An ape partial postcranial skeleton (KNM-NP 64631) from the Middle Miocene of Napudet, northern Kenya.

An ape partial postcranial skeleton (KNM-NP 64631) was recovered during the 2015-2021 field seasons at Napudet, a Middle Miocene (∼13 Ma) locality in northern Kenya. Bony elements representing the shoulder, elbow, hip, and ankle joints, thoracic and lumbar vertebral column, and hands and feet, offer valuable new information about the body plan and positional behaviors of Middle Miocene apes. Body mass estimates from femoral head dimensions suggest that the KNM-NP 64631 individual was smaller-bodied (c. 13-17 kg) than some Miocene taxa from eastern Africa, including Ekembo nyanzae, and probably Equatorius africanus or Kenyapithecus wickeri, and was more comparable to smaller-bodied male Nacholapithecus kerioi individuals. Similar to many Miocene apes, the KNM-NP 64631 individual had hip and hallucal tarsometatarsal joints reflecting habitual hindlimb loading in a variety of postures, a distal tibia with a large medial malleolus, an inflated humeral capitulum, probably a long lumbar spine, and a long pollical proximal phalanx relative to femoral head dimensions. The KNM-NP 64631 individual departs from most Early Miocene apes in its possession of a more steeply beveled radial head and deeper humeral zona conoidea, reflecting enhanced supinating-pronating abilities at the humeroradial joint. The KNM-NP 64631 individual also differs from Early Miocene Ekembo heseloni in having a larger elbow joint (inferred from radial head size) relative to the mediolateral width of the lumbar vertebral bodies and a more asymmetrical talar trochlea, and in these ways recalls inferred joint proportions for, and talocrural morphology of, N. kerioi. Compared to most Early Miocene apes, the KNM-NP 64631 individual likely relied on more forelimb-dominated arboreal behaviors, perhaps including vertical climbing (e.g., extended elbow, hoisting). Moreover, the Napudet ape partial postcranial skeleton suggests that an arboreally adapted body plan characterized by relatively large (here, based on joint size) forelimbs, but lacking orthograde suspensory adaptations, may not have been 'unusual' among Middle Miocene apes.

Kanapoi revisited: Paleoecological and biogeographical inferences from the fossil fish.

Fish fossils were recovered from three different depositional contexts at the Pliocene Kanapoi site to: 1) test the assumption that habitat and ecology of modern fish taxa can predict habitat and ecology of fossil taxa; 2) reconstruct the lake and river environments in the Kanapoi Formation, with reference to fish fossils from the nearby Lothagam site deposits; and 3) investigate biogeographical inferences from the fossils. We compare the Kanapoi fish taxa and their depositional environments with the taxa and environments in modern Lake Turkana, and with another Plio-Pleistocene fauna from the eastern Turkana Basin. Taphonomic caveats are discussed. Our results support the use of ecological preferences of modern fish to predict past preferences. Our analysis of the Kanapoi fossils also indicates that the Pliocene Lonyumun Lake had a diverse fauna, with an unusual mix of taxa compared to the modern lake. The presence of possibly endemic species in the Pliocene lake may additionally represent a period of isolation during this epoch. Few fish fossils were recovered in the deposits of the ancestral Kerio River, a primary affluent of Lonyumun Lake then as now, but those present indicate a different ecology than that interpreted for the modern lake. Previously unknown fish taxa which enter the lake during the Pliocene suggest the existence of a connection between the Nile River and the Turkana Basin, which may have been viable for other vertebrates, including hominins.

Proboscidea from Kanapoi, Kenya.

The early Pliocene site of Kanapoi (Turkana Basin, Kenya) has a large, diverse vertebrate sample that contains the earliest representatives of the hominin genus Australopithecus. Included in this sample is an impressive assemblage of fossil proboscideans, comprised of deinotheres (Deinotherium bozasi), anancine gomphotheres (Anancus ultimus), and at least three species of elephant (Loxodonta adaurora, a primitive morph of Loxodonta exoptata, and Elephas ekorensis). A single specimen from high in the sequence could plausibly belong to a primitive stage of Elephas recki. A review of dental carbon isotope analyses indicates a range of dietary habits for these taxa, from dedicated browsing (deinotheres) to mixed feeding/grazing (elephants and gomphotheres), which in early Pliocene elephants corresponds to molars with greater crown height and more plates than in late Miocene confamilials, bringing their morphology more in phase with feeding behavior than was the case in their earlier relatives. Variation in feeding preferences among Kanapoi proboscideans corresponds to evidence for habitat heterogeny, including inferred substantial presence of grasses; the occurrence of multiple megaherbivores may have contributed to the fragmentation of ecosystems, positively affecting early hominin success and aiding diversification of other ungulate groups.

Revisiting the pedogenic carbonate isotopes and paleoenvironmental interpretation of Kanapoi.

Reconstructed habitats of Australopithecus anamensis at Kanapoi by Wynn (2000) yielded evidence for both wooded and grassy environments. Wynn's study was based on stable isotopic (δ13CPC, δ18OPC) analyses of a small sample of pedogenic nodules (n = 14) collected from paleosols spanning Kanapoi's stratigraphic interval. Whether this small sample size adequately characterized Kanapoi's vegetation or was the result of time averaging remains unclear. To address this uncertainty, we sampled Kanapoi paleosols at 39 locations (78 analyses) from laterally extensive units. Our data demonstrate that Kanapoi offered A. anamensis diverse habitats distributed in temporally discrete stratigraphic horizons. Habitat heterogeneity appears to have been a real aspect of Kanapoi paleoenvironments and not an artifact of Wynn's (2000) small sample size or time averaging. We suggest habitat heterogeneity was influenced by the location of Kanapoi at the confluence of fluvial, deltaic, and lacustrine depositional environments. We also compared Kanapoi's δ13CPC and δ18OPC values to those of other Pliocene hominin localities in eastern Africa dated to 4.5-3.7 Ma. Kanapoi's δ18OPC values are significantly higher than most sites, potentially reflecting regional variability in water source δ18O values and/or more arid climatic conditions. Kanapoi's δ13CPC values indicate significantly more woody cover than at all other sites except those in the Turkana Basin. Kanapoi provided A. anamensis with a wide range of C3-C4 resources as the C4 biome spread across eastern Africa. [Wynn, J.G., 2000. Paleosols, stable carbon isotopes and paleoenvironmental interpretation of Kanapoi, Northern Kenya. J. Hum. Evol. 39, 411-432.].

Isotopic equifinality and rethinking the diet of Australopithecus anamensis.

OBJECTIVES: Australopithecus anamensis has comparable δ13 Cenamel values to Ardipithecus ramidus, and both have been characterized as C3 feeders in open woodland habitats similar to "savanna" chimps. Unlike Ar. ramidus and "savanna" chimps, A. anamensis shows a derived dentognathic morphology for tough foods and a dental microwear pattern similar to the C3 -C4 -mixed-feeding A. afarensis. Here I test the hypothesis that changing the variables (ε*enamel-diet , δ13 CC3, δ13 CC4 values) used to calculate the percentage of dietary C4 foods (%C4 diet) by 1-2‰ does not make a substantial difference for hominin diet reconstructions [van der Merwe, Masao, & Bamford, 2008, South African Journal of Science 104:153-155]. MATERIALS AND METHODS: I estimate vegetation structures for A. anamensis with pedogenic carbonate and faunal enamel δ13 C values from the Pliocene Omo-Turkana Basin (4.2-3.9 Ma). I recalculate A. anamensis' %C4 diet based on new body size-dependent estimates of the ε*enamel-diet value and alternative δ13 CC3 and δ13 CC4 values. RESULTS: The Pliocene Omo-Turkana Basin shows evidence for dietary resources with a wide range of δ13 C values including canopy-driven 13 C depleted ground C3 foods. Alternative equation variables changed by 1-2‰ yield higher C4 estimates for A. anamensis (15-31%) than previously thought (0-10%). The choice of δ13 CC3 and δ13 CC4 values for estimating %C4 is not a perfunctory task and potentially explains the δ13 C isotopic equifinality of A. anamensis and "savanna" chimps. DISCUSSION: My integrative diet model reconciles the carbon isotopic data with the dentognathic and microwear evidence of A. anamensis' diet and suggests that "savanna" chimps are not proper dietary analogs of A. anamensis. A foraging strategy across heterogeneous habitats of the Pliocene Omo-Turkana Basin incorporating an array of 13 C-depleted and 13 C-enriched C3 foods and a portion of C4 resources may have served as one of the selective pressures for A. anamensis, the earliest habitual biped [van der Merwe et al., 2008, South African Journal of Science 104:153-155].

Diversity analysis of Plio-Pleistocene large mammal communities in the Omo-Turkana Basin, eastern Africa.

Knowing how the diversity of large mammal communities changes across space and time provides an important ecological framework for studying hominin evolution. However, diversity studies that apply methods currently used by neoecologists are rare in paleoanthropology and are also challenging due to diversity's unusual statistical properties. Here, we apply up-to-date analytical methods for understanding how species- and genus-level large mammalian diversity in the Omo-Turkana Basin changed through time and across space at multiple spatiotemporal scales (within each formation:102-3 km2 and 104-5 years; and within the basin as a whole: 103 km2 and 105 years). We found that, on average, Koobi Fora's large mammal community was more diverse than Nachukui's, which in turn was more diverse than Shungura's. Diversity was stable through time within each of these formations (alpha diversity), as was diversity in the basin as a whole (gamma diversity). Compositional dissimilarity between these three formations (beta diversity) was relatively low through time, with a 0.6 average proportion of shared species, suggesting dispersal acted to homogenize the region. Though alpha and gamma diversity were fairly stable through time, we do observe several notable peaks: during the KBS Member in Koobi Fora (30% increase), the Lokalalei Member in Nachukui (120% increase), and at 1.7 Ma in the entire basin (100% increase). We conclude by (1) demonstrating that habitat heterogeneity was an important factor influencing alpha diversity within each of the three formations, and (2) hypothesizing that diversity stability may have been driven by equilibrial dynamics in which overall diversity was constrained by resource availability, implying biotic interactions were an important factor in structuring the communities that included hominins as members. Our findings demonstrate the need to quantify how large mammal diversity changes across time and space in order to further our understanding of hominin ecology and evolution.

Influence of Plio-Pleistocene basin hydrology on the Turkana hominin enamel carbonate δ(18)O values.

Stable oxygen isotopes of hominin enamel carbonate (δ(18)OEC) provide a window into aspects of past drinking behavior and diet, body size, breastfeeding and weaning, mobility, and paleoclimate. It is tempting to compare all hominins across time and space in order to gauge species-level adaptations to changing environments and niche separation between those living sympatrically. Basinal, sub-basinal, and micro-environmental differences, however, may exert an influence on variation in enamel carbonate isotopic values that must be reconciled before hominin species across Africa can be meaningfully compared. Plio-Pleistocene Turkana hominin δ(18)OEC values show a considerable spread, potentially revealing many intrinsic and extrinsic contributing factors operating on different scales. In this study, I examine Turkana hominin δ(18)OEC values relative to identity (taxon, tooth type and number, body size of taxon), dietary (δ(13)C value, Turkana coeval and modern mammalian δ(18)OEC values), and contextual (time, depositional environment) information of each specimen and collection locality and discuss various potential influences. Turkana hominin δ(18)OEC values may primarily reflect differences in imbibed water sources (lake vs. river) as a function of evolving basin hydrology.

Chronostratigraphy of KNM-ER 3733 and other Area 104 hominins from Koobi Fora.

A magnetostratigraphy for ∼ 60 m of Koobi Fora Formation sediment in Area 104 was derived from 46 oriented samples that produced well-resolved characteristic magnetizations from progressive thermal demagnetization. Approximately 59 m below the Morte Tuff, previously dated to ∼ 1.51 Ma (millions of years ago), the Olduvai-Matuyama boundary (∼ 1.78 Ma) was found to be at the level of marker bed A2--inconsistent with the Area 102 type section and thus contrary to fossil dating schemes that utilize temporal equivalence between A2 [104] and A2 [102]. The magnetostratigraphic data, coupled with the Morte Tuff, provide a means to interpolate new ages for marker beds A2 [104] and the White Tuff, as well as multiple Area 104 hominin fossils. Noteworthy is the new date of ∼ 1.63 Ma for KNM-ER 3733, which now implicates KNM-ER 2598 as the sole early African Homo erectus fossil demonstrably older than Dmanisi and Java Homo specimens. Re-dating KNM-ER 3733 creates a ∼ 300-kyr gap at 1.9 to 1.6 Ma in the African fossil record of H. erectus, which might be partially spanned by hand axes recently dated at ∼ 1.76 Ma, if the Acheulian is indeed proprietary to this species.

Improved age control on early Homo fossils from the upper Burgi Member at Koobi Fora, Kenya.

To address questions regarding the evolutionary origin, radiation and dispersal of the genus Homo, it is crucial to be able to place the occurrence of hominin fossils in a high-resolution chronological framework. The period around 2 Ma (millions of years ago) in eastern Africa is of particular interest as it is at this time that a more substantial fossil record of the genus Homo is first found. Here we combine magnetostratigraphy and strontium (Sr) isotope stratigraphy to improve age control on hominin-bearing upper Burgi (UBU) deposits in Areas 105 and 131 on the Karari Ridge in the eastern Turkana Basin (Kenya). We identify the base of the Olduvai subchron (bC2n) plus a short isolated interval of consistently normal polarity that we interpret to be the Pre-Olduvai event. Combined with precession-forced (~20 kyr [thousands of years]) wet-dry climate cycles resolved by Sr isotope ratios, the magnetostratigraphic data allow us to construct an age model for the UBU deposits. We provide detailed age constraints for 15 hominin fossils from Area 131, showing that key specimens such as cranium KNM-ER 1470, partial face KNM-ER 62000 and mandibles KNM-ER 1482, KNM-ER 1801, and KNM-ER 1802 can be constrained between 1.945 ± 0.004 and 2.058 ± 0.034 Ma, and thus older than previously estimated. The new ages are consistent with a temporal overlap of two species of early Homo that can be distinguished by their facial morphology. Further, our results show that in this time interval, hominins occurred throughout the wet-dry climate cycles, supporting the hypothesis that the lacustrine Turkana Basin was a refugium during regionally dry periods. By establishing the observed first appearance datum of a marine-derived stingray in UBU deposits at 2.058 ± 0.034 Ma, we show that at this time the Turkana Basin was hydrographically connected to the Indian Ocean, facilitating dispersal of fauna between these areas. From a biogeographical perspective, we propose that the Indian Ocean coastal strip should be considered as a possible source area for one or more of the multiple Homo species in the Turkana Basin from over 2 Ma onwards.

Interactions between climate change and infrastructure projects in changing water resources: An ethnobiological perspective from the Daasanach, Kenya.

The fast and widespread environmental changes that have intensified in the last decades are bringing disproportionate impacts to Indigenous Peoples and Local Communities. Changes that affect water resources are particularly relevant for subsistence-based peoples, many of whom already suffer from constraints regarding reliable access to safe water. Particularly in areas where water is scarce, climate change is expected to amplify existing stresses in water availability, which are also exacerbated by multiple socioeconomic drivers. In this paper, we look into the local perceptions of environmental change expressed by the Daasanach people of northern Kenya, where the impacts of climate change overlap with those brought by large infrastructure projects recently established in the Omo River. We show that the Daasanach have rich and detailed understanding of changes in their environment, especially in relation to water resources. Daasanach understand observations of change in different elements of the social-ecological system as an outcome of complex interactions between climatic and non-climatic drivers of change. Our findings highlight the perceived synergistic effects of climate change and infrastructure projects in water resources, driving multiple and cascading impacts on biophysical elements and local livelihoods. Our results also demonstrate the potential of Local Ecological Knowledge in enhancing the understanding of complex social-ecological issues, such as the impacts of environmental change in local communities. To minimize and mitigate the social-ecological impacts of development projects, it is essential to consider potential synergies between climatic and socioeconomic factors and to ensure inclusive governance rooted in local understandings of environmental change.

The bioarchaeology of mid-Holocene pastoralist cemeteries west of Lake Turkana, Kenya.

Early herders in eastern Africa built elaborate megalithic cemeteries ~ 5000 BP overlooking what is now Lake Turkana in northwestern Kenya. At least six 'pillar sites' were constructed during a time of rapid change: cattle, sheep, and goats were introduced to the basin as the lake was shrinking at the end of the African Humid Period. Cultural changes at this time include new lithic and ceramic technologies and the earliest monumentality in eastern Africa. Isolated human remains previously excavated from pillar sites east of Lake Turkana seemed to indicate that pillar site platforms were ossuaries for secondary burials. Recent bioarchaeological excavations at four pillar sites west of the lake have now yielded ≥49 individuals, most from primary and some from secondary interments, challenging earlier interpretations. Here we describe the mortuary cavities, and burial contexts, and included items such as adornments from Lothagam North, Lothagam West, Manemanya, and Kalokol pillar sites. In doing so, we reassess previous hypotheses regarding pillar site construction, use, and inter-site variability. We also present the first osteological analyses of skeletons buried at these sites. Although the human remains are fragmentary, they are nevertheless informative about the sex, age, and body size of the deceased and give evidence for health and disease processes. Periosteal moulds of long bone midshafts (n = 34 elements) suggest patterns of terrestrial mobility. Pillar site deposits provide important new insights into early herder lifeways in eastern Africa and the impact of the transition to pastoralism on past human populations.