Homo heterogenus: Variability in early Pleistocene Homo environments.

To understand the ecological dominance of Homo sapiens, we need to investigate the origins of the plasticity that has enabled our colonization of the planet. We can approach this by exploring the variability of habitats to which different hominin populations have adapted over time. In this article, we draw upon and synthesize the current research on habitats of genus Homo during the early Pleistocene. We examined 121 published environmental reconstructions from 74 early Pleistocene sites or site phases to assess the balance of arguments in the research community. We found that, while grasslands and savannahs were prominent features of Homo habitats in the early Pleistocene, current research does not place early Pleistocene Homo, in any single environmental type, but in a wide variety of environments, ranging from open grasslands to forests. Our analysis also suggests that the first known dispersal of Homo out of Africa was accompanied by niche expansion.

Do species factories exist? Detecting exceptional patterns of evolution in the mammalian fossil record.

A species factory refers to the source that gives rise to an exceptionally large number of species. However, what is it exactly: a place, a time or a combination of places, times and environmental conditions, remains unclear. Here we search for species factories computationally, for which we develop statistical approaches to detect origination, extinction and sorting hotspots in space and time in the fossil record. Using data on European Late Cenozoic mammals, we analyse where, how and how often species factories occur, and how they potentially relate to the dynamics of environmental conditions. We find that in the Early Miocene origination hotspots tend to be located in areas with relatively low estimated net primary productivity. Our pilot study shows that species first occurring in origination hotspots tend to have a longer average longevity and a larger geographical range than other species, thus emphasizing the evolutionary importance of the species factories.

Taphonomic and spatial analyses from the Early Pleistocene site of Venta Micena 4 (Orce, Guadix-Baza Basin, southern Spain).

Venta Micena is an area containing several palaeontological sites marking the beginning of the Calabrian stage (Early Pleistocene). The richness of the fossil accumulation including species of Asian, African and European origin, makes Venta Micena a key site for the the palaeoecological and palaeoenvironmental study of southern Europe during the Early Pleistocene. Thus, research has been focused on Venta Micena 3, which was originally interpreted as a single palaeosurface associated with a marshy context, in which most of the fauna was accumulated by Pachycrocuta brevirostris. Recent excavations have unearthed a new site, Venta Micena 4, located in the same stratigraphic unit (Unit C) and in close proximity to Venta Micena 3. Here we show the first analyses regarding the taphonomic and spatial nature of this new site, defining two stratigraphic boundaries corresponding to two different depositional events. Furthermore, the taphonomic analyses of fossil remains seem to indicate a different accumulative agent than Pachycrocuta, thus adding more complexity to the palaeobiological interpretation of the Venta Micena area. These results contribute to the discussion of traditional interpretations made from Venta Micena 3.

The case of the grass-eating suids in the Plio-Pleistocene Turkana Basin: 3D dental topography in relation to diet in extant and fossil pigs.

Two separate subfamilies of Plio-Pleistocene African pigs (suids) consecutively evolved hypsodont and horizodont molars with flat occlusal surfaces, commonly interpreted as an adaptive trait to a grazing diet, similar to that of the present warthogs (Phacochoerus spp.). To investigate this in detail, we studied the 3D-dental topography of fossil specimens from the Turkana Basin, using geographic information systems-based methods. To establish baselines for interpretation of the Turkana Basin suids, topography of third molars of extant suids with known diets were analyzed: grazing warthog (Phacochoerus africanus), herbivorous mixed-feeder forest hog (Hylochoerus meinertzhageni), omnivorous generalist wild boar (Sus scrofa), omnivorous fruit and tuber eater bush pig (Potamochoerus spp.), and omnivorous fruit eater babirusa (Babyrousa spp.) In addition, we analyzed supposedly browsing Miocene suids, Listriodon spp. The same topographic measures were applied to Plio-Pleistocene specimens from the Turkana Basin, Kenya: Notochoerus euilus, Notochoerus scotti, Kolpochoerus heseloni, and Metridiochoerus andrewsi. With some differences between techniques, 3D-dental topography analysis of extant suid molars mostly predicts the dietary differences between the species correctly. The grazing P. africanus differs from both the omnivorous suids and the herbivorous mixed-feeder H. meinertzhageni in all except one metrics. The omnivorous mostly tropical suids, Potamochoerus and Babyrousa, primarily differ from the generalist, S. scrofa, in the orientation patch count analysis, showing higher occlusal complexity in the latter. Although, there might be significant gaps between the morphological changes and the ecological changes, we conclude that based on comparison of dental topography with the present-day suids, N. scotti and M. andrewsi were most likely highly specialized grazers, while N. euilus and K. heseloni retained more of their ancestral, omnivorous heritage, but consumed grasses more than the extant omnivorous suids. RESEARCH HIGHLIGHTS: Dental topography can predict different diets in present-day wild pigs. The Plio-Pleistocene pigs in the Turkana Basin had dental topography mostly similar to extant grazing warthog, although some species also had resemblances to omnivorous forest pigs.

The phylogenetic signal in tooth wear: What does it mean?

A new study by Fraser et al (2018) urges the use of phylogenetic comparative methods, whenever possible, in analyses of mammalian tooth wear. We are concerned about this for two reasons. First, this recommendation may mislead the research community into thinking that phylogenetic signal is an artifact of some sort rather than a fundamental outcome of the evolutionary process. Secondly, this recommendation may set a precedent for editors and reviewers to enforce phylogenetic adjustment where it may unnecessarily weaken or even directionally alter the results, shifting the emphasis of analysis from common patterns manifested by large clades to rare cases.

The rise and fall of the Old World savannah fauna and the origins of the African savannah biome.

Despite much interest in the ecology and origins of the extensive grassland ecosystems of the modern world, the biogeographic relationships of savannah palaeobiomes of Africa, India and mainland Eurasia have remained unclear. Here we assemble the most recent data from the Neogene mammal fossil record in order to map the biogeographic development of Old World mammalian faunas in relation to palaeoenvironmental conditions. Using genus-level faunal similarity and mean ordinated hypsodonty in combination with palaeoclimate modelling, we show that savannah faunas developed as a spatially and temporally connected entity that we term the Old World savannah palaeobiome. The Old World savannah palaeobiome flourished under the influence of middle and late Miocene global cooling and aridification, which resulted in the spread of open habitats across vast continental areas. This extensive biome fragmented into Eurasian and African branches due to increased aridification in North Africa and Arabia during the late Miocene. Its Eurasian branches had mostly disappeared by the end of the Miocene, but the African branch survived and eventually contributed to the development of Plio-Pleistocene African savannah faunas, including their early hominins. The modern African savannah fauna is thus a continuation of the extensive Old World savannah palaeobiome.

Author Correction: The rise and fall of the Old World savannah fauna and the origins of the African savannah biome.

In the version of this Article originally published, each of the five panels in Fig. 5 incorrectly contained a black diagonal line across the plot. This has now been corrected.

Paleoecology of the Serengeti during the Oldowan-Acheulean transition at Olduvai Gorge, Tanzania: The mammal and fish evidence.

Eight years of excavation work by the Olduvai Geochronology and Archaeology Project (OGAP) has produced a rich vertebrate fauna from several sites within Bed II, Olduvai Gorge, Tanzania. Study of these as well as recently re-organized collections from Mary Leakey's 1972 HWK EE excavations here provides a synthetic view of the faunal community of Olduvai during Middle Bed II at ∼1.7-1.4 Ma, an interval that captures the local transition from Oldowan to Acheulean technology. We expand the faunal list for this interval, name a new bovid species, clarify the evolution of several mammalian lineages, and record new local first and last appearances. Compositions of the fish and large mammal assemblages support previous indications for the dominance of open and seasonal grassland habitats at the margins of an alkaline lake. Fish diversity is low and dominated by cichlids, which indicates strongly saline conditions. The taphonomy of the fish assemblages supports reconstructions of fluctuating lake levels with mass die-offs in evaporating pools. The mammals are dominated by grazing bovids and equids. Habitats remained consistently dry and open throughout the entire Bed II sequence, with no major turnover or paleoecological changes taking place. Rather, wooded and wet habitats had already given way to drier and more open habitats by the top of Bed I, at 1.85-1.80 Ma. This ecological change is close to the age of the Oldowan-Acheulean transition in Kenya and Ethiopia, but precedes the local transition in Middle Bed II. The Middle Bed II large mammal community is much richer in species and includes a much larger number of large-bodied species (>300 kg) than the modern Serengeti. This reflects the severity of Pleistocene extinctions on African large mammals, with the loss of large species fitting a pattern typical of defaunation or 'downsizing' by human disturbance. However, trophic network (food web) analyses show that the Middle Bed II community was robust, and comparisons with the Serengeti community indicate that the fundamental structure of food webs remained intact despite Pleistocene extinctions. The presence of a generalized meat-eating hominin in the Middle Bed II community would have increased competition among carnivores and vulnerability among herbivores, but the high generality and interconnectedness of the Middle Bed II food web suggests this community was buffered against extinctions caused by trophic interactions.

Dental functional morphology predicts the scaling of chewing rate in mammals.

How food intake and mastication scale to satisfy the metabolic needs of mammals has been the subject of considerable scientific debate. Existing theory suggests that the negative allometric scaling of metabolic rate with body mass is compensated by a matching allometric scaling of the chewing rate. Why empirical studies have found that the scaling coefficients of the chewing rate seem to be systematically smaller than expected from theory remains unknown. Here we explain this imparity by decoupling the functional surface area of teeth from overall surface area. The functional surface area is relatively reduced in forms emphasizing linear edges (e.g., lophodont) compared with forms lacking linear structures (e.g., bunodont). In forms with reduced relative functional surface, the deficit in food processed per chew appears to be compensated for by increased chewing rate, such that the metabolic requirements are met. This compensation accounts for the apparent difference between theoretically predicted and observed scaling of chewing rates. We suggest that this reflects adaptive functional evolution to plant foods with different fracture properties and extend the theory to incorporate differences in functional morphology.

Reconciling taxon senescence with the Red Queen's hypothesis.

In the fossil record, taxa exhibit a regular pattern of waxing and waning of occupancy, range or diversity between their origin and extinction. This pattern appears to contradict the law of constant extinction, which states that the probability of extinction in a given taxon is independent of that taxon's age. It is nevertheless well established for species, genera and higher taxa of terrestrial mammals, marine invertebrates, marine microorganisms, and recent Hawaiian clades of animals and plants. Here we show that the apparent contradiction between a stochastically constant extinction rate and the seemingly deterministic waxing and waning pattern of taxa disappears when we consider their peak of expansion rather than their final extinction. To a first approximation, we find that biotic drivers of evolution pertain mainly to the peak of taxon expansion, whereas abiotic drivers mainly apply to taxon extinction. The Red Queen's hypothesis, which emphasizes biotic interactions, was originally proposed as an explanation of the law of constant extinction. Much effort has since been devoted to determining how this hypothesis, emphasizing competition for resources, relates to the effects of environmental change. One proposed resolution is that biotic and abiotic processes operate at different scales. By focusing attention on taxon expansion rather than survival, we resolve an apparent contradiction between the seemingly deterministic waxing and waning patterns over time and the randomness of extinction that the Red Queen's hypothesis implies.

The palaeoenvironment of the middle Miocene pliopithecid locality in Damiao, Inner Mongolia, China.

Damiao, Inner Mongolia, has three main fossil horizons representing the early, middle, and late Miocene. The middle Miocene locality DM01 is the only primate locality from the region and also represents the latest occurrence of pliopithecoids in northern China. The presence of pliopithecoid primates in central Asia after the middle Miocene climatic optimum seems to contradict the general trend of strengthening climatic zonality and increasing aridity. To investigate this enigma, we employ faunal similarity, ecometrics, and stable isotope analysis. Our results support previous inferences concerning the presence of locally humid environments within the increasingly arid surroundings that characterized central Asia. Hypsodonty, estimated mean annual precipitation (MAP), local sedimentology, and large mammal fossils suggest more humid and possibly more forested and wooded environments for the DM01 locality. We compared our results with the adjacent fossil-rich middle Miocene Tunggur localities. However, the small mammal fauna and isotope data are consistent with a mosaic of forest and grassland environment for all Damiao localities. Based on our results, Tunggur may have been too seasonal or not sufficiently humid for pliopithecids. This is supported by the higher mean hypsodonty and lower estimated MAP estimates, as well as slightly higher δ13C values. We suggest that DM01, the driest known Asian pliopithecid locality, may have been a more humid refugium within a generally drier regional context.

Merging paleobiology with conservation biology to guide the future of terrestrial ecosystems.

Conservation of species and ecosystems is increasingly difficult because anthropogenic impacts are pervasive and accelerating. Under this rapid global change, maximizing conservation success requires a paradigm shift from maintaining ecosystems in idealized past states toward facilitating their adaptive and functional capacities, even as species ebb and flow individually. Developing effective strategies under this new paradigm will require deeper understanding of the long-term dynamics that govern ecosystem persistence and reconciliation of conflicts among approaches to conserving historical versus novel ecosystems. Integrating emerging information from conservation biology, paleobiology, and the Earth sciences is an important step forward on the path to success. Maintaining nature in all its aspects will also entail immediately addressing the overarching threats of growing human population, overconsumption, pollution, and climate change.

Herbivore teeth predict climatic limits in Kenyan ecosystems.

A major focus in evolutionary biology is to understand how the evolution of organisms relates to changes in their physical environment. In the terrestrial realm, the interrelationships among climate, vegetation, and herbivores lie at the heart of this question. Here we introduce and test a scoring scheme for functional traits present on the worn surfaces of large mammalian herbivore teeth to capture their relationship to environmental conditions. We modeled local precipitation, temperature, primary productivity, and vegetation index as functions of dental traits of large mammal species in 13 national parks in Kenya over the past 60 y. We found that these dental traits can accurately estimate local climate and environment, even at small spatial scales within areas of relatively uniform climate (within two ecoregions), and that they predict limiting conditions better than average conditions. These findings demonstrate that the evolution of key functional properties of organisms may be more reflective of demands during recurring adverse episodes than under average conditions or during isolated severe events.

Mechanical modelling of tooth wear.

Different diets wear teeth in different ways and generate distinguishable wear and microwear patterns that have long been the basis of palaeodiet reconstructions. Little experimental research has been performed to study them together. Here, we show that an artificial mechanical masticator, a chewing machine, occluding real horse teeth in continuous simulated chewing (of 100 000 chewing cycles) is capable of replicating microscopic wear features and gross wear on teeth that resemble wear in specimens collected from nature. Simulating pure attrition (chewing without food) and four plant material diets of different abrasives content (at n = 5 tooth pairs per group), we detected differences in microscopic wear features by stereomicroscopy of the chewing surface in the number and quality of pits and scratches that were not always as expected. Using computed tomography scanning in one tooth per diet, absolute wear was quantified as the mean height change after the simulated chewing. Absolute wear increased with diet abrasiveness, originating from phytoliths and grit. In combination, our findings highlight that differences in actual dental tissue loss can occur at similar microwear patterns, cautioning against a direct transformation of microwear results into predictions about diet or tooth wear rate.

An ecometric analysis of the fossil mammal record of the Turkana Basin.

Although ecometric methods have been used to analyse fossil mammal faunas and environments of Eurasia and North America, such methods have not yet been applied to the rich fossil mammal record of eastern Africa. Here we report results from analysis of a combined dataset spanning east and west Turkana from Kenya between 7 and 1 million years ago (Ma). We provide temporally and spatially resolved estimates of temperature and precipitation and discuss their relationship to patterns of faunal change, and propose a new hypothesis to explain the lack of a temperature trend. We suggest that the regionally arid Turkana Basin may between 4 and 2 Ma have acted as a 'species factory', generating ecological adaptations in advance of the global trend. We show a persistent difference between the eastern and western sides of the Turkana Basin and suggest that the wetlands of the shallow eastern side could have provided additional humidity to the terrestrial ecosystems. Pending further research, a transient episode of faunal change centred at the time of the KBS Member (1.87-1.53 Ma), may be equally plausibly attributed to climate change or to a top-down ecological cascade initiated by the entry of technologically sophisticated humans.This article is part of the themed issue 'Major transitions in human evolution'.

Cope's Rule and the Universal Scaling Law of Ornament Complexity.

Luxuriant, bushy antlers, bizarre crests, and huge, twisting horns and tusks are conventionally understood as products of sexual selection. This view stems from both direct observation and from the empirical finding that the size of these structures grows faster than body size (i.e., ornament size shows positive allometry). We contend that the familiar evolutionary increase in the complexity of ornaments over time in many animal clades is decoupled from ornament size evolution. Increased body size comes with extended growth. Since growth scales to the quarter power of body size, we predicted that ornament complexity should scale according to the quarter power law as well, irrespective of the role of sexual selection in the evolution and function of the ornament. To test this hypothesis, we selected three clades (ammonites, deer, and ceratopsian dinosaurs) whose species bore ornaments that differ in terms of the importance of sexual selection to their evolution. We found that the exponent of the regression of ornament complexity to body size is the same for the three groups and is statistically indistinguishable from 0.25. We suggest that the evolution of ornament complexity is a by-product of Cope's rule. We argue that although sexual selection may control size in most ornaments, it does not influence their shape.

Modeling the Population-Level Processes of Biodiversity Gain and Loss at Geological Timescales.

The path of species diversification is commonly observed by inspecting the fossil record. Yet, how species diversity changes at geological timescales relate to lower-level processes remains poorly understood. Here we use mathematical models of spatially structured populations to show that natural selection and gradual environmental change give rise to discontinuous phenotype changes that can be connected to speciation and extinction at the macroevolutionary level. In our model, new phenotypes arise in the middle of the environmental gradient, while newly appearing environments are filled by existing phenotypes shifting their adaptive optima. Slow environmental change leads to loss of phenotypes in the middle of the extant environmental range, whereas fast change causes extinction at one extreme of the environmental range. We compared our model predictions against a well-known yet partially unexplained pattern of intense hoofed mammal diversification associated with grassland expansion during the Late Miocene. We additionally used the model outcomes to cast new insight into Cope's law of the unspecialized. Our general finding is that the rate of environmental change determines where generation and loss of diversity occur in the phenotypic and physical spaces.

Co-occurrence of pliopithecoid and hominoid primates in the fossil record: An ecometric analysis.

Both pliopithecoid and hominoid primates were widely distributed throughout Eurasia during the Miocene but are known to have coexisted at only a few localities. It has been speculated that their different habitat preferences permitted only minimal overlap under special environmental conditions. Here we study the context for pliopithecoid and hominoid co-occurrence by assessing taxonomically-based palaeoecological diversity of associated fossil mammals and by direct ecometric analysis based on hypsodonty of mammalian herbivores. Our results show that pliopithecoids persistently inhabited more humid environments compared to the other primate groups studied, suggesting an inability to adapt to changing environmental conditions. The opportunity for hominoids and pliopithecoids to co-occur appears to have been restricted by niche conservatism in the latter group. Our study also indicates that direct ecometric analysis gives a better separation of the ecological preferences of these primate clades than do analyses of taxonomically-based community structure.