other hand six of the incisors suggest the presence of species of the size of the large Phiomyidae of Rusinga and Songhor. Nine mandibles belong to the Cricetodontini, one represents a small Phiomydae similar to one of the forms from Songhor, one represents a Sciuridae, while three of the mandibles belong to a new genus, Leakeymys ternani nov. gen., nov. sp., to which genus and species I also provisionally attribute the skull fragment. (The collection also includes a mandible of a murine, but in this specimen the bone is not completely fossilized as it is in all the other material, and it is, therefore, I believe, an intrusive specimen which can be ignored at the present time.)

The 'White Highlands' of Kenya

The term 'the White Highlands* was derived from the official policy that certain agricultural lands in Kenya should be reserved for settlers of European origin. The termination of this policy after a period of nearly sixty years makes this an appropriate time to examine which were the areas included in the White Highlands and why. To early explorers and administrators, the cool climate and absence of population over large areas of the highlands of what is now called Kenya suggested the possibility of European settlement. In 1901 much of this land lay within the boundaries of Uganda and Sir Harry Johnston was able to report (Cmd. 671)'. . . here we have a territory (now that the Uganda Railway is built) admirably suited for a white man's country, and I can say this, with no thought of injustice to any native race, for the country in question is either utterly uninhabited for miles and miles or at most its inhabitants are wandering hunters who have no settled home, or whose fixed habitation is the lands outside the healthy The first Land Regulations (of the East Africa Protectorate) were published as early as 1897 but little alienation took place until after the construction of the Uganda Railway, which reached Nairobi in 1899 and Lake Victoria in 1901, when settlement was greatly encouraged and the regula? tions were replaced by the Crown Lands Ordinance, 1902. Under this Ordinance, grants of Crown Lands could be made freehold or by leases of up to ninety-nine years. The Commissioner of Lands was not empowered to sell or lease any land in the actual occupation of the natives and further, if any grants were made which were subsequently found to contain African settlements, these settlements were deemed to be excluded from the lease so long as they were occupied. It was never contemplated that grants of land could be made to Africans, who had already selected the areas they chose to occupy. However, another immigrant community was interested in farming land in the highlands, the Asians, particularly as represented by the labourers brought from India to assist in the building of the railway. Under pressure from the Government of India, the contracts of these workers could be terminated in East Africa instead of in India, if the worker so desired. It is clear that by 1902 grants of Crown Land for agricultural purposes in the highlands were being restric? ted to Europeans or Americans, so that, although applications were invited from In? dians to lease agricultural land, the area lying between Kiu and Fort Ternan was specifically excluded. This policy was approved by Lord Elgin, the Secretary of State for the Colonies, in July 1906. Thus the Commissioner of Lands would only grant Crown Land for agricultural purposes within the highlands area, which was understood to extend from approximately Kiu to Fort Ternan railway stations, to Europeans. Furthermore, the lease could not be transferred without the approval of the Commissioner, who would presumably apply the same policy to the transfer as he had to the original grant. When the Crown Land Ordinance, 1915, replaced that of 1902 a further extension of this policy was written into section 36 as follows: There shall by virtue of this Ordinance be implied in every lease granted under this part to a European a covenant that he shall not without the consent of the Governor in Council appoint or allow a non-European to be a manager or otherwise to occupy or be in control of the land leased*.