The first Miocene fossils from coastal woodlands in the southern East African Rift.

The Miocene was a key time in the evolution of African ecosystems witnessing the origin of the African apes and the isolation of eastern coastal forests through an expanding arid corridor. Until recently, however, Miocene sites from the southeastern regions of the continent were unknown. Here, we report the first Miocene fossil teeth from the shoulders of the Urema Rift in Gorongosa National Park, Mozambique. We provide the first 1) radiometric ages of the Mazamba Formation, 2) reconstructions of paleovegetation in the region based on pedogenic carbonates and fossil wood, and 3) descriptions of fossil teeth. Gorongosa is unique in the East African Rift in combining marine invertebrates, marine vertebrates, reptiles, terrestrial mammals, and fossil woods in coastal paleoenvironments. The Gorongosa fossil sites offer the first evidence of woodlands and forests on the coastal margins of southeastern Africa during the Miocene, and an exceptional assemblage of fossils including new species.

Pliocene hominins from East Turkana were associated with mesic environments in a semiarid basin.

During the middle Pliocene (∼3.8-3.2 Ma), both Australopithecus afarensis and Kenyanthropus platyops are known from the Turkana Basin, but between 3.60 and 3.44 Ma, most hominin fossils are found on the west side of Lake Turkana. Here, we describe a new hominin locality (ET03-166/168, Area 129) from the east side of the lake, in the Lokochot Member of the Koobi Fora Formation (3.60-3.44 Ma). To reconstruct the paleoecology of the locality and its surroundings, we combine information from sedimentology, the relative abundance of associated mammalian fauna, phytoliths, and stable isotopes from plant wax biomarkers, pedogenic carbonates, and fossil tooth enamel. The combined evidence provides a detailed view of the local paleoenvironment occupied by these Pliocene hominins, where a biodiverse community of primates, including hominins, and other mammals inhabited humid, grassy woodlands in a fluvial floodplain setting. Between <3.596 and 3.44 Ma, increases in woody vegetation were, at times, associated with increases in arid-adapted grasses. This suggests that Pliocene vegetation included woody species that were resilient to periods of prolonged aridity, resembling vegetation structure in the Turkana Basin today, where arid-adapted woody plants are a significant component of the ecosystem. Pedogenic carbonates indicate more woody vegetation than other vegetation proxies, possibly due to differences in temporospatial scale and ecological biases in preservation that should be accounted for in future studies. These new hominin fossils and associated multiproxy paleoenvironmental indicators from a single locale through time suggest that early hominin species occupied a wide range of habitats, possibly including wetlands within semiarid landscapes. Local-scale paleoecological evidence from East Turkana supports regional evidence that middle Pliocene eastern Africa may have experienced large-scale, climate-driven periods of aridity. This information extends our understanding of hominin environments beyond the limits of simple wooded, grassy, or mosaic environmental descriptions.

Genomic variation in baboons from central Mozambique unveils complex evolutionary relationships with other Papio species.

BACKGROUND: Gorongosa National Park in Mozambique hosts a large population of baboons, numbering over 200 troops. Gorongosa baboons have been tentatively identified as part of Papio ursinus on the basis of previous limited morphological analysis and a handful of mitochondrial DNA sequences. However, a recent morphological and morphometric analysis of Gorongosa baboons pinpointed the occurrence of several traits intermediate between P. ursinus and P. cynocephalus, leaving open the possibility of past and/or ongoing gene flow in the baboon population of Gorongosa National Park. In order to investigate the evolutionary history of baboons in Gorongosa, we generated high and low coverage whole genome sequence data of Gorongosa baboons and compared it to available Papio genomes. RESULTS: We confirmed that P. ursinus is the species closest to Gorongosa baboons. However, the Gorongosa baboon genomes share more derived alleles with P. cynocephalus than P. ursinus does, but no recent gene flow between P. ursinus and P. cynocephalus was detected when available Papio genomes were analyzed. Our results, based on the analysis of autosomal, mitochondrial and Y chromosome data, suggest complex, possibly male-biased, gene flow between Gorongosa baboons and P. cynocephalus, hinting to direct or indirect contributions from baboons belonging to the "northern" Papio clade, and signal the presence of population structure within P. ursinus. CONCLUSIONS: The analysis of genome data generated from baboon samples collected in central Mozambique highlighted a complex set of evolutionary relationships with other baboons. Our results provided new insights in the population dynamics that have shaped baboon diversity.

Early Pleistocene large mammals from Maka'amitalu, Hadar, lower Awash Valley, Ethiopia.

The Early Pleistocene was a critical time period in the evolution of eastern African mammal faunas, but fossil assemblages sampling this interval are poorly known from Ethiopia's Afar Depression. Field work by the Hadar Research Project in the Busidima Formation exposures (~2.7-0.8 Ma) of Hadar in the lower Awash Valley, resulted in the recovery of an early Homo maxilla (A.L. 666-1) with associated stone tools and fauna from the Maka'amitalu basin in the 1990s. These assemblages are dated to ~2.35 Ma by the Bouroukie Tuff 3 (BKT-3). Continued work by the Hadar Research Project over the last two decades has greatly expanded the faunal collection. Here, we provide a comprehensive account of the Maka'amitalu large mammals (Artiodactyla, Carnivora, Perissodactyla, Primates, and Proboscidea) and discuss their paleoecological and biochronological significance. The size of the Maka'amitalu assemblage is small compared to those from the Hadar Formation (3.45-2.95 Ma) and Ledi-Geraru (2.8-2.6 Ma) but includes at least 20 taxa. Bovids, suids, and Theropithecus are common in terms of both species richness and abundance, whereas carnivorans, equids, and megaherbivores are rare. While the taxonomic composition of the Maka'amitalu fauna indicates significant species turnover from the Hadar Formation and Ledi-Geraru deposits, turnover seems to have occurred at a constant rate through time as taxonomic dissimilarity between adjacent fossil assemblages is strongly predicted by their age difference. A similar pattern characterizes functional ecological turnover, with only subtle changes in dietary proportions, body size proportions, and bovid abundances across the composite lower Awash sequence. Biochronological comparisons with other sites in eastern Africa suggest that the taxa recovered from the Maka'amitalu are broadly consistent with the reported age of the BKT-3 tuff. Considering the age of BKT-3 and biochronology, a range of 2.4-1.9 Ma is most likely for the faunal assemblage.

Estimating origination times from the early hominin fossil record.

The age of the earliest recovered fossil evidence of a hominin taxon is all too often equated with that taxon's origination. However, the earliest known fossil record nearly always postdates, sometimes by a substantial period of time, the true origination of a taxon. Here we evaluate the first appearance records of the earliest potential hominins (Sahelanthropus, Ardipithecus, Orrorin), as well as of the genera Australopithecus, Homo, and Paranthropus, to illustrate the considerable uncertainty regarding the actual timing of origin of these taxa. By placing confidence intervals on the first appearance records of early hominin taxa, we can better evaluate patterns of hominin diversity, turnover, and potential correlations with climatic and environmental changes.

Divergence-time estimates for hominins provide insight into encephalization and body mass trends in human evolution.

Quantifying speciation times during human evolution is fundamental as it provides a timescale to test for the correlation between key evolutionary transitions and extrinsic factors such as climatic or environmental change. Here, we applied a total evidence dating approach to a hominin phylogeny to estimate divergence times under different topological hypotheses. The time-scaled phylogenies were subsequently used to perform ancestral state reconstructions of body mass and phylogenetic encephalization quotient (PEQ). Our divergence-time estimates are consistent with other recent studies that analysed extant species. We show that the origin of the genus Homo probably occurred between 4.30 and 2.56 million years ago. The ancestral state reconstructions show a general trend towards a smaller body mass before the emergence of Homo, followed by a trend towards a greater body mass. PEQ estimations display a general trend of gradual but accelerating encephalization evolution. The obtained results provide a rigorous temporal framework for human evolution.

The environments of Australopithecus anamensis at Allia Bay, Kenya: A multiproxy analysis of early Pliocene Bovidae.

Australopithecus anamensis, among the earliest fully bipedal hominin species, lived in eastern Africa around 4 Ma. Much of what is currently known about the paleoecology of A. anamensis comes from the type locality, Kanapoi, Kenya. Here, we extend knowledge of the range of environments occupied by A. anamensis by presenting the first multiproxy paleoecological analysis focusing on Bovidae excavated from another important locality where A. anamensis was recovered, locality 261-1 (ca. 3.97 Ma) at Allia Bay, East Turkana, Kenya. Paleoenvironments are reconstructed using astragalar ecomorphology, mesowear, hypsodonty index, and oxygen and carbon isotopes from dental enamel. We compare our results to those obtained from Kanapoi. Our results show that the bovid community composition is similar between the two fossil assemblages. Allia Bay and Kanapoi bovid astragalar ecomorphology spans the spectrum of modern morphologies indicative of grassland, woodland, and even forest-adapted forms. Dietary reconstructions based on stable isotopes, mesowear, and hypsodonty reveal that these bovids' diet encompassed the full C3 to C4 dietary spectrum and overlap in the two data sets. Our results allow us to confidently extend our reconstructions of the paleoenvironments of A. anamensis at Kanapoi to Allia Bay, where this pivotal hominin species is associated with heterogeneous settings including habitats with varying degrees of tree cover, including grasslands, bushlands, and woodlands.

Middle Pliocene hominin distribution patterns in Eastern Africa.

Abundance distributions of large mammals are underused in exploring how ecological pressures vary across contemporaneous sites in the fossil record. To investigate variation in relative abundance across contemporaneous Pliocene mammal communities, we examine the time interval between ∼3.6 and 3.22 Ma at four sites in the Afar and Turkana basins: Hadar and the lower Omo Valley in Ethiopia and East Turkana and West Turkana in Kenya. Taphonomic and collection biases are examined using skeletal parts, body size, and taxonomic data from database collections. Taphonomic biases due to geologic conditions and fossil collection affected all sites, but those in the Turkana Basin appeared particularly affected by collecting bias. As a result, hominin relative abundance is calculated separately using a taphonomic control taxon, which shares similar collection biases and size. Comparisons of mammalian taxonomic groups revealed that the Omo region was dominated by suids and cercopithecids. The other sites are dominated by open habitat and mixed habitat associated bovids. Hominins had higher abundance wherein the dominant mammal taxa indicate a mix of woodland and grassland environments (Hadar) and were rarer at sites where the majority of taxa are associated with woodland vegetation (the Omo Valley). West Turkana is characterized by mixed habitats and the highest relative abundance of hominins relative to control taxa, but sampling issues due to the collection and reporting of papionins likely drive this result. East Turkana has few hominins relative to the control taxon and has dominant habitats indicative of floodplain grasslands but has a small sample size compared with the other sites. These analyses suggest that Kenyanthropus platyops and Australopithecus afarensis inhabited similar types of habitats across different rift basins. Most convincingly, this study contributes to a growing body of evidence suggesting that early hominins diverged from their great ape counterparts by abandoning woodland-dominated habitats.

Dietary trends in herbivores from the Shungura Formation, southwestern Ethiopia.

Diet provides critical information about the ecology and environment of herbivores. Hence, understanding the dietary strategies of fossil herbivores and the associated temporal changes is one aspect of inferring paleoenvironmental conditions. Here, we present carbon isotope data from more than 1,050 fossil teeth that record the dietary patterns of nine herbivore families in the late Pliocene and early Pleistocene (3.6 to 1.05 Ma) from the Shungura Formation, a hominin-bearing site in southwestern Ethiopia. An increasing trend toward C4 herbivory has been observed with attendant reductions in the proportions of browsers and mixed feeders through time. A high proportion of mixed feeders has been observed prior to 2.9 Ma followed by a decrease in the proportion of mixed feeders and an increase in grazers between 2.7 and 1.9 Ma, and a further increase in the proportion of grazers after 1.9 Ma. The collective herbivore fauna shows two major change points in carbon isotope values at ∼2.7 and ∼2.0 Ma. While hominin fossils from the sequence older than 2.7 Ma are attributed to Australopithecus, the shift at ∼2.7 Ma indicating the expansion of C4 grasses on the landscape was concurrent with the first appearance of Paranthropus The link between the increased C4 herbivory and more open landscapes suggests that Australopithecus lived in more wooded landscapes compared to later hominins such as Paranthropus and Homo, and has implications for key morphological and behavioral adaptations in our lineage.

Isotopic evidence for the timing of the dietary shift toward C4 foods in eastern African Paranthropus.

New approaches to the study of early hominin diets have refreshed interest in how and when our diets diverged from those of other African apes. A trend toward significant consumption of C4 foods in hominins after this divergence has emerged as a landmark event in human evolution, with direct evidence provided by stable carbon isotope studies. In this study, we report on detailed carbon isotopic evidence from the hominin fossil record of the Shungura and Usno Formations, Lower Omo Valley, Ethiopia, which elucidates the patterns of C4 dietary utilization in the robust hominin Paranthropus The results show that the most important shift toward C4 foods occurred at ∼2.37 Ma, within the temporal range of the earliest known member of the genus, Paranthropus aethiopicus, and that this shift was not unique to Paranthropus but occurred in all hominins from this fossil sequence. This uptake of C4 foods by hominins occurred during a period marked by an overall trend toward increased C4 grazing by cooccurring mammalian taxa from the same sequence. However, the timing and geographic patterns of hominin diets in this region differ from those observed elsewhere in the same basin, where environmental controls on the underlying availability of various food sources were likely quite different. These results highlight the complexities of dietary responses by hominins to changes in the availability of food resources.

Fossils from Mille-Logya, Afar, Ethiopia, elucidate the link between Pliocene environmental changes and Homo origins.

Several hypotheses posit a link between the origin of Homo and climatic and environmental shifts between 3 and 2.5 Ma. Here we report on new results that shed light on the interplay between tectonics, basin migration and faunal change on the one hand and the fate of Australopithecus afarensis and the evolution of Homo on the other. Fieldwork at the new Mille-Logya site in the Afar, Ethiopia, dated to between 2.914 and 2.443 Ma, provides geological evidence for the northeast migration of the Hadar Basin, extending the record of this lacustrine basin to Mille-Logya. We have identified three new fossiliferous units, suggesting in situ faunal change within this interval. While the fauna in the older unit is comparable to that at Hadar and Dikika, the younger units contain species that indicate more open conditions along with remains of Homo. This suggests that Homo either emerged from Australopithecus during this interval or dispersed into the region as part of a fauna adapted to more open habitats.

The ecology of Australopithecus anamensis in the early Pliocene of Kanapoi, Kenya.

Australopithecus anamensis is a pivotal species in human evolution. It is likely to be the direct ancestor of Australopithecus afarensis and the species that may have given rise to the Homo and Paranthropus lineages. It had a suite of adaptations for habitual bipedalism and a diet that differed from that of earlier hominin species. Under what environmental and ecological conditions did this suite of adaptations arise? The early Pliocene site of Kanapoi in the Lake Turkana Basin of Kenya has the largest sample of A. anamensis in eastern Africa and a rich record of fossil vertebrates. Most Kanapoi fossils are chronologically well constrained by radiometrically dated tephras between the ages of 4.2 and 4.1 million years ago. Sedimentological, isotopic, and faunal data indicate that the environments of Kanapoi during the early Pliocene had a complex range of vegetation types that included closed woodlands, shrubs, and grasslands near a river (for most of the sequence) or lake. These were dynamic landscapes that could shift rapidly from fluvial to lacustrine conditions, and then back. Australopithecus anamensis shared its environments with at least 10 species of very large herbivores, which undoubtedly played a major role in modifying the landscape by opening wooded areas and providing pathways for bipedal hominins. Hominins may have competed for terrestrial resources with abundant suids (Nyanzachoerus and Notochoerus) and for arboreal resources with monkeys (Parapapio being the most common cercopithecid). Kanapoi had a formidable group of predators that included a very abundant species of hyena (Parahyaena howelli), two sabre-tooth felids (Dinofelis and Homotherium), a giant otter (Enhydriodon cf. dikikae), and three species of crocodiles. Various measures of abundance indicate that A. anamensis was an important component of the Kanapoi early Pliocene ecosystems, and that its key adaptations allowed this species to thrive in complex and dynamic landscapes.

Paleoecological implications of dental mesowear and hypsodonty in fossil ungulates from Kanapoi.

The Pliocene site of Kanapoi is key to our understanding of the environmental context of the earliest species of Australopithecus. Various approaches have been used to reconstruct the environments of this site, and here we contribute new data and analyses using mesowear and hypsodonty. The dental traits of 98 bovids, suids and rhinocerotids from Kanapoi were analyzed using these proxies. Results indicate that most of the animals analyzed had a relatively abrasive diet. Bovids in the assemblage incorporated more grass into their diet than do modern species of the same tribe or genus. Although Pliocene Kanapoi likely had complex environments, our analysis indicates that grassy habitats were a dominant component of the ecosystem, a conclusion that supports the results of previous investigations of the paleoecology of the site.

Ruminants (Giraffidae and Bovidae) from Kanapoi.

We update here our recent revision of the Kanapoi ruminants and describe recently collected material. We now regard the occurrence of reduncins as doubtful, we revise the identification of a large raphicerin as being more probably Gazella, and we add Gazella cf. janenschi and the Cephalophini to the faunal list. New material of Tragelaphus kyaloi suggests that this species held its head unlike other tragelaphins, and was not an exclusive dedicated browser, but Kanapoi pre-dates the Pliocene change of Sivatherium, Aepyceros, Alcelaphini, and even Tragelaphini toward more grazing diets. Kanapoi shares several ruminant taxa with sites in Ethiopia and Tanzania, attesting to latitudinal exchanges.

Suidae from Kanapoi.

The whole collection of Suidae from Kanapoi is revised in the context of the systematics and evolution of Nyanzachoerus in the Pliocene of Eastern Africa. It contains only two species, Nyanzachoerus kanamensis and Notochoerus jaegeri. The size and morphology of their premolars overlap, but not those of their m3s. No transitional form between them is known in Kenya, but some populations from Uganda and Ethiopia display intermediate characters, suggesting that No. jaegeri could be descended from a kanamensis-like ancestor. However, the cranial remains of No. jaegeri from Kanapoi are insufficient to formally establish the affinities of the species. On the basis of the dentition, Notochoerus euilus could be descended from No. jaegeri. The noticeable absence of Kolpochoerus at Kanapoi (and in the whole Turkana Basin at that time) remains unexplained. The presence of a species with affinity to Nyanzachoerus tulotos at Ekora raises the possibility that uppermost Miocene sediments occur there.

Reorganization of surviving mammal communities after the end-Pleistocene megafaunal extinction.

Large mammals are at high risk of extinction globally. To understand the consequences of their demise for community assembly, we tracked community structure through the end-Pleistocene megafaunal extinction in North America. We decomposed the effects of biotic and abiotic factors by analyzing co-occurrence within the mutual ranges of species pairs. Although shifting climate drove an increase in niche overlap, co-occurrence decreased, signaling shifts in biotic interactions. Furthermore, the effect of abiotic factors on co-occurrence remained constant over time while the effect of biotic factors decreased. Biotic factors apparently played a key role in continental-scale community assembly before the extinctions. Specifically, large mammals likely promoted co-occurrence in the Pleistocene, and their loss contributed to the modern assembly pattern in which co-occurrence frequently falls below random expectations.

Comparative isotopic evidence from East Turkana supports a dietary shift within the genus Homo.

It has been suggested that a shift in diet is one of the key adaptations that distinguishes the genus Homo from earlier hominins, but recent stable isotopic analyses of fossils attributed to Homo in the Turkana Basin show an increase in the consumption of C4 resources circa 1.65 million years ago, significantly after the earliest evidence for Homo in the eastern African fossil record. These data are consistent with ingesting more C4 plants, more animal tissues of C4 herbivores, or both, but it is also possible that this change reflects factors unrelated to changes in the palaeobiology of the genus Homo. Here we use new and published carbon and oxygen isotopic data (n = 999) taken from large-bodied fossil mammals, and pedogenic carbonates in fossil soils, from East Turkana in northern Kenya to investigate the context of this change in the isotope signal within Homo. By targeting taxa and temporal intervals unrepresented or undersampled in previous analyses, we were able to conduct the first comprehensive analysis of the ecological context of hominin diet at East Turkana during a period crucial for detecting any dietary and related behavioural differences between early Homo (H. habilis and/or H. rudolfensis) and Homo erectus. Our analyses suggest that the genus Homo underwent a dietary shift (as indicated by δ13Cena and δ18Oena values) that is (1) unrelated to changes in the East Turkana vegetation community and (2) unlike patterns found in other East Turkana large mammals, including Paranthropus and Theropithecus. These data suggest that within the Turkana Basin a dietary shift occurred well after we see the first evidence of early Homo in the region.