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The archaeological and paleontological records from the west coast of South Africa have potential to provide insights into ecosystem dynamics in the region during the mid Pleistocene. Although the fossil record suggests an ecosystem quite different than that of the region today, we understand little about the ecological factors that contributed to this disparity. The site of Elandsfontein (EFT) dates to between 1.0 and 0.6 million years ago (Ma), preserves in situ lithic and faunal materials found in direct association with each other, and provides the rare opportunity to examine the relationship between hominin behavioural variability and landscape heterogeneity in a winter rainfall ecosystem. In this study, we examine the stable carbon isotopic composition of a large sample (n = 81) of Cape dune mole-rats (Bathyergus suillus) and contemporaneous large mammals (> 6 kg; n = 194) from EFT. We find that δ13C values of B. suillus are significantly different to those of contemporaneous large mammals from EFT indicating a significant presence of plants utilizing the C4 photosynthetic pathway during the mid-Pleistocene, in contrast to present C3 dominated ecosystems along the west coast of South Africa. Additionally, we find that artifact density at EFT localities is positively correlated with δ13C values in B. suillus enamel suggesting that evidence of more intense hominin occupation may be associated with the presence of more C4 vegetation. Lastly, we hypothesize that this unique distribution of vegetation 1) provided abundant resources for both hominin and non-hominin taxa and 2) may have concentrated hominin and animal behavior in certain places on the ancient landscape.

Nearly all primates are ecologically dependent on trees, but they are nonetheless found in an enormous range of habitats, from highly xeric environments to dense rainforest. Most primates have a relatively ‘generalised’ skeleton, enabling locomotor flexibility and facilitating other crucial functions, such as manual foraging and grooming. This paper explores the associations between habitat, locomotion and morphology in the forelimbs of cercopithecids (Old World monkeys), contextualising their skeletal ecomorphological patterns with those of other mammals, and complementing functional morphological analyses with phylogenetic comparative techniques. The ecomorphological signals present in the generalised primate postcranium, and how an ancestral arboreal ‘bauplan’ might be modified to incorporate terrestriality or exploit distinct arboreal substrates, are investigated. Analysis of ecomorphological variation in guenons indicates that terrestrial Chlorocebus species retain core elements of a general guenon form, with modifications for terrestriality that vary by species. Adaptation to different modes of arboreality has also occurred in Cercopithecus. The considerable morphological similarity in the guenons sampled emphasises the importance of generality in the primate postcranium – much forelimb variation appears to have emerged stochastically, with a smaller number of traits having a strong functional signal. Analysis of a broader sample of cercopithecids and comparison with felids, suids and bovids indicates that although the cercopithecid humerus has functional morphological signals that enable specimens to be assigned with a reasonable degree of certainty to habitat groups, there is considerable overlap in the specimens assigned to each habitat group. This probably reflects ecological dependence on trees, even in predominantly terrestrial species, as well as the multiple functions of the forelimb and, in some cases, wide geographic distributions that promote intraspecific variation. The use of phylogenetic correction reduced the discriminatory power of the models, indicating that, like allometry, phylogeny contains important ecomorphological information, and should not necessarily be factored out of analyses.

Mammalian carnivores are rarely incorporated in paleoenvironmental reconstructions, largely because of their rarity within the fossil record. However, multivariate statistical modeling can be successfully used to quantify specific anatomical features as environmental predictors. Here we explore morphological variability of the humerus in a closely related group of predators (Felidae) to investigate the relationship between morphometric descriptors and habitat categories. We analyze linear measurements of the humerus in three different morphometric combinations (log-transformed, size-free, and ratio), and explore four distinct ways of categorizing habitat adaptations. Open, Mixed, and Closed categories are defined according to criteria based on traditional descriptions of species, distributions, and biome occupancy. Extensive exploratory work is presented using linear discriminant analyses and several fossils are included to provide paleoecological reconstructions.We found no significant differences in the predictive power of distinct morphometric descriptors or habitat criteria, although sample splitting into small and large cat guilds greatly improves the stability of the models. Significant insights emerge for three long-canine cats:Smilodon populator,Paramachairodus orientalis, andDinofelissp. from Olduvai Gorge (East Africa).S. populatorandP. orientalisare both predicted to have been closed-habitat adapted taxa. The false “sabertooth”Dinofelissp. from Olduvai Gorge is predicted to be adapted to mixed habitat. The application of felid humerus ecomorphology to the carnivoran record of Olduvai Gorge shows that the older stratigraphic levels (Bed I, 1.99–1.79 Ma) included a broader range of environments than Beds II or V, where there is an abundance of cats adapted to open environments.

The middle Pleistocene fossil mammal assemblage from Lainyamok in the southern Kenya rift has previously been considered the oldest (330-392 ka) African mammal community consisting entirely of extant species, with the dominant bovid tentatively attributed to the southern African blesbok (Damaliscus cf. dorcas). We show that the blesbok-like fossils from Lainyamok belong to an extinct species, described here as Damaliscus hypsodon sp. nov. The D. hypsodon hypodigm includes the previously unnamed small alcelaphine material known from late Pleistocene sites elsewhere in Kenya and Tanzania. Its dental anatomy, together with an ecomorphological analysis of its postcrania, indicates that D. hypsodon grazed in open and arid grassland environments. Although Lainyamok is no longer represented entirely by extant species, the absence of species common earlier in the middle Pleistocene of East Africa suggests substantial faunal turnover between 500 and 400 ka. Damaliscus hypsodon persisted in East Africa until the end of the Pleistocene and its extinction can be attributed to a loss of arid grassland environments at the onset of the Holocene. The fossil evidence from southern Kenya suggests that the development of the taxonomically modern large mammal community was a long-term process characterized by the extinction of grazing specialists, with marked turnover occurring between ~ 500 and 400 ka and near the end of the Pleistocene.

Morphological convergence amongst species inhabiting similar environments but having different evolutionary histories is a concept central to evolutionary biology. Cases of divergent evolution, where there is morphological divergence between closely related species exploiting different environments, are less well studied. Here we show divergent evolution in the morphology of the proximal phalanges of several closely related African antelope species inhabiting different environments. This morphological divergence was consistently observed in both a neutral morphospace and an externally ordinated morphospace. Divergence, but not convergence, was also observed when size and shape were considered independently. Finally, convergent evolution of the morphology of the proximal phalanges was observed, but only in the externally ordinated morphospace. Size shows less correlation with phylogeny than does shape. Therefore, we suggest that divergence in size will occur more readily when a species encounters new environmental conditions than divergence in shape. These findings are compatible with observations of rapid dwarfing on islands (Foster’s rule).

Analysis of dental mesowear is a useful tool to examine dietary strategies in ungulates, and provides a palaeoenvironmental proxy at archaeological and palaeontological sites. We examine the distribution of traditional qualitative mesowear variables amongst 33 species of extant African antelope. We also examine the relationship of our dataset with the original dataset of Fortelius and Solounias (2000) . We find that confining mesowear analyses to antelope species alone reduces their discriminating power relative to that found in analyses using a broader range of taxa. Variability in mesowear scores along the toothrow suggest that different teeth do not wear equivalently over an individual’s life-time, so it is important to confine comparisons to individual tooth positions for palaeodietary classifications. Mesowear scores from the third upper molar were found to have the greatest discriminating power. Frugivores displayed unique mesowear signatures, suggesting that this trophic guild should be taken into consideration separately in future mesowear analyses. These results have implications which can improve the application and the discriminatory power of mesowear analyses for the determination of the palaeodiets of extinct taxa, and, by extension, their palaeoenvironments.

Aim To quantify how mammal community structure relates to heterogeneity of vegetation for palaeoecological reconstructions, and to test whether historical or environmental factors are more important in structuring communities. Location Sixty-three natural protected areas in Asia, Africa and South and Central America. Methods We defined faunal communities by allocating species to ecological guilds and calculating proportional representation within each guild. Vegetation heterogeneity for each natural protected area was calculated from satellite images. The relationship between these ecospaces was calculated using canonical correlations analysis, redundancy analysis and principal components analysis. We expected that large, herbivorous mammals would be most strongly correlated with open areas. Convergence was tested by independently eliminating the effects of geography and vegetation heterogeneity on the structure of the mammal communities. We expected that vegetation would more strongly structure communities than geographical position. Results We show that the guild structure of communities across habitats is significantly correlated with vegetation heterogeneity. The highest correlation was between small, scansorial-arboreal secondary consumers and heavy tree cover. The first convergence analysis shows American communities distinguished from Asian and African communities; these latter communities show a remarkable convergence in structure. Historical factors only affected the continent whose mammals had experienced a long period of isolation. The second convergence analysis shows that almost all biomes have the same or very similar community structure regardless of continent. Main conclusions Communities from the same environments in different continents showed remarkable convergence. Communities from the same continents only converged when those continents shared a recent geological and biological history. These results suggest that historical and environmental factors are operating over different timescales. This study confirms that environmental reconstructions made on the basis of whole communities will accurately reflect the environment that the community lived in. However, reconstructions made for fossil sites in deep time need to take historical factors into consideration. Small, arboreal and scansorial secondary consumers show the strongest correlation with vegetation, correlating with continuous tree canopy cover. This relationship allows simple reconstructions of the amount of tree cover occurring in a landscape from the proportion of species from the community falling in this ecological guild.

The habitat preferences of fauna found at palaeontological and archaeological sites can be used to investigate ancient environments and hominin habitat preferences. Here we present a discriminant function model linking astragalus morphology to four broadly defined habitat categories (open, light cover, heavy cover, and closed) using modern bovids of known ecology. Twenty-four measurements were taken on a sample of 286 astragali from 36 extant African antelope species. These measurements were used to generate ratios reflecting shape. An 11 variable discriminant function model was developed that had high classification success rates for complete astragali. Resubstitution analysis, jackknife analysis, and the classification of several “test samples” of specimens suggest that the predictive accuracy of this model is around 87%. The total classification success rates of 87% (jackknifed) or 93% (resubstitution) are considerably higher than those derived in another study of bovid astragalus ecomorphology (67%; [DeGusta, D., Vrba, E.S., 2003. A method for inferring palaeohabitats from the functional morphology of bovid astragali. J. Archaeol. Sci. 30, 1009–1022]) that used a more limited measurement scheme and a smaller sample of bovids than the present study. Different approaches to operationalizing ecomorphic analyses are considered in order to best extract accurate palaeoenvironmental information from palaeontological and archaeological datasets.

Kolpochoerus heseloni is known from the later Pliocene into the middle Pleistocene of Africa. It is widely distributed in time and space, and it is known from many hominin palaeontological and archaeological sites in Africa. We use three methods to investigate the diet and habitat preferences of this species. Dental microwear analysis and carbon stable isotope analysis are used to examine diet, and limb bone ecomorphological analysis is used to reconstruct habitat preference. Results suggest that there is synchronic variability and change through time in the diet and habit preferences of K. heseloni. The results also reinforce how different this taxon is from extant pig species.

One of the distinguishing features of Homo sapiens is its absolutely and relatively large brain. This feature is also seen in less extreme form in some fossil Homo species. However, are increases in brain size during the Plio-Pleistocene only seen in Homo, and is brain enlargement among Plio-Pleistocene primates confined to hominins? This study examines evidence for changes in brain size for species and lineage samples of three synchronic East African fossil primate groups, the two hominin genera Homo and Paranthropus, and the cercopithecoid genus Theropithecus. Hominin endocranial capacity data were taken from the literature, but it was necessary to develop an indirect method for estimating the endocranial volume of Theropithecus. Bivariate and multivariate regression equations relating measured endocranial volume to three external cranial dimensions were developed from a large (ca. 340) sample of modern African cercopithecoids. These equations were used to estimate the endocranial volumes of 20 Theropithecus specimens from the African Plio-Pleistocene. Spearman's rho and the Hubert nonparametric test were used to search for evidence of temporal trends in both the hominin and Theropithecus data. Endocranial volume apparently increased over time in both Homo and Paranthropus boisei, but there was no evidence for temporal trends in the endocranial volume of Theropithecus. Thus, hypotheses which suggest a mix of environmental, social, dietary, or other factors as catalysts for increasing brain in Plio-Pleistocene primates must accommodate evidence of brain enlargement in both Homo and Paranthropus, and explain why this phenomenon appears to be restricted to hominins.

The late Pliocene is notable for the appearance of two new hominid genera as well as the first archaeological sites, generally attributed to the Oldowan Industrial Complex. However, the behavioral ecology of Oldowan hominids has been little explored, particularly at sites older than 2.0 Ma. Moreover, debates on Oldowan hominid foraging ecology and behavior have centered on data from only two regions, and often from single site levels. Here we describe the preliminary results of our investigation of Oldowan occurrences at Kanjera South. These occurrences preserve the oldest known traces of hominid activity in southwestern Kenya, and unlike most of the Oldowan sites in the 2.0-2.5 Ma time interval, artefacts are found in spatial association with a well-preserved fauna. In 1996 and 1997, this project initiated the first excavation program for Kanjera South. Magneto- and biostratigraphy indicate that deposition began approximately 2.2 Ma, substantially earlier than previously thought. At Excavation 1, artefacts were found in spatial association with a taxonomically diverse faunal assemblage in Beds KS-1 and KS-2. Excavation 2 yielded a partial hippopotamus axial skeleton with artefacts in KS-3. Cores from both sites were incidentally flaked and represent a Mode I lithic technology indistinguishable from the Oldowan. Approximately 15% of the artefacts were manufactured from non-local raw materials, indicating a flow of resources into the area. Stable isotopic analysis of KS-1 and KS-2 pedogenic carbonates suggests that the Excavation 1 assemblages formed in a relatively open (>75% C4 grass) habitat. The Excavation 1 and 2 faunas contain a high proportion of equids relative to Oldowan accumulations from Bed I Olduvai Gorge, Tanzania. Beds KS-1 and KS-2 thus preserve traces of Oldowan hominid activities in a more open setting than has been previously documented.

A review of the middle Pleistocene vertebrate faunas of Africa is presented. After a summary of the climate and conditions prevalent during the middle Pleistocene, a regional approach is taken to the evidence, focusing in turn on northern Africa, eastern Africa, and southern Africa. Western Africa is not discussed due to the lack of faunal remains from the region. The changes in geographic range of some species and the composition of the evolving African fauna are emphasized, and modern African terrestrial faunas are shown to be archaic in comparison with those of Eurasia and the Americas.

We examine the relationship between mesowear variables and carbon and nitrogen isotopes in 16 species of African antelope (Mammalia: Bovidae). We show significant differences in carbon and nitrogen isotope values between individuals exhibiting sharp versus round cusps, and high versus low occlusal relief. We show significant correlations between mesowear variables and both carbon and nitrogen isotopes. We find significant correlations between mesowear score and nitrogen, but not carbon isotopes. Finally, we find no significant correlations between hypsodonty index and either isotope examined. Our results provide strong support for the use of mesowear variables in palaeodietary reconstructions of antelopes. Our results further suggest that for the antelopes examined here, mesowear signals are a direct result of diet, while hyposodonty may be the result of phylogenetic legacy.

General ecological theories have paid scant attention to the information preserved in the fossil record. However, in order for an ecological theory to be truly general, it must hold in any ecosystem at any point in time. Here, we make the case that all modern ecological theories should be tested in geological time. We explore some of the limitations of the fossil record when examined in light of modern ecology. While there are fundamental differences between the way modern ecosystems and fossil ones are studied, we demonstrate that comparisons between the two are not impossible. We present three major research areas where fossil information has been successfully used to inform modern ecological thought; namely community ecology, biogeography and extinction studies. These examples also serve to highlight ecological issues that could not have been conceived purely on the basis of modern data. We advocate a much stronger interaction between modern ecologists and paleontologists in addressing present and future ecological questions.

We examined the fossil remains of antelope (Mammalia: Bovidae) postcrania recovered from the Upper Laetolil Beds and the overlying Upper Ndolanya Beds in northern Tanzania. We used analyses of the ecomorphology of extinct antelopes to determine their habitat preferences. The most common postcranial elements - the phalanges, astragali, and distal radii - were examined. A total sample of 446 specimens was analyzed. Changes in the relative proportion of habitat preferences through time are suggestive of temporal fluctuations in habitat availability. As has been noted elsewhere, the mammals from Laetoli do not show the expected distribution of body sizes, so this paper also examined how such size biases, assumed to be taphonomic in origin, might have affected the reconstructions proposed here. We conclude that, on the basis of habitat preferences of the antelopes recovered from Laetoli, there is evidence for the continuous regional presence of woodland and forest throughout deposition of the Upper Laetolil Beds. Antelopes preferring forest and heavy cover habitats dominate the assemblage. Antelopes that locomoted principally in open and light cover habitats are largely in the minority, with a combined frequency never exceeding around 25% of the total in each stratigraphic unit of the Upper Laetolil Beds. The relative proportions of heavy cover and forest preferring antelopes in particular are likely affected by body size biases caused by taphonomic processes active during the deposition of the Upper Laetolil Beds, which favoured the preservation of the postcrania of smaller antelopes. During the formation of the Upper Ndolanya Beds, the proportion of antelopes preferring more open habitats is greatly increased, although forest-preferring antelopes are still present. The conclusion that forest and woodland were always present throughout the sequence is robust.

The manufacture of stone tools and their use to access animal tissues by Pliocene hominins marks the origin of a key adaptation in human evolutionary history. Here we report an in situ archaeological assemblage from the Koobi Fora Formation in northern Kenya that provides a unique combination of faunal remains, some with direct evidence of butchery, and Oldowan artifacts, which are well dated to 1.95 Ma. This site provides the oldest in situ evidence that hominins, predating Homo erectus , enjoyed access to carcasses of terrestrial and aquatic animals that they butchered in a well-watered habitat. It also provides the earliest definitive evidence of the incorporation into the hominin diet of various aquatic animals including turtles, crocodiles, and fish, which are rich sources of specific nutrients needed in human brain growth. The evidence here shows that these critical brain-growth compounds were part of the diets of hominins before the appearance of Homo ergaster/erectus and could have played an important role in the evolution of larger brains in the early history of our lineage.

BACKGROUND: Major biological and cultural innovations in late Pliocene hominin evolution are frequently linked to the spread or fluctuating presence of C(4) grass in African ecosystems. Whereas the deep sea record of global climatic change provides indirect evidence for an increase in C(4) vegetation with a shift towards a cooler, drier and more variable global climatic regime beginning approximately 3 million years ago (Ma), evidence for grassland-dominated ecosystems in continental Africa and hominin activities within such ecosystems have been lacking. METHODOLOGY/PRINCIPAL FINDINGS: We report stable isotopic analyses of pedogenic carbonates and ungulate enamel, as well as faunal data from approximately 2.0 Ma archeological occurrences at Kanjera South, Kenya. These document repeated hominin activities within a grassland-dominated ecosystem. CONCLUSIONS/SIGNIFICANCE: These data demonstrate what hitherto had been speculated based on indirect evidence: that grassland-dominated ecosystems did in fact exist during the Plio-Pleistocene, and that early Homo was active in open settings. Comparison with other Oldowan occurrences indicates that by 2.0 Ma hominins, almost certainly of the genus Homo, used a broad spectrum of habitats in East Africa, from open grassland to riparian forest. This strongly contrasts with the habitat usage of Australopithecus, and may signal an important shift in hominin landscape usage.

The earliest archaeological traces and two new hominin genera (Homo and Paranthropus) appear in the late Pliocene of Africa. These first appearances may reflect novel hominin adaptive responses to shifting resource bases over geological time and/or an increasingly seasonal distribution of food over the annual cycle. Whereas regional environmental change has been documented during the Plio-Pleistocene of East Africa, it is difficult to resolve relative proportions of specific habitats at a given place and time, how these may have changed over time, and the explicit nature of particular habitats. Detailed reconstructions of paleohabitats based on paleontological, geological and geochemical evidence are necessary in order to better understand the interplay between environmental change and hominin biological and behavioral evolution.

This contribution discusses suid specimens from the Manonga Valley, Tanzania. The 1990, 1992, and 1994 expeditions to the Wembere-Manonga Valley recovered 65 suid specimens from ten sites (Table I). Fossils suids derive from horizons throughout the Wembere-Manonga Formation, although the majority were recovered from the Ibole Member (Harrison 1991, 1993, 1994; Harrison and Verniers, 1993; Harrison et al., 1993; Haileab and Harrison, 1993; Mbago et al., 1993; Verniers, this volume, Chapter 2). There were no differences between specimens from the different members so they are treated together here. All suid remains can be attributed to the Pliocene species Nyanzachoerus kanamensis.

Bone morphology of the cats (Mammalia: Felidae) is influenced by many factors, including locomotor mode, body size, hunting methods, prey size and phylogeny. Here, we investigate the shape of the proximal and distal humeral epiphyses in extant species of the felids, based on two-dimensional landmark configurations. Geometric morphometric techniques were used to describe shape differences in the context of phylogeny, allometry and locomotion. The influence of these factors on epiphyseal shape was assessed using Principal Component Analysis, Linear Discriminant functions and multivariate regression. Phylogenetic Generalised Least Squares was used to examine the association between size or locomotion and humeral epiphyseal shape, after taking a phylogenetic error term into account. Results show marked differences in epiphyseal shape between felid lineages, with a relatively large phylogenetic influence. Additionally, the adaptive influences of size and locomotion are demonstrated, and their influence is independent of phylogeny in most, but not all, cases. Several features of epiphyseal shape are common to the largest terrestrial felids, including a relative reduction in the surface area of the humeral head and increased robusticity of structures that provide attachment for joint-stabilising muscles, including the medial epicondyle and the greater and lesser tubercles. This increased robusticity is a functional response to the increased loading forces placed on the joints due to large body mass.

The Shungura Formation in the lower Omo River Valley, southern Ethiopia, has yielded an important paleontological and archeological record from the Pliocene and Pleistocene of eastern Africa. Fossils are common throughout the sequence and provide evidence of paleoenvironments and environmental change through time. This study developed discriminant function ecomorphology models that linked astragalus morphology to broadly defined habitat categories (open, light cover, heavy cover, forest, and wetlands) using modern bovids of known ecology. These models used seven variables suitable for use on fragmentary fossils and had overall classification success rates of >82%. Four hundred and one fossils were analyzed from Shungura Formation members B through G (3.4-1.9 million years ago). Analysis by member documented the full range of ecomorph categories, demonstrating that a wide range of habitats existed along the axis of the paleo-Omo River. Heavy cover ecomorphs, reflecting habitats such as woodland and heavy bushland, were the most common in the fossil sample. The trend of increasing open cover habitats from Members C through F suggested by other paleoenvironmental proxies was documented by the increase in open habitat ecomorphs during this interval. However, finer grained analysis demonstrated considerable variability in ecomorph frequencies over time, suggesting that substantial short-term variability is masked when grouping samples by member. The hominin genera Australopithecus, Homo, and Paranthropus are associated with a range of ecomorphs, indicating that all three genera were living in temporally variable and heterogeneous landscapes. Australopithecus finds were predominantly associated with lower frequencies of open habitat ecomorphs, and high frequencies of heavy cover ecomorphs, perhaps indicating a more woodland focus for this genus.