Hominin dental remains from the Pliocene localities at Lomekwi, Kenya (1982-2009).

Increasing evidence for both taxonomic diversity and early stone manufacture during the Pliocene highlights the importance of the hominin fossil record from this epoch in eastern Africa. Here, we describe dental remains from Lomekwi (West Turkana, Kenya), which date from between 3.2 and 3.5 Ma. The sample was collected between 1982 and 2009 and includes five gnathic specimens and a total of 67 teeth (mostly isolated permanent postcanine teeth). Standard linear dimensions indicate that, although the Lomekwi teeth are relatively small, there is broad overlap in size with contemporary Australopithecus afarensis and Australopithecus deyiremeda specimens at most tooth positions. However, some dental characters differentiate this sample from these species, including a relatively large P4 and M3 compared with the M1, a high incidence of well-developed protostylids, and specific accessory molar cuspules. Owing to a lack of well-preserved tooth crowns (and the complete absence of mandibular teeth) in the holotype and paratype of Kenyanthropus platyops, and limited comparable gnathic morphology in the new specimens, it cannot be determined whether these Lomekwi specimens should be attributed to this species. Attribution of these specimens is further complicated by a lack of certainty about position along the tooth row of many of the molar specimens. More comprehensive shape analyses of the external and internal morphology of these specimens, and additional fossil finds, would facilitate the taxonomic attribution of specimens in this taxonomically diverse period of human evolution.

N-glycolyl groups of nonhuman chondroitin sulfates survive in ancient fossils.

Biosynthesis of the common mammalian sialic acid N-glycolylneuraminic acid (Neu5Gc) was lost during human evolution due to inactivation of the CMAH gene, possibly expediting divergence of the Homo lineage, due to a partial fertility barrier. Neu5Gc catabolism generates N-glycolylhexosamines, which are potential precursors for glycoconjugate biosynthesis. We carried out metabolic labeling experiments and studies of mice with human-like Neu5Gc deficiency to show that Neu5Gc degradation is the metabolic source of UDP-GlcNGc and UDP-GalNGc and the latter allows an unexpectedly selective incorporation of N-glycolyl groups into chondroitin sulfate (CS) over other potential glycoconjugate products. Partially N-glycolylated-CS was chemically synthesized as a standard for mass spectrometry to confirm its natural occurrence. Much lower amounts of GalNGc in human CS can apparently be derived from Neu5Gc-containing foods, a finding confirmed by feeding Neu5Gc-rich chow to human-like Neu5Gc-deficient mice. Unlike the case with Neu5Gc, N-glycolyl-CS was also stable enough to be detectable in animal fossils as old as 4 My. This work opens the door for investigating the biological and immunological significance of this glycosaminoglycan modification and for an "ancient glycans" approach to dating of Neu5Gc loss during the evolution of Homo.

Complete permanent mandibular dentition of early Homo from the upper Burgi Member of the Koobi Fora Formation, Ileret, Kenya.

The KNM-ER 64060 dentition derives from a horizon that most likely dates to between 2.02 and 2.03 Ma. A proximate series of postcranial bones (designated KNM-ER 64061) derives from the same siltstone unit and may be associated with the dentition, but their separation on the surface of the site leaves some room for doubt. KNM-ER 64060 is one of fewer than ten hominin specimens from the Early Pleistocene of East Africa that comprises a full or nearly complete mandibular dentition. Its taxonomic attribution is potentially significant, especially if the postcranial elements are related. At least three, and probably four hominin species, including Paranthropus boisei and Homo erectus (= H. ergaster), are known at about this time in East Africa. Other penecontemporaneous fossils have been referred to a single, highly variable species, H. habilis, or two taxa, namely H. habilis and H. rudolfensis. Although the weight of evidence supports the attribution of these specimens to two species, there is notable lack of agreement over the assignation of individual fossils. We take a conservative approach and group all such specimens under the designation "early Homo sp." for comparative purposes. KNM-ER 64060 is clearly attributable to Homo rather than Paranthropus. The preponderance of the evidence suggests that the affinities of KNM-ER 64060 are with fossils assigned to the early Homo sp. category rather than with H. erectus. This is indicated by the overall sizes of the KNM-ER 64060 canine, premolar and molar crowns, the size relationships of the P3 to P4, the relative narrowness of its premolar crowns, the cusp proportions of the M1 and especially those of the M2 and M3, and seemingly the possession of a two-rooted P4. Some of these comparisons suggest further that among the fossils comprising the early Homo sp. sample, the KNM-ER 64060 dentition exhibits greater overall similarity to specimens such as OH 7 and OH 16 that represent Homo habilis sensu stricto.

New fossils from Koobi Fora in northern Kenya confirm taxonomic diversity in early Homo.

Since its discovery in 1972 (ref. 1), the cranium KNM-ER 1470 has been at the centre of the debate over the number of species of early Homo present in the early Pleistocene epoch of eastern Africa. KNM-ER 1470 stands out among other specimens attributed to early Homo because of its larger size, and its flat and subnasally orthognathic face with anteriorly placed maxillary zygomatic roots. This singular morphology and the incomplete preservation of the fossil have led to different views as to whether KNM-ER 1470 can be accommodated within a single species of early Homo that is highly variable because of sexual, geographical and temporal factors, or whether it provides evidence of species diversity marked by differences in cranial size and facial or masticatory adaptation. Here we report on three newly discovered fossils, aged between 1.78 and 1.95 million years (Myr) old, that clarify the anatomy and taxonomic status of KNM-ER 1470. KNM-ER 62000, a well-preserved face of a late juvenile hominin, closely resembles KNM-ER 1470 but is notably smaller. It preserves previously unknown morphology, including moderately sized, mesiodistally long postcanine teeth. The nearly complete mandible KNM-ER 60000 and mandibular fragment KNM-ER 62003 have a dental arcade that is short anteroposteriorly and flat across the front, with small incisors; these features are consistent with the arcade morphology of KNM-ER 1470 and KNM-ER 62000. The new fossils confirm the presence of two contemporary species of early Homo, in addition to Homo erectus, in the early Pleistocene of eastern Africa.

Dietary changes of large herbivores in the Turkana Basin, Kenya from 4 to 1 Ma.

A large stable isotope dataset from East and Central Africa from ca. 30 regional collection sites that range from forest to grassland shows that most extant East and Central African large herbivore taxa have diets dominated by C4 grazing or C3 browsing. Comparison with the fossil record shows that faunal assemblages from ca. 4.1-2.35 Ma in the Turkana Basin had a greater diversity of C3-C4 mixed feeding taxa than is presently found in modern East and Central African environments. In contrast, the period from 2.35 to 1.0 Ma had more C4-grazing taxa, especially nonruminant C4-grazing taxa, than are found in modern environments in East and Central Africa. Many nonbovid C4 grazers became extinct in Africa, notably the suid Notochoerus, the hipparion equid Eurygnathohippus, the giraffid Sivatherium, and the elephantid Elephas. Other important nonruminant C4-grazing taxa switched to browsing, including suids in the lineage Kolpochoerus-Hylochoerus and the elephant Loxodonta. Many modern herbivore taxa in Africa have diets that differ significantly from their fossil relatives. Elephants and tragelaphin bovids are two groups often used for paleoecological insight, yet their fossil diets were very different from their modern closest relatives; therefore, their taxonomic presence in a fossil assemblage does not indicate they had a similar ecological function in the past as they do at present. Overall, we find ecological assemblages of C3-browsing, C3-C4-mixed feeding, and C4-grazing taxa in the Turkana Basin fossil record that are different from any modern ecosystem in East or Central Africa.

Diet of Theropithecus from 4 to 1 Ma in Kenya.

Theropithecus was a common large-bodied primate that co-occurred with hominins in many Plio-Pleistocene deposits in East and South Africa. Stable isotope analyses of tooth enamel from T. brumpti (4.0-2.5 Ma) and T. oswaldi (2.0-1.0 Ma) in Kenya show that the earliest Theropithecus at 4 Ma had a diet dominated by C4 resources. Progressively, this genus increased the proportion of C4-derived resources in its diet and by 1.0 Ma, had a diet that was nearly 100% C4-derived. It is likely that this diet was comprised of grasses or sedges; stable isotopes cannot, by themselves, give an indication of the relative importance of leaves, seeds, or underground storage organs to the diet of this primate. Theropithecus throughout the 4- to 1-Ma time range has a diet that is more C4-based than contemporaneous hominins of the genera Australopithecus, Kenyanthropus, and Homo; however, Theropithecus and Paranthropus have similar proportions of C4-based resources in their respective diets.

Stable isotope-based diet reconstructions of Turkana Basin hominins.

Hominin fossil evidence in the Turkana Basin in Kenya from ca. 4.1 to 1.4 Ma samples two archaic early hominin genera and records some of the early evolutionary history of Paranthropus and Homo. Stable carbon isotopes in fossil tooth enamel are used to estimate the fraction of diet derived from C3 or C4 resources in these hominin taxa. The earliest hominin species in the Turkana Basin, Australopithecus anamensis, derived nearly all of its diet from C3 resources. Subsequently, by ca. 3.3 Ma, the later Kenyanthropus platyops had a very wide dietary range--from virtually a purely C3 resource-based diet to one dominated by C4 resources. By ca. 2 Ma, hominins in the Turkana Basin had split into two distinct groups: specimens attributable to the genus Homo provide evidence for a diet with a ca. 65/35 ratio of C3- to C4-based resources, whereas P. boisei had a higher fraction of C4-based diet (ca. 25/75 ratio). Homo sp. increased the fraction of C4-based resources in the diet through ca. 1.5 Ma, whereas P. boisei maintained its high dependency on C4-derived resources.

Associated ilium and femur from Koobi Fora, Kenya, and postcranial diversity in early Homo.

During the evolution of hominins, it is generally accepted that there was a shift in postcranial morphology between Australopithecus and the genus Homo. Given the scarcity of associated remains of early Homo, however, relatively little is known about early Homo postcranial morphology. There are hints of postcranial diversity among species, but our knowledge of the nature and extent of potential differences is limited. Here we present a new associated partial ilium and femur from Koobi Fora, Kenya, dating to 1.9 Ma (millions of years ago) that is clearly attributable to the genus Homo but documents a pattern of morphology not seen in eastern African early Homo erectus. The ilium and proximal femur share distinctive anatomy found only in Homo. However, the geometry of the femoral midshaft and contour of the pelvic inlet do not resemble that of any specimens attributed to H. erectus from eastern Africa. This new fossil confirms the presence of at least two postcranial morphotypes within early Homo, and documents diversity in postcranial morphology among early Homo species that may reflect underlying body form and/or adaptive differences.

Diet of Paranthropus boisei in the early Pleistocene of East Africa.

The East African hominin Paranthropus boisei was characterized by a suite of craniodental features that have been widely interpreted as adaptations to a diet that consisted of hard objects that required powerful peak masticatory loads. These morphological adaptations represent the culmination of an evolutionary trend that began in earlier taxa such as Australopithecus afarensis, and presumably facilitated utilization of open habitats in the Plio-Pleistocene. Here, we use stable isotopes to show that P. boisei had a diet that was dominated by C(4) biomass such as grasses or sedges. Its diet included more C(4) biomass than any other hominin studied to date, including its congener Paranthropus robustus from South Africa. These results, coupled with recent evidence from dental microwear, may indicate that the remarkable craniodental morphology of this taxon represents an adaptation for processing large quantities of low-quality vegetation rather than hard objects.

Late Miocene to Pliocene carbon isotope record of differential diet change among East African herbivores.

Stable isotope and molecular data suggest that C(4) grasses first appeared globally in the Oligocene. In East Africa, stable isotope data from pedogenic carbonate and fossil tooth enamel suggest a first appearance between 15-10 Ma and subsequent expansion during the Plio-Pleistocene. The fossil enamel record has the potential to provide detailed information about the rates of dietary adaptation to this new resource among different herbivore lineages. We present carbon isotope data from 452 fossil teeth that record differential rates of diet change from C(3) to mixed C(3)/C(4) or C(4) diets among East African herbivore families at seven different time periods during the Late Miocene to the Pliocene (9.9-3.2 Ma). Significant amounts of C(4) grasses were present in equid diets beginning at 9.9 Ma and in rhinocerotid diets by 9.6 Ma, although there is no isotopic evidence for expansive C(4) grasslands in this part of the Late Miocene. Bovids and hippopotamids followed suit with individuals that had C(4)-dominated (>65%) diets by 7.4 Ma. Suids adopted C(4)-dominated diets between 6.5 and 4.2 Ma. Gomphotheriids and elephantids had mostly C(3)-dominated diets through 9.3 Ma, but became dedicated C(4) grazers by 6.5 Ma. Deinotheriids and giraffids maintained a predominantly C(3) diet throughout the record. The sequence of differential diet change among herbivore lineages provides ecological insight into a key period of hominid evolution and valuable information for future studies that focus on morphological changes associated with diet change.

An exploratory study on the combined effects of external and internal morphology on load dissipation in primate capitates: its potential for an understanding of the positional and locomotor repertoire of early hominins.

This pilot study explored whether the redirection of stress through trabeculae within morphologically constrained capitates provides information about habitual/positional behaviours unavailable from the study of external morphology alone. To assess this possibility, an experimental finite element approach was taken, whereby no attempt was made to reconstruct the actual magnitudes and loading conditions experienced by the capitates in vivo. Rather, this work addressed fundamental biological questions relating to bone plasticity, i.e. internal versus external bone morphology. The capitates of 7 species with different and - in the case of fossils - inferred locomotor behaviours were selected. Virtual models of capitates were created, scaled to the same size and subjected to the same theoretical load. In the first set of analyses, models were assigned the material properties of bone throughout, whereas in the second set, models were assigned 11 different material properties representing the trabecular architecture derived from high-resolution CT. Species with arboreal behaviours consistently redirected loads towards the ulnar aspect of the capitate when trabeculae were introduced, while terrestrial species, and the bipedal Homo, redirected stress towards the radial side. From these preliminary analyses, it is tentatively concluded that Australopithecus anamensis habitually engaged in arboreal behaviours, whereas Australopithecus afarensis did not.

Middle Pliocene hominin diversity: Australopithecus deyiremeda and Kenyanthropus platyops.

Geometric morphometric shape analyses are used to compare the maxillae of the Kenyanthropus platyops holotype KNM-WT 40000, the Australopithecus deyiremeda holotype BRT-VP-3/1 and other australopiths. The main aim is to explore the relationship between these two specimens and contemporary Australopithecus afarensis Five landmarks placed on lateral views of the maxillae quantify key aspects of the morphology. Generalized Procrustes analyses and principal component analyses of the resulting shape coordinates were performed. The magnitudes of differences in shape and their significances were assessed using Procrustes and Mahalanobis' distances, respectively. Both KNM-WT 40000 and BRT-VP-3/1 show statistically significant differences in maxillary shape from A. afarensis, but do so in dissimilar ways. Moreover, the former differs more from A. afarensis than the latter. KNM-WT 40000 has a more anteriorly positioned zygomatic process with a transversely flat, and more orthognathic subnasal clivus. BRT-VP-3/1 has a more inferiorly positioned zygomatic process, a slightly retracted dental arcade, but without shortening of the anterior maxilla. These findings are consistent with previous conclusions that the two fossils should be attributed to separate species, rather than to A. afarensis, and with the presence of three contemporary hominin species in the Middle Pliocene of eastern Africa.This article is part of the themed issue 'Major transitions in human evolution'.

Browsing and grazing in elephants: the isotope record of modern and fossil proboscideans.

The diet of extant elephants (Loxodonta in Africa, Elephas in Asia) is dominated by C3 browse although some elephants have a significant C4 grass component in their diet. This is particularly noteworthy because high-crowned elephantid cheek teeth represent adaptation to an abrasive grazing diet and because isotopic analysis demonstrates that C4 vegetation was the dominant diet for Elephas in Asia from 5 to 1 Ma and for both Loxodonta and Elephas in Africa between 5-1 Ma. Other proboscideans in Africa and southern Asia, except deinotheres, also had a C4-dominated diet from about 7 Ma (when the C4 biomass radiated in tropical and subtropical regions) until their subsequent extinction.

Enamel and dentine development and the life history profile of Victoriapithecus macinnesi from Maboko Island, Kenya.

Two permanent lower second molar teeth (KNM-MB 19841 and KNM-MB 27844) attributed to Victoriapithecus macinnesi were prepared for histological analysis. A further five male and three female canine teeth were replicated with a silicone impression material and perikymata counts subsequently made on epoxy casts over the whole crown surface. Daily enamel cross striations averaged 6 microm apart in both cuspal and lateral enamel of the molars and the total crown formation times were approximately 1.24 years in the molars and, using the same periodicity determined from the molars, 1.84 and 1.36 years respectively in the male and female canines. Rates of dentine formation matched those known for extant macaques and were used to calculate root extension rates, which averaged 11.5 microm per day over the whole 8 mm root length of KNM-MB 19841. The period between M2 initiation and gingival emergence was estimated to be approximately 1.95 years in Victoriapithecus which is greater than estimates for Cebus albifrons and Chlorocebus aethiops, (which are similar in body mass to Victoriapithecus), but less than estimates made here for several macaque species. A speculative picture of dental development in Victoriapithecus emerges that is slower than that known for modern vervet monkeys and may have been more similar to that in some smaller modern macaque species.

Hominin diversity in the Middle Pliocene of eastern Africa: the maxilla of KNM-WT 40000.

The 3.5-Myr-old hominin cranium KNM-WT 40000 from Lomekwi, west of Lake Turkana, has been assigned to a new hominin genus and species, Kenyanthropus platyops, on the basis of a unique combination of derived facial and primitive neurocranial features. Central to the diagnosis of K. platyops is the morphology of the maxilla, characterized by a flat and relatively orthognathic subnasal region, anteriorly placed zygomatic processes and small molars. To study this morphology in more detail, we compare the maxillae of African Plio-Pleistocene hominin fossils and samples of modern humans, chimpanzees and gorillas, using conventional and geometric morphometric methods. Computed tomography scans and detailed preparation of the KNM-WT 40000 maxilla enable comprehensive assessment of post-mortem changes, so that landmark data characterizing the morphology can be corrected for distortion. Based on a substantially larger comparative sample than previously available, the results of statistical analyses show that KNM-WT 40000 is indeed significantly different from and falls outside the known range of variation of species of Australopithecus and Paranthropus, contemporary Australopithecus afarensis in particular. These results support the attribution of KNM-WT 40000 to a separate species and the notion that hominin taxonomic diversity in Africa extends back well into the Middle Pliocene.

Was Australopithecus anamensis ancestral to A. afarensis? A case of anagenesis in the hominin fossil record.

We tested the hypothesis that early Pliocene Australopithecus anamensis was ancestral to A. afarensis by conducting a phylogenetic analysis of four temporally successive fossil samples assigned to these species (from earliest to latest: Kanapoi, Allia Bay, Laetoli, Hadar) using polarized character-state data from 20 morphological characters of the dentition and jaws. If the hypothesis that A. anamensis is ancestral to A. afarensis is true, then character-state changes between the temporally ordered site-samples should be congruent with hypothesized polarity transformations based on outgroup (African great ape) conditions. The most parsimonious reconstruction of character-state evolution suggests that each of the hominin OTUs shares apomorphies only with geologically younger OTUs, as predicted by the hypothesis of ancestry (tree length=31; Consistency Index=0.903). This concordance of stratigraphic and character-state data supports the idea that the A. anamensis and A. afarensis samples represent parts of an anagenetically evolving lineage, or evolutionary species. Each site-sample appears to capture a different point along this evolutionary trajectory. We discuss the implications of this conclusion for the taxonomy and adaptive evolution of these early-middle Pliocene hominins.

Enamel thickness, microstructure and development in Afropithecus turkanensis.

Afropithecus turkanensis, a 17-17.5 million year old large-bodied hominoid from Kenya, has previously been reported to be the oldest known thick-enamelled Miocene ape. Most investigations of enamel thickness in Miocene apes have been limited to opportunistic or destructive studies of small samples. Recently, more comprehensive studies of enamel thickness and microstructure in Proconsul, Lufengpithecus, and Dryopithecus, as well as extant apes and fossil humans, have provided information on rates and patterns of dental development, including crown formation time, and have begun to provide a comparative context for interpretation of the evolution of these characters throughout the past 20 million years of hominoid evolution. In this study, enamel thickness and aspects of the enamel microstructure in two A. turkanensis second molars were quantified and provide insight into rates of enamel apposition, numbers of cells actively secreting enamel, and the time required to form regions of the crown. The average value for relative enamel thickness in the two molars is 21.4, which is a lower value than a previous analysis of this species, but which is still relatively thick compared to extant apes. This value is similar to those of several Miocene hominoids, a fossil hominid, and modern humans. Certain aspects of the enamel microstructure are similar to Proconsul nyanzae, Dryopithecus laietanus, Lufengpithecus lufengensis, Graecopithecus freybergi and Pongo pygmaeus, while other features differ from extant and fossil hominoids. Crown formation times for the two teeth are 2.4-2.6 years and 2.9-3.1 years respectively. These times are similar to a number of extant and fossil hominoids, some of which appear to show additional developmental similarities, including thick enamel. Although thick enamel may be formed through several developmental pathways, most Miocene hominoids and fossil hominids with relatively thick enamel are characterized by a relatively long period of cuspal enamel formation and a rapid rate of enamel secretion throughout the whole cusp, but a shorter total crown formation time than thinner-enamelled extant apes.

A new skeleton of Theropithecus brumpti (Primates: Cercopithecidae) from Lomekwi, West Turkana, Kenya.

A relatively complete skeleton of the fossil papionin, Theropithecus brumpti, from the site of Lomekwi, west of Lake Turkana, Kenya, is here described. The specimen, KNM-WT 39368, was recovered at the site of LO 5 (3 degrees 51'N and 35 degrees 45'E), from sediments dated to approximately 3.3Ma. The skeleton is that of an old adult male and preserves a number of articulated elements, including most of the forelimbs and tail. The cranial morphology is that of a large, early T. brumpti, exhibiting a deep mandible with a deeply excavated mandibular corpus fossa, and mandibular alveoli and cheek teeth arrayed in a reversed Curve of Spee. The forelimb skeleton exhibits a unique mixture of characteristics generally associated with a terrestrial locomotor habitus, such as a narrow scapula and a highly stable elbow joint, combined with those more representative of habitual arborealists, such as muscle attachments reflecting a large rotator cuff musculature and a flexible shoulder joint. The forelimb of KNM-WT 39368 also presents several features, unique toTheropithecus, which represent adaptations for manual grasping and fine manipulation. These features include a large, retroflexed medial humeral epicondyle (to which large pronator, and carpal and digital flexor muscles attached) and proportions of the digital rays that denote capabilities for precise opposition between the thumb and index finger. Taken together, these features indicate that one of the earliest recognized representatives of Theropithecus exhibited the food harvesting and processing anatomy that distinguished the genus through time and that contributed to its success throughout the later Pliocene and Pleistocene. Based on the anatomy of KNM-WT 39368 and the known habitat preference of T. brumpti, the species is reconstructed as being a generally terrestrial but highly dexterous, very large-bodied, sexually dimorphic, and possibly folivorous papionin. T. brumpti was adapted for propulsive quadrupedal locomotion over generally even ground, and yet was highly adept at manual foraging. The estimate of 43.8kg body mass for KNM-WT 39368 renders unlikely the possibility that the species, or at least adult males of the species, were highly arboreal. T. brumpti, as represented by KNM-WT 39368, is seen as a large, colorfully decorated, and basically terrestrial papionin that was restricted to riverine forest habitats in the Lake Turkana Basin from the middle to latest Pliocene.