Plio-Pleistocene environmental variability in Africa and its implications for mammalian evolution.

Understanding the climatic drivers of environmental variability (EV) during the Plio-Pleistocene and EV's influence on mammalian macroevolution are two outstanding foci of research in African paleoclimatology and evolutionary biology. The potential effects of EV are especially relevant for testing the variability selection hypothesis, which predicts a positive relationship between EV and speciation and extinction rates in fossil mammals. Addressing these questions is stymied, however, by 1) a lack of multiple comparable EV records of sufficient temporal resolution and duration, and 2) the incompleteness of the mammalian fossil record. Here, we first compile a composite history of Pan-African EV spanning the Plio-Pleistocene, which allows us to explore which climatic variables influenced EV. We find that EV exhibits 1) a long-term trend of increasing variability since ∼3.7 Ma, coincident with rising variability in global ice volume and sea surface temperatures around Africa, and 2) a 400-ky frequency correlated with seasonal insolation variability. We then estimate speciation and extinction rates for fossil mammals from eastern Africa using a method that accounts for sampling variation. We find no statistically significant relationship between EV and estimated speciation or extinction rates across multiple spatial scales. These findings are inconsistent with the variability selection hypothesis as applied to macroevolutionary processes.

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.

Geographically divergent evolutionary and ecological legacies shape mammal biodiversity in the global tropics and subtropics.

Studies of the factors governing global patterns of biodiversity are key to predicting community responses to ongoing and future abiotic and biotic changes. Although most research has focused on present-day climate, a growing body of evidence indicates that modern ecological communities may be significantly shaped by paleoclimatic change and past anthropogenic factors. However, the generality of this pattern is unknown, as global analyses are lacking. Here we quantify the phylogenetic and functional trait structure of 515 tropical and subtropical large mammal communities and predict their structure from past and present climatic and anthropogenic factors. We find that the effects of Quaternary paleoclimatic change are strongest in the Afrotropics, with communities in the Indomalayan realm showing mixed effects of modern climate and paleoclimate. Malagasy communities are poorly predicted by any single factor, likely due to the atypical history of the island compared with continental regions. Neotropical communities are mainly codetermined by modern climate and prehistoric and historical human impacts. Overall, our results indicate that the factors governing tropical and subtropical mammalian biodiversity are complex, with the importance of past and present factors varying based on the divergent histories of the world's biogeographic realms and their native biotas. Consideration of the evolutionary and ecological legacies of both the recent and ancient past are key to understanding the forces shaping global patterns of present-day biodiversity and its response to ongoing and future abiotic and biotic changes in the 21st century.

Earliest known Oldowan artifacts at >2.58 Ma from Ledi-Geraru, Ethiopia, highlight early technological diversity.

The manufacture of flaked stone artifacts represents a major milestone in the technology of the human lineage. Although the earliest production of primitive stone tools, predating the genus Homo and emphasizing percussive activities, has been reported at 3.3 million years ago (Ma) from Lomekwi, Kenya, the systematic production of sharp-edged stone tools is unknown before the 2.58-2.55 Ma Oldowan assemblages from Gona, Ethiopia. The organized production of Oldowan stone artifacts is part of a suite of characteristics that is often associated with the adaptive grade shift linked to the genus Homo Recent discoveries from Ledi-Geraru (LG), Ethiopia, place the first occurrence of Homo ∼250 thousand years earlier than the Oldowan at Gona. Here, we describe a substantial assemblage of systematically flaked stone tools excavated in situ from a stratigraphically constrained context [Bokol Dora 1, (BD 1) hereafter] at LG bracketed between 2.61 and 2.58 Ma. Although perhaps more primitive in some respects, quantitative analysis suggests the BD 1 assemblage fits more closely with the variability previously described for the Oldowan than with the earlier Lomekwian or with stone tools produced by modern nonhuman primates. These differences suggest that hominin technology is distinctly different from generalized tool use that may be a shared feature of much of the primate lineage. The BD 1 assemblage, near the origin of our genus, provides a link between behavioral adaptations-in the form of flaked stone artifacts-and the biological evolution of our ancestors.

Late Pliocene environmental change during the transition from Australopithecus to Homo.

It has long been hypothesized that the transition from Australopithecus to Homo in eastern Africa was linked to the spread of open and arid environments near the Plio-Pleistocene boundary, but data for the latest Pliocene are scarce. Here we present new stable carbon isotope data from the late Pliocene mammalian fauna from Ledi-Geraru, in the lower Awash Valley (LAV), Ethiopia, and mammalian community analyses from the LAV and Turkana Basin. These data, combined with pedogenic carbonate stable isotopes, indicate that the two regions were largely similar through the Plio-Pleistocene, but that important environmental differences existed during the emergence of Homo around 2.8 million years ago. The mid-Pliocene to late Pliocene interval in the LAV was characterized by increasingly C4-dominated, arid and seasonal environments. The early Homo mandible LD 350-1 has a carbon isotope value similar to that of earlier Australopithecus from the LAV, possibly indicating that the emergence of Homo from Australopithecus did not involve a dietary shift. Late Pliocene LAV environments contrast with contemporaneous environments in the Turkana Basin, which were more woody and mesic. These findings have important implications for the environmental conditions surrounding the emergence of Homo, as well as recent hypotheses regarding Plio-Pleistocene environmental change in eastern Africa.

Strong influence of palaeoclimate on the structure of modern African mammal communities.

Ecological research often assumes that species are adapted to their current climatic environments. However, climate fluctuations over geologic timescales have influenced species dispersal and extinction, which in turn may affect community structure. Modern community structure is likely to be the product of both palaeoclimate and modern climate, with the relative degrees of influence of past and present climates unknown. Here, we assessed the influence of climate at different time periods on the phylogenetic and functional trait structure of 203 African mammal communities. We found that the climate of the mid-Holocene (approx. 6000 years ago) and Last Glacial Maximum (approx. 22 000 years ago) were frequently better predictors of community structure than modern climate for mammals overall, carnivorans and ungulates. Primate communities were more strongly influenced by modern climate than palaeoclimate. Overall, community structure of African mammals appears to be related to the ecological flexibility of the groups considered here and the regions of continental Africa that they occupy. Our results indicate that the future redistribution, expansion and contraction of particular biomes due to human activity, such as climate and land-use change, will differentially affect mammal groups that vary in their sensitivity to environmental change.

Dietary flexibility of Australopithecus afarensis in the face of paleoecological change during the middle Pliocene: Faunal evidence from Hadar, Ethiopia.

One approach to understanding the context of changes in hominin paleodiets is to examine the paleodiets and paleohabitats of contemporaneous mammalian taxa. Recent carbon isotopic studies suggest that the middle Pliocene was marked by a major shift in hominin diets, characterized by a significant increase in C4 foods in Australopithecus-grade species, including Australopithecus afarensis. To contextualize previous isotopic studies of A. afarensis, we employed stable isotopes to examine paleodiets of the mammalian fauna contemporaneous with A. afarensis at Hadar, Ethiopia. We used these data to inform our understanding of paleoenvironmental change through the deposition of the Hadar Formation. While the majority of the taxa in the Hadar fauna were C4 grazers, most show little change in the intensity of C4 food consumption over the 0.5 million-year interval sampled. Two taxa (equids and bovins) do show increases in C4 consumption through the Hadar Formation and into the younger, overlying Busidima Formation. Changes in the distributions of C4-feeders, C3-feeders and mixed-C3/C4-feeders in the sampled intervals are consistent with evidence of dietary reconstructions based on ecomorphology, and with habitats reconstructed using community structure analyses. Meanwhile, A. afarensis is one of many mammalian taxa whose C4 consumption does not show directional change over the intervals sampled. In combination with a wide range of carbon and oxygen isotopic composition for A. afarensis as compared to the other large mammal taxa, these results suggest that the C3/C4 dietary flexibility of A. afarensis was relatively unusual among most of its mammalian cohort.

PALEOANTHROPOLOGY. Response to Comment on "Early Homo at 2.8 Ma from Ledi-Geraru, Afar, Ethiopia".

Hawks et al. argue that our analysis of Australopithecus sediba mandibles is flawed and that specimen LD 350-1 cannot be distinguished from this, or any other, Australopithecus species. Our reexamination of the evidence confirms that LD 350-1 falls outside of the pattern that A. sediba shares with Australopithecus and thus is reasonably assigned to the genus Homo.

Climate and species richness predict the phylogenetic structure of African mammal communities.

We have little knowledge of how climatic variation (and by proxy, habitat variation) influences the phylogenetic structure of tropical communities. Here, we quantified the phylogenetic structure of mammal communities in Africa to investigate how community structure varies with respect to climate and species richness variation across the continent. In addition, we investigated how phylogenetic patterns vary across carnivores, primates, and ungulates. We predicted that climate would differentially affect the structure of communities from different clades due to between-clade biological variation. We examined 203 communities using two metrics, the net relatedness (NRI) and nearest taxon (NTI) indices. We used simultaneous autoregressive models to predict community phylogenetic structure from climate variables and species richness. We found that most individual communities exhibited a phylogenetic structure consistent with a null model, but both climate and species richness significantly predicted variation in community phylogenetic metrics. Using NTI, species rich communities were composed of more distantly related taxa for all mammal communities, as well as for communities of carnivorans or ungulates. Temperature seasonality predicted the phylogenetic structure of mammal, carnivoran, and ungulate communities, and annual rainfall predicted primate community structure. Additional climate variables related to temperature and rainfall also predicted the phylogenetic structure of ungulate communities. We suggest that both past interspecific competition and habitat filtering have shaped variation in tropical mammal communities. The significant effect of climatic factors on community structure has important implications for the diversity of mammal communities given current models of future climate change.

Paleoanthropology. Early Homo at 2.8 Ma from Ledi-Geraru, Afar, Ethiopia.

Our understanding of the origin of the genus Homo has been hampered by a limited fossil record in eastern Africa between 2.0 and 3.0 million years ago (Ma). Here we report the discovery of a partial hominin mandible with teeth from the Ledi-Geraru research area, Afar Regional State, Ethiopia, that establishes the presence of Homo at 2.80 to 2.75 Ma. This specimen combines primitive traits seen in early Australopithecus with derived morphology observed in later Homo, confirming that dentognathic departures from the australopith pattern occurred early in the Homo lineage. The Ledi-Geraru discovery has implications for hypotheses about the timing and place of origin of the genus Homo.

Paleoanthropology. Late Pliocene fossiliferous sedimentary record and the environmental context of early Homo from Afar, Ethiopia.

Sedimentary basins in eastern Africa preserve a record of continental rifting and contain important fossil assemblages for interpreting hominin evolution. However, the record of hominin evolution between 3 and 2.5 million years ago (Ma) is poorly documented in surface outcrops, particularly in Afar, Ethiopia. Here we present the discovery of a 2.84- to 2.58-million-year-old fossil and hominin-bearing sediments in the Ledi-Geraru research area of Afar, Ethiopia, that have produced the earliest record of the genus Homo. Vertebrate fossils record a faunal turnover indicative of more open and probably arid habitats than those reconstructed earlier in this region, which is in broad agreement with hypotheses addressing the role of environmental forcing in hominin evolution at this time. Geological analyses constrain depositional and structural models of Afar and date the LD 350-1 Homo mandible to 2.80 to 2.75 Ma.

Diet of Australopithecus afarensis from the Pliocene Hadar Formation, Ethiopia.

The enhanced dietary flexibility of early hominins to include consumption of C4/crassulacean acid metabolism (CAM) foods (i.e., foods derived from grasses, sedges, and succulents common in tropical savannas and deserts) likely represents a significant ecological and behavioral distinction from both extant great apes and the last common ancestor that we shared with great apes. Here, we use stable carbon isotopic data from 20 samples of Australopithecus afarensis from Hadar and Dikika, Ethiopia (>3.4-2.9 Ma) to show that this species consumed a diet with significant C4/CAM foods, differing from its putative ancestor Au. anamensis. Furthermore, there is no temporal trend in the amount of C4/CAM food consumption over the age of the samples analyzed, and the amount of C4/CAM food intake was highly variable, even within a single narrow stratigraphic interval. As such, Au. afarensis was a key participant in the C4/CAM dietary expansion by early australopiths of the middle Pliocene. The middle Pliocene expansion of the eastern African australopith diet to include savanna-based foods represents a shift to use of plant food resources that were already abundant in hominin environments for at least 1 million y and sets the stage for dietary differentiation and niche specialization by subsequent hominin taxa.

On the industrial attributions of the Aterian and Mousterian of the Maghreb.

North Africa is quickly emerging as one of the more important regions yielding information on the origins of modern Homo sapiens. Associated with significant fossil hominin remains are two stone tool industries, the Aterian and Mousterian, which have been differentiated, respectively, primarily on the basis of the presence and absence of tanged, or stemmed, stone tools. Largely because of historical reasons, these two industries have been attributed to the western Eurasian Middle Paleolithic rather than the African Middle Stone Age. In this paper, drawing on our recent excavation of Contrebandiers Cave and other published data, we show that, aside from the presence or absence of tanged pieces, there are no other distinctions between these two industries in terms of either lithic attributes or chronology. Together, these results demonstrate that these two 'industries' are instead variants of the same entity. Moreover, several additional characteristics of these assemblages, such as distinctive stone implements and the manufacture and use of bone tools and possible shell ornaments, suggest a closer affinity to other Late Pleistocene African Middle Stone Age industries rather than to the Middle Paleolithic of western Eurasia.

The enigmatic molar from Gondolin, South Africa: implications for Paranthropus paleobiology.

The specific attribution of the large hominin M(2) (GDA-2) from Gondolin has significant implications for the paleobiology of Paranthropus. If it is a specimen of Paranthropus robustus it impacts that species' size range, and if it belongs to Paranthropus boisei it has important biogeographic implications. We evaluate crown size, cusp proportions and the likelihood of encountering a large-bodied mammal species in both East and South Africa in the Early Pleistocene. The tooth falls well outside the P. robustus sample range, and comfortably within that for penecontemporaneous P. boisei. Analyses of sample range, distribution and variability suggest that it is possible, albeit unlikely to find a M(2) of this size in the current P. robustus sample. However, taphonomic agents - carnivore (particularly leopard) feeding behaviors - have likely skewed the size distribution of the Swartkrans and Drimolen P. robustus assemblage. In particular, assemblages of large-bodied mammals accumulated by leopards typically display high proportions of juveniles and smaller adults. The skew in the P. robustus sample is consistent with this type of assemblage. Morphological evidence in the form of cusp proportions is congruent with GDA-2 representing P. robustus rather than P. boisei. The comparatively small number of large-bodied mammal species common to both South and East Africa in the Early Pleistocene suggests a low probability of encountering an herbivorous australopith in both. Our results are most consistent with the interpretation of the Gondolin molar as a very large specimen of P. robustus. This, in turn, suggests that large, presumptive male, specimens are rare, and that the levels of size variation (sexual dimorphism) previously ascribed to this species are likely to be gross underestimates.

Middle and late Pleistocene faunas of Pinnacle Point and their paleoecological implications.

The Western Cape region of South Africa is home to a unique type of mediterranean vegetation called fynbos, as well as some of the earliest sites of modern human occupation in southern Africa. Reconstructing the paleohabitats during occupations of these early anatomically modern Homo sapiens is important for understanding the availability of resources to the humans during the development of behaviors that are often considered advanced. These reconstructions are critical to understanding the nature of the changes in the environment and resources over time. Here we analyze the craniodental fossils of the larger mammals recovered from two Pleistocene assemblages in the Pinnacle Point complex, Mossel Bay, Western Cape Region, South Africa. We reconstruct the paleohabitats as revealed by multivariate analyses of the mammalian community structures. Pinnacle Point 30 is a carnivore assemblage and Pinnacle Point 13B includes early evidence of a suite of modern human behavior; together they present an opportunity to identify environmental change over time at a localized geographic scale. Further, this is the first such study to include dated Western Cape localities from Marine Isotope Stage 6, a time of environmental pressure that may have marginalized human populations. Results indicate that environmental change in the Western Cape was more complex than generalized C(4) grassland expansions replacing fynbos habitats during glacial lowered sea levels, and thus, resources available to early modern humans in the region may not have been entirely predictable.

The large-mammal fauna from the Kibish Formation.

The Kibish faunal remains are useful for reconstructing the habitat of the earliest documented Homo sapiens and for understanding the community within which early modern humans existed. A diverse assemblage of large mammals, including many species of bovids, suids, and equids, has been recovered from the Kibish Formation. There are no extinct large mammals represented in the fossil assemblage, and the overall taxonomic composition of the fossil fauna is similar to the modern-day wildlife community living near the Omo River. The fossil faunal assemblage shows a paucity of arboreal primates, and carnivore species are rare. However, the faunal sample includes possible Cephalophus (duiker) remains and Hylochoerus meinertzhageni (giant forest hog), taxa that are extremely rare in the African fossil record, and both indicate more closed habitats. Comparative analyses of the Kibish faunal remains using the ecological-diversity approach document close associations with edaphic grassland and woodland vegetation types. These vegetation forms are similar to current habitats surrounding the Omo River.

Paleoecological patterns at the Hadar hominin site, Afar Regional State, Ethiopia.

Reconstructing paleoecological patterns associated with hominin taxa, such as Australopithecus afarensis, is important for understanding possible evolutionary mechanisms involved in extinction and speciation events. It is critical to identify local, regional, or pan-African causal factors because patterns at these different levels may affect separate populations of the same species of hominin in unique ways. Habitat reconstructions of 12 submembers of the Hadar and Busidima formations (approximately 3.8-2.35 Ma) are presented here along with faunal differences in these submembers through time. Habitats with medium density tree and bush cover dominated the landscape through much of the earlier time period in the Hadar Formation. The lowermost Sidi Hakoma Member is the most closed habitat. The Denen Dora Member shows the influence of frequent floodplain edaphic grasslands with high abundances of reducin bovids. There is an influx of ungulates in the Kada Hadar Member (approximately 3.2--approximately 2.96 Ma) that indicates a more arid habitat populated by mammals that were recovered from earlier deposits further south in Ethiopia and Kenya. In the younger deposits from the Busidima Formation at Hadar, the landscape was open wooded grassland with some floodplain environments. The fossil assemblages from the Busidima Formation show a substantial species turnover. Although high numbers of A. afarensis specimens are associated with the lower Sidi Hakoma Member, they clearly inhabited a variety of habitats throughout the entire Hadar Formation. Australopithecus afarensis from Laetoli through Hadar times appears to have been a eurytopic species.

Primate communities: past, present, and possible future.

An understanding of the fundamental causes of the structure of primate communities is important for studies of primate evolutionary history, primate behavioral ecology, and development of conservation strategies. Research into these structuring factors has benefited from new perspectives such as consideration of primate phylogenetic history, metacommunities, and interactions with predators and nonprimate competitors. This review presents the underlying factors of primate community structure within the biogeographic regions of Madagascar, the Neotropics, Africa, and Asia. One of the major differences among these locations likely resulted from the initial primate taxa that colonized each region (a single colonization event in the case of Madagascar and South America, and multiple radiations of higher-level taxa in Africa and Asia). As most primates live in forests, the differences among the forests in these locations, caused by various climatic influences, further influenced speciation and the development of primate communities. Within these habitats, species interactions with different groups of organisms were also instrumental in developing community dynamics. Through an investigation of these fundamental factors, we identify some of the most important effects on primate communities in each region. These findings suggest that low primate richness in Asia may be caused by either the abundance of dipterocarp trees or high levels of monsoon rains. High numbers of frugivores and a lack of folivores in neotropical communities may be associated with competition with sloths that were already present at the time of initial radiation. Climatic patterns which affect forest structure and productivity in Madagascar may be responsible for high numbers of folivorous lemurs. The identification of these factors are important for the conservation of existing primate communities, and indicate directions for future studies.