Miocene cercopithecoidea from the Tugen Hills, Kenya

Miocene to Pleistocene fossiliferous sediments in the Tugen Hills span the time period from at least 15.5 Ma to 0.25 Ma, including time periods unknown or little known elsewhere in Africa. Consequently, the Tugen Hills deposits hold the potential to inform us about crucial phylogenetic events in African faunal evolution and about long-term environmental change. Among the specimens collected from this region are a number of discoveries already important to the understanding of primate evolution. Here, we describe additional cercopithecoid material from the Miocene deposits in the Tugen Hills sequence, including those from securely dated sites in the Muruyur Beds (16-13.4 Ma), the Mpesida Beds (7-6.2 Ma) and the Lukeino Formation (∼6.2-5.7 Ma). We also evaluate previously described material from the Ngorora Formation (13-8.8 Ma). Identified taxa include Victoriapithecidae gen. et sp. indet., cf. Parapapio lothagamensis, and at least two colobines. Specimens attributed to cf. Pp. lothagamensis would extend the species’ geographic range beyond its type locality. In addition, we describe specimens sharing derived characters with modern African colobines (Tribe: Colobina), a finding that is congruent with previous molecular estimates of colobine divergence dates. These colobine specimens represent some of the earliest known members of the modern African colobine radiation and, in contrast to previous hypotheses, suggest that early African colobines were mainly arboreal and that semi-terrestrial Late Miocene and Plio-Pleistocene colobine taxa were secondarily derived in their locomotor adaptations.     

Geology and geochronology of the middle Miocene Kipsaramon site complex, Muruyur Beds, Tugen Hills, Kenya.

The Muruyur Beds are a substantial sedimentary deposit within a middle Miocene sequence of mafic volcanic flows associated with early stages of rifting in the central Kenyan Rift Valley. They are best represented in the Muruyur region, near Bartabwa, north of Kipsaramon, where dates range from 16.0 to 13.4 Ma. At Kipsaramon, located about 10 km south of Muruyur along the crest of the Tugen Hills, the upper Muruyur Beds are absent and the lower part can be divided into three members. Important fossil sites within Member 1 are dated between 15.8 and 15.6 Ma, and within Member 3 between 15.6 and 15.4 Ma. BPRP#89, in Member 1, is a bonebed at least 2500 m(2)in areal extent and up to 30 cm thick, which constitutes one of the richest concentrations of in situ fossil vertebrate bones in eastern Africa. BPRP#91, at approximately the same level at BPRP#89, is the source of a hominoid talus and other mammal and bird fossils. In Member 3, BPRP#122 has produced specimens of at least five individuals of the hominoid Equatorius, including a partial skeleton. The Muyuyur Beds were deposited near the western margin of a lake that was formed during the early stages of faulting and volcanism in the African Rift system. The bonebed in Member 1 appears to represent the influx of fluvially transported vertebrate and plant remains into a shallow portion of the lake. Elements of the fauna as well as stable isotopes that indicate both forest and more open environments occurred in proximity to the lake during the time of deposition of Member 1.     

New cercopithecoids and a hominoid from 12.5 Ma in the Tugen Hills succession, Kenya.

The early evolutionary history of the cercopithecoids is poorly understood, primarily due to a lack of fossil material from between 15 and about 9 Ma. Cercopithecoid primate specimens from a fossil site in the Ngorora Formation of the Tugen Hills, Kenya, belong to the genus Victoriapithecus, possibly a new species. These fossils are associated with a hominoid specimen that resembles Proconsul, and another tooth of a catarrhine, also probably hominoid. The locality is BPRP#38, in the Kabasero type section of the Ngorora Formation, and well dated at 12.5 Ma. If the hominoid specimen is confirmed as Proconsul, it would be one of the most recent recorded. The relatively diverse mammal fauna from the site in some ways resembles that of Fort Ternan. The site is underlain, and not far removed in time, from one of the best fossil macro-floras in Africa, which indicates lowland rainforest conditions in this portion of the Rift Valley at 12.6 Ma.     

The age of Orrorin tugenensis,an early hominid from the Tugen Hills,Kenya

Throughout the history of the East African Rift valley, tectonic depressions and volcanic dams formed within the graben in which Middle Miocene to Recent volcanic rocks and lacustrine to fluvial sediments accumulated. During the Late Miocene, voluminous trachytes erupted in the vicinity of Kabarnet and almost filled the valley to its brim. Continued tectonic activity formed new basins floored by the Kabarnet Trachyte, one of which was located in the region immediately east of the present day Tugen Hills. The Lukeino sediments that accumulated in this basin crop out over an area of 44 km × 13 km. In 2000, Orrorin tugenensis, which is important for understanding the earliest stages of human evolution, was found in the Lukeino Formation 〚12〛, 〚13〛. It is concluded that the Lukeino formation accumulated between 6.0 and 5.7 Ma.     

Stratigraphy, age and environments of the late Miocene Mpesida Beds, Tugen Hills, Kenya.

Interpretations of faunal assemblages from the late Miocene Mpesida Beds in the Tugen Hills of the Central Kenyan Rift Valley have figured prominently in discussions of faunal turnover and establishment of the modern East African communities. These faunal changes have important implications for the divergence of the human lineage from the African apes ca. 8-5 Ma. While fossil material recovered from the Mpesida Beds has traditionally been analyzed collectively, accumulating evidence indicates that Mpesida facies span the 7-6 Ma interval and are scattered more than 25 km along the eastern flanks of the Tugen Hills. Stratigraphic distinctions between Mpesida facies and younger sediments in the sequence, such as the Lukeino Formation, are not yet fully resolved, further complicating temporal assessments and stratigraphic context of Mpesida facies. These issues are discussed with specific reference to exposures of Mpesida facies at Rurmoch, where large fossil tree fragments were swept up in an ancient ash flow. Preserved anatomical features of the fossil wood as well as estimated tree heights suggest a wet, lowland rainforest in this portion of the rift valley. Stable isotopic analyses of fossil enamel and paleosol components indicate the presence of more open habitats locally. Overlying air-fall tuffs and epiclastic debris, possibly associated with the ash flow, have yielded an assemblage of vertebrate fossils including two teeth belonging to one of the earliest colombines of typical body size known from Africa, after the rather small Microcolobus. Single-crystal, laser-fusion,(40)Ar/(39)Ar dates from a capping trachyte flow as well as tuffs just below the lava contact indicate an age of greater than 6.37 Ma for the fossil material.     

Partial skeleton of Theropithecus brumpti (Primates, Cercopithecidae) from the Chemeron Formation of the Tugen Hills, Kenya

Here we describe a complete skull and partial skeleton of a large cercopithecoid monkey (KNM-TH 46700) discovered in the Chemeron Formation of the Tugen Hills at BPRP Site #152 (2.63 Ma). Associated with the skeleton was a mandible of an infant cercopithecoid (KNM-TH 48364), also described here. KNM-TH 46700 represents an aged adult female of Theropithecus brumpti, a successful Pliocene papionin taxon better known from the Omo Shungura Formation in Ethiopia and sites east and west of Lake Turkana, Kenya. While the morphology of male T. brumpti is well-documented, including a partial skeleton with both cranial and postcranial material, the female T. brumpti morphotype is not well-known. This skeleton represents some of the first associated evidence of cranial and postcranial female T. brumpti remains. In addition to the complete skull, postcranial material includes elements of the axial skeleton and lower limb. While aspects of the skeleton conform to those of specimens previously assigned to T. brumpti, other features on the femur and tibia appear to differ from those previously described for this species. It is unclear whether these differences represent general variation within the T. brumpti population, variation between the sexes in T. brumpti, or the incorrect assignment of previous isolated hindlimb specimens. In total, the observable morphological features of the hindlimb suggest that KNM-TH 46700 was a terrestrial quadruped similar to modern savannah baboons (Papio). From the available evidence, it is difficult to assess whether or not KNM-TH 46700 frequently engaged in the specialized squatting and shuffling behavior observed in extant geladas (Theropithecus gelada).     

Pliocene hominid partial mandible from Tabarin, Baringo, Kenya

Sediments in the Tugen Hills, west of Lake Baringo, Kenya, form one of the best fossiliferous successions known in Africa spanning the period from 14 my to less than 4 my. Hominoid fossils have previously been recovered from a number of localities in the region. We describe here a new hominid mandible (KNM-TH 13150) from the site of Tabarin, in the Chemeron Formation. Isotopic determinations on a tuff below the fossiliferous horizon gives dates of 4.96 my and 5.25 my. The associated fauna is consistent with these results and independently suggests a minimum age for the specimen of 4.15 my. Although fragmentary, the preserved morphology of the Tabarin mandible is consistent with the diagnosis of the Pliocene hominid Australopithecus afarensis. It can be distinguished from all other currently recognized hominoid taxa.     

Creodonts and carnivores from the Middle Miocene Muruyur Formation at Kipsaraman and Cheparawa,Baringo District,Kenya

The Middle Miocene Muruyur Formation (ca 14.5 Ma), Tugen Hills, Kenya, has yielded a huge creodont and a variety of carnivores ranging in size from mongoose-sized viverrids and herpestids to lion-sized amphicyonids. The fauna partly fills what used to be a major gap in our knowledge of Neogene African carnivores, spanning the period between the better known Early Miocene assemblages of western Kenya and eastern Uganda, and the Late Miocene and Plio–Pleistocene faunas of East Africa. Present in the deposits are Megistotherium, two species of Hecubides, one species of Agnotherium, Herpestes, Vishnuictis, and one or two undetermined felids.     

New species of Listriodon (Suidae,Mammalia) from Bartule,Member A,Ngorora Formation (ca 13 Ma), Tugen Hills,Kenya

Hitherto the only listriodont known from the Ngorora Formation, Kenya, was Lopholistriodon kidogosana, a highly derived form represented by a skull with dentition and some isolated upper and lower teeth. The only available mandibular material of the species was a fragment of symphysis. A recently discovered listriodont mandible with most of the dentition from Bartule, near Bartabwa, belongs to a new species of a different genus, Listriodon.     

On the limnology of Lake Baringo (Kenya): II. Pelagic primary production and algal composition of Lake Baringo,Kenya

Sicilian reservoirs, a vital resource for this densely populated island, are undergoing increased eutrophication processes, due to the lack of urban waste treatment plants and to intense agriculture. Massive blooms of toxic cyanobacteria represent one of the most obvious consequences, and can be dangerous for population health. The possibility of a fast, even if only short term, recovery of water quality through application of lime, was tested through analysis of ion dynamics in two reservoirs, to develop an `emergency plan', to counteract cyanobacterial blooms and to decrease the risk for human health. The results indicate that these water bodies are suitable for lime treatments, even though the high content of sulphate and their managed hydrological cycles, have to be considered.     

Preliminary observations on the BK 8518 mandible from Baringo, Kenya

A newly discovered adult hominid mandible (BK 8518) from Baringo, Kenya, is described and assessed. The corpus, many of the tooth crowns, and most of the left ascending ramus are preserved. The teeth are heavily and asymmetrically worn. Compared with BK 67 (the 1966 mandible) the body of BK 8518 is more robust; the internal symphyseal buttressing is more pronounced; the M3s have seven cusps and exceed the M2s in size. There are no compelling reasons, however, to attribute the two mandibles to different taxa and, in view of the lack of any comprehensive taxonomic diagnosis for Homo erectus, "erectus-like," and habiline mandibular remains, the new specimen is also best regarded as Homo sp. indet. (aff. erectus).     

40Ar/(39)Ar dating of Chemeron Formation strata encompassing the site of hominid KNM-BC 1, Tugen Hills, Kenya.

A fossil hominid temporal bone (KNM-BC 1) from surface exposures at Baringo Paleontological Research Project site BPRP#2 in the Chemeron Formation outcropping in a tributary drainage of the Kapthurin River west of Lake Baringo, Kenya has been attributed to Homo sp. indet. K-feldspar phenocrysts from lapilli tuffs bracketing the inferred fossiliferous horizon yield single-crystal(40)Ar/(39)Ar ages of 2.456+/-0.006 and 2.393+/-0.013 Ma. These age determinations are supported by stratigraphically consistent ages on higher tuff horizons and from nearby sections. In addition, new(40)Ar/(39)Ar ages on tuffaceous units near the base and top of the formation along the Kapthurin River yield 3.19+/-0.03 and 1.60+/-0.05 Ma respectively. The base of the formation along the Kapthurin River is thus approximately 0.5 Ma younger than the uppermost Chemeron Formation strata exposed at Tabarin, 23 km to the north-northwest. The upper half of the formation along the Kapthurin River was deposited at an average rate of approximately 11 cm/ka, compared to 21-23 cm/ka at Tabarin.     

Neogene paleobiogeography and East African paleoenvironments: contributions from the Tugen Hills rodents and lagomorphs.

A minimum of 28 genera of rodents and one genus of lagomorph were recovered from the Tugen Hills, Baringo District, Kenya, from localities dating from over 15.5 to about 4.4 Ma. The middle Miocene (sites dated between 15.8 and 15.3 Ma) rodent fauna recovered primarily from the Kipsaramon site complex, Muruyur Formation, includes a mixture of characteristically early Miocene taxa, and more derived forms. Composition of the African rodent fauna changes dramatically with the introduction of myocricetodontines, democricetodontines, and dendromurines, immigrants primarily from southern Asia. In the Tugen Hills, these taxa are first found in the Kabasero localities, Ngorora Formation, at sites dating from 12.5-12.33 Ma. A second major change in the African rodent fauna reflects the introduction of murines, immigrants from southern Asia. In the Tugen Hills murines are first encountered at Kapcheberek, Lukeino Formation, dated to 5.9-5.7 Ma. One rodent genus from the Lukeino Formation (Arvicanthis), and two from the Tabarin locality, Chemeron Formation (Heliosciurus, Paraxerus; 4.5-4.4 Ma), represent the earliest records of these extant African genera. A cricetomyine from the Ngorora Formation (12.5 Ma) is likely the earliest report of this exclusively African group. One of the earliest African records of porcupines (Hystricide) is from the Lukeino Formation. Lagomorphs are poorly represented, but include one of the earliest African occurrences of the family Leporidae from the Mpesida Beds (bracketed by dates of 7-6.2 Ma), and possibly a new genus of leporid from the Kapcheberek locality. Analysis of the Tugen Hills small mammals in association with other African records suggests several episodes of dispersal between Africa and Eurasia during the middle and late Miocene. Rodents from Kipsaramon are indicative of forests in conjunction with more open habitats. Those from the Kapcheberek locality are suggestive of a savanna habitat. The rodents from the Tabarin locality suggest a woodland environment.     

New wrist bones of Proconsul africanus and P. nyanzae from Rusinga Island, Kenya

The anatomy of the wrist of two species of the early Miocene hominoid Proconsul is described based on new material collected on Rusinga Island, Kenya. These fossils generally confirm previous findings that the wrist of Proconsul is monkey-like in much of its morphology. However, the structure of the ulnar side of the wrist, particularly the ulnocarpal joint, is significantly different from that of extant monkeys and suggests some functional affinities with extant hominoids. Thus the wrist of Proconsul is neither monkey-like nor ape-like in its total morphology. Instead, it shows a unique combination of features which once again point to the oversimplicity of forcing fossil forms into categories based only on extant taxa.     

Fossil leaves from Lukeino,a 6-million-year-old Formation in the Baringo Basin,Kenya

Thirty-eight dicotyledonous leaf morphotypes and some monocot leaf fragments are described from the Kapcharar, Kabogongoi, Koibo Chepkweny and Inoswa Kamelon localities in the upper third of the Lukeino Formation, Tugen Hills, Baringo Basin (Kenya). The leaves are preserved in fine-grained paper shales and range in length from 9 to 160 mm. Some of the leaves show affinities with extant species in the Burseraceae, Ebenaceae, Euphorbiaceae, Fabaceae, Loganiaceae, Malvaceae and Sapotaceae families that are common in the local flora today. There are compound and simple leaves, mostly entire-margined and with only 10% non-entire margins. A CLAMP analysis of the Kapcharar leaves indicates that the local climate was seasonal, summer wet with 1723 mm mean annual rainfall and with a mean annual temperature of 19.8 °C. The vegetation was a deciduous forest or woodland with open areas nearby. A monocot leaf and two fragments of fern pinnae are also described; they imply locally wetter environments. The flora described here is directly associated with the early hominid Orrorin tugenensis.     

Remarks apropos of Pliocene hominoid humerus from Chemeron (basin of Lake Baringo, Kenya

Detailed study of a proximal fragment of a Pliocene left humerus and comparison with extant and fossil primates suggests that the specimen, although definitely hominoid, cannot be clearly associated with any known Plio-Pleistocene hominid or with any modern hominoids.     

Trophic state and nutrient limitation in Lake Baringo,Kenya

The trophic state of Lake Baringo and factors that could be limiting the development of algal biomass in it were investigated during one wet/dry hydrological cycle in 2014–2015. Water samples were analysed for dissolved inorganic nutrients, including , and , total phosphorus and Chlorophyll a. Light attenuation was estimated using Secchi depth. The trophic state was determined using Carlson trophic state indices (CTSI). Deviations in CTSI, nutrient ratios and ambient nutrient concentrations were used to identify factors limiting phytoplankton growth. The mean values measured for Secchi depth, nitrate, total phosphorus and Chlorophyll a showed significant seasonal variation (p < 0.05). Based on the Carlson trophic state index, the results show that Lake Baringo is eutrophic. However, the lake is also experiencing phosphorus limitation and poor light penetration, because of high turbidity, which is more pronounced during the wet season.