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.

Vrba was right: Historical climate fragmentation, and not current climate, explains mammal biogeography.

Climate plays a crucial role in shaping species distribution and evolution over time. Dr Vrba's Resource-Use hypothesis posited that zones at the extremes of temperature and precipitation conditions should host a greater number of climate specialist species than other zones because of higher historical fragmentation. Here, we tested this hypothesis by examining climate-induced fragmentation over the past 5 million years. Our findings revealed that, as stated by Vrba, the number of climate specialist species increases with historical regional climate fragmentation, whereas climate generalist species richness decreases. This relationship is approximately 40% stronger than the correlation between current climate and species richness for climate specialist species and 77% stronger for generalist species. These evidences suggest that the effect of climate historical fragmentation is more significant than that of current climate conditions in explaining mammal biogeography. These results provide empirical support for the role of historical climate fragmentation and physiography in shaping the distribution and evolution of life on Earth.

Diversification and evolutionary history of the African laminated-toothed rats (Rodentia, Otomyini).

The African continent was subjected to periodic climatic shifts during the Pliocene and Pleistocene. These habitat changes greatly affected the evolutionary processes and tempo of diversification in numerous, widely distributed mammals. The Otomyini (Family Muridae) comprises three African rodent genera, Parotomys, Otomys and Myotomys, characterized by unique laminated-shaped molars. Species within this tribe generally prefer open-habitat and show low dispersal capabilities, with previous studies suggesting that their diversification was closely associated with climatic oscillations over the last four million years. Our phylogenetic reconstructions, based on three mitochondrial (mtDNA) genes (Cytb, COI and 12S) and four nuclear introns (EF, SPTBN, MGF and THY), identified eight major genetic clades that are distributed across southern, eastern and western Africa. Our data permit the re-examination of the taxonomic status of the three genera as well as the previously proposed mesic-arid dichotomy of the 10 South African species. Moreover, multiple mtDNA species delimitation methods incorporating 168 specimens estimated the number of Otomyini species to be substantially higher than the âˆ¼ 30 recognized, suggesting that the current taxonomy will necessitate an integrative approach to delimit extant species diversity within the Otomyini. The data suggests that the origin of the tribe can be dated back to âˆ¼ 5.7 million years ago (Ma) in southern Africa. The distribution and phylogenetic associations among the eight major otomyine evolutionary lineages can best be explained by several waves of northward colonization from southern Africa, complemented by independent reversed dispersals from eastern back to southern Africa at different time periods. There is strong support for the hypothesis that the radiation, dispersion, and diversification of the otomyine rodents is closely linked to recent Plio-Pleistocene climatic oscillations.

The anatomy of the hindlimb of Theropithecus brumpti (Cercopithecidae, Papionini): Morphofunctional implications.

Theropithecus brumpti is a primate known from numerous craniodental specimens in the Plio-Pleistocene Shungura Formation (Lower Omo Valley, Ethiopia), but the anatomy of its hindlimb is documented only by a few associated and mostly incomplete postcranial specimens. The adaptations of T. brumpti are still debated, with its substrate preferences and its use of squatting postures recently discussed based on anatomical differences when compared with its extant representative, Theropithecus gelada. Here, we describe an associated femur and tibia (L 869-1 and L 869-2) of a presumed T. brumpti male and a partial foot (L 865-1r and L 865-1t) of a male T. brumpti, dated to ca. 2.6 Ma and ca. 2.32 Ma respectively. Based on univariate and bivariate morphometric analyses, we provide new data on the morphological correlates of substrate preferences and postural behaviors of this fossil species. Our results are in agreement with previous analyses and present T. brumpti as a predominantly terrestrial primate. We demonstrate the presence of osteological correlates associated with the use of squatting behaviors in T. brumpti but also point to significant anatomical differences between this paleontological species and T. gelada. These differences blur the functional value of characters previously identified as diagnostic of T. gelada and its postural behavior. We further document the postcranial distinctiveness of the Theropithecus clade in relation to the Papio clade. This study thus provides new insights into the postcranial anatomy and paleoecology of an abundant fossil primate from the Plio-Pleistocene of eastern Africa.

The Australopithecus assemblage from Sterkfontein Member 4 (South Africa) and the concept of variation in palaeontology.

Interpreting morphological variation within the early hominin fossil record is particularly challenging. Apart from the fact that there is no absolute threshold for defining species boundaries in palaeontology, the degree of variation related to sexual dimorphism, temporal depth, geographic variation or ontogeny is difficult to appreciate in a fossil taxon mainly represented by fragmentary specimens, and such variation could easily be conflated with taxonomic diversity. One of the most emblematic examples in paleoanthropology is the Australopithecus assemblage from the Sterkfontein Caves in South Africa. Whereas some studies support the presence of multiple Australopithecus species at Sterkfontein, others explore alternative hypotheses to explain the morphological variation within the hominin assemblage. In this review, I briefly summarize the ongoing debates surrounding the interpretation of morphological variation at Sterkfontein Member 4 before exploring two promising avenues that would deserve specific attention in the future, that is, temporal depth and nonhuman primate diversity.

Recurrent founder-event speciation across the Mediterranean likely shaped the species diversity and geographic distribution of the freshwater snail genus Mercuria Boeters, 1971 (Caenogastropoda: Hydrobiidae).

Dispersal is known to play an important role in shaping the diversity and geographic range of freshwater gastropods. Here, we used phylogenetic methods to test for the influence of dispersal and other biogeographic processes (such as vicariance) on the speciation and distribution patterns of Mercuria Boeters, 1971, a snail genus widely distributed in the western Palaearctic. The 25 extant species traditionally thought to comprise the genus, which were described mainly on the basis of morphology, have been recorded from lowland waters in both the Mediterranean and Atlantic river basins of Europe and North Africa. Using molecular phylogenies based on three gene fragments (COI, 16S rRNA and 28S rRNA) from 209 individuals, four molecular species delimitation methods and a shell characterization, we identified 14 putative species in our dataset, nine of which correspond to species classified by traditional taxonomy. Furthermore, biogeographical modelling favoured a scenario in which recurrent founder-event speciation since the late Miocene is the most probable process explaining the species diversity and distribution of the Mediterranean clades, whereas episodes of postglacial northward colonization from Iberian refugia by the species M.tachoensis may explain the current presence of the genus in Atlantic lowlands. The dispersal events inferred for Mercuria, probably promoted by multiple factors such as the changing connectivity of drainage basins driven by climate change or better access for avian dispersal vectors in lowlands, may explain the rare case among hydrobiids of a species-rich genus containing individual species with a large distribution area.

Do rates of dental wear in extant African great apes inform the time of weaning?

Reconstructing the life histories of extinct hominins remains one of the main foci of paleoanthropological inquiry, as an extended juvenile period impacts the social and cognitive development of species. However, the paucity of hominin remains, the lack of comparative hominoid data, and the destructive nature of many life history approaches have limited our understanding of the relationship between dental development (eruption) and weaning in primates. Alternatively, the rate of dental wear in early-forming teeth has been suggested a good proxy for the timing of weaning. Here we test this hypothesis on an ontogenetic series of Gorilla gorilla gorilla and Pan troglodytes troglodytes, using geographic information systems-based shape descriptors of M1s in relation to the nitrogen (δ15N) and carbon (δ13C) isotope composition of their associated hair. Results show that Gorilla g. gorilla are fully weaned considerably later than Pan t. troglodytes, that is, after M1s had been in full functional occlusion for some time. Yet, throughout ontogeny, gorilla dental wear rates are greater than they are in chimpanzees. This refutes the hypothesis that the rates of wear of early-forming teeth inform the time of weaning (i.e., nutritional independence). Instead, dietary breadth and seasonal variation in resource availability are implicated. This finding has implications for interpreting the hominin fossil record and raises questions about the triggers for, and the mechanisms of, life history change in hominin evolution. As a case in point, commonalities in life history patterns between early hominins and Western lowland gorillas seem to be a means to mitigate the effects of recurrent (i.e., seasonal) resource limitations and-conceivably-to prevent high infant mortality rates. Taken further, difference between hominid life histories are likely to be of degree, not kind.

Feeding ecology of the last European colobine monkey, Dolichopithecus ruscinensis.

Currently, very little is known about the ecology of extinct Eurasian cercopithecids. Dietary information is crucial in understanding the ecological adaptations and diversity of extinct cercopithecids and the evolution of this family. For example, the colobine genus Dolichopithecus is represented by multiple large-bodied species that inhabited Eurasia during the Pliocene-Early Pleistocene. The available evidence, though limited, suggests semiterrestrial locomotion, which contrasts with most extant African and Asian colobines that exhibit morphological and physiological adaptations for arboreality and folivory. These differences raise questions regarding the dietary specialization of early colobine taxa and how/if that influenced their dispersion out of Africa and into Eurasia. To further our understanding of the ecology of Plio-Pleistocene cercopithecids, we characterized the dental capabilities and potential dietary adaptations of Dolichopithecus ruscinensis through dental topographic and enamel thickness analyses on an M1 from the locality of Serrat d'en Vacquer, Perpignan (France). We also assessed the feeding behavior of D. ruscinensis through dental microwear texture analysis on a broad sample of fossil molars from fossil sites in France, Greece, Bulgaria, and Romania. Dental topographic and enamel thickness analyses suggest that D. ruscinensis could efficiently process a wide range of foods. Results of the dental microwear texture analysis suggest that its diet ranged from folivory to the consumption of more mechanically challenging foods. Collectively, this suggests a more opportunistic feeding behavior for Dolichopithecus than characteristic of most extant colobines.

The enigmatic Leiosaurae clade: Phylogeography, species delimitation, phylogeny and historical biogeography of its southernmost species.

The clade Leiosaurae is composed of poorly-known species endemic to the southern region of South America. The difficulties of finding these lizards in the field, and their highly conserved morphology, have limited our taxonomic knowledge and understanding of their evolutionary histories. Here, we use data collected over 9 years to study the phylogenetic history, genetic diversity, and biogeographic history of almost all the southernmost species of Leiosaurae (except P. nigroigulus), including: Leiosaurus bellii, Diplolaemus darwinii, D. bibronii, D. sexcinctus and D. leopardinus. We use a fragment of the mitochondrial cytochrome-b gene to resolve general phylogeographic patterns, and add another mitochondrial gene and eight nuclear genes to perform species delimitation and phylogenetic analyses associated with divergence times. We found evidence for three putative new species-level taxa within L. bellii and five within Diplolaemus species, indicating high levels of geographic structure. We use a time-calibrated phylogeny to estimate ranges of ancestral distributions and to generate new hypotheses about their historical biogeography.

Evolution of Sri Lanka's Giant Danios (Teleostei: Cyprinidae: Devario): Teasing apart species in a recent diversification.

The small, colourful freshwater fishes of the cyprinid genus Devario are among the many vertebrate groups that appear to have diversified on Sri Lanka, a continental Indian Ocean island, which is part of the Western Ghats-Sri Lanka Biodiversity Hotspot. Despite Sri Lanka having been connected with India via a wide isthmus intermittently until the Plio-Pleistocene and almost continuously since then, during sea-level low-stands, the number of species of Devario on Sri Lanka is comparable with that on the Indian Peninsula, some 25 times its size. Here, from a sampling of 27 Devario populations across Sri Lanka's major river basins and climatic zones, we present and analyze a phylogeny based on mitochondrial and nuclear DNA data. We show that five species of Devario occur on the island, all but one of which are endemic: Devario malabaricus is widespread throughout the lowlands and parts of central hills of Sri Lanka and southern India. A new narrow-range endemic, here described as D. memorialis sp. nov., was discovered in a remnant rainforest habitat at Aranayake (Ma Oya basin) in this study. It is immediately distinguished from Sri Lankan congeners by having only 8 (vs 9-12) branched dorsal-fin rays and by uncorrected pairwise genetic distances of more than 4.0% and 7.8% for cytochrome c oxidase subunit 1 and cytochrome b, respectively. Our results provide strong support for the monophyly of the entire Sri Lankan diversification of Devario. Divergence-timing analysis suggests the common ancestor dispersed to the island in the late Miocene, with the insular diversification in the island's south-western wet zone taking place during the Plio-pleistocene. There are signs of gene flow between the Indian and Sri Lankan populations of D. malabaricus until the late Pleistocene. Phylogenetic and haplotype-network analyses suggest basin-centric phylogeographic structure within the endemic species; D. malabaricus, however, shows little such structure in the island. Molecular and morphological analyses failed to identify D. annnataliae and D. udenii confidently as distinct species: they are considered synonyms of D. micronema. The morphological variation observed within D. micronema is likely attributable to polymorphism. The discordance between the mitochondrial and nuclear phylogenies for some samples of Devario in the present study suggest signs of mitochondrial introgression.

Multiple radiations of spiny mice (Rodentia: Acomys) in dry open habitats of Afro-Arabia: evidence from a multi-locus phylogeny.

BACKGROUND: Spiny mice of the genus Acomys are distributed mainly in dry open habitats in Africa and the Middle East, and they are widely used as model taxa for various biological disciplines (e.g. ecology, physiology and evolutionary biology). Despite their importance, large distribution and abundance in local communities, the phylogeny and the species limits in the genus are poorly resolved, and this is especially true for sub-Saharan taxa. The main aims of this study are (1) to reconstruct phylogenetic relationships of Acomys based on the largest available multilocus dataset (700 genotyped individuals from 282 localities), (2) to identify the main biogeographical divides in the distribution of Acomys diversity in dry open habitats in Afro-Arabia, (3) to reconstruct the historical biogeography of the genus, and finally (4) to estimate the species richness of the genus by application of the phylogenetic species concept. RESULTS: The multilocus phylogeny based on four genetic markers shows presence of five major groups of Acomys called here subspinosus, spinosissimus, russatus, wilsoni and cahirinus groups. Three of these major groups (spinosissimus, wilsoni and cahirinus) are further sub-structured to phylogenetic lineages with predominantly parapatric distributions. Combination of alternative species delimitation methods suggests the existence of 26 molecular operational taxonomic units (MOTUs), potentially corresponding to separate species. The highest genetic diversity was found in Eastern Africa. The origin of the genus Acomys is dated to late Miocene (ca. 8.7 Ma), when the first split occurred between spiny mice of eastern (Somali-Masai) and south-eastern (Zambezian) savannas. Further diversification, mostly in Plio-Pleistocene, and the current distribution of Acomys were influenced by the interplay of global climatic factors (e.g., Messinian salinity crisis, intensification of Northern Hemisphere glaciation) with local geomorphology (mountain chains, aridity belts, water bodies). Combination of divergence dating, species distribution modelling and historical biogeography analysis suggests repeated "out-of-East-Africa" dispersal events into western Africa, the Mediterranean region and Arabia. CONCLUSIONS: The genus Acomys is very suitable model for historical phylogeographic and biogeographic reconstructions of dry non-forested environments in Afro-Arabia. We provide the most thorough phylogenetic reconstruction of the genus and identify major factors that influenced its evolutionary history since the late Miocene. We also highlight the urgent need of integrative taxonomic revision of east African taxa.

Evolution of the Grey-bellied pygmy mouse group: Highly structured molecular diversity with predictable geographic ranges but morphological crypsis.

The grey-bellied pygmy mouse (Mus triton) from the endemic African subgenus Nannomys is a widespread rodent species inhabiting the highlands of eastern and central Africa. Although it has long been considered as a single species, recent data has suggested the existence of a species complex. In order to evaluate the geographical structure and current taxonomy of M. triton, we analysed one mitochondrial and six nuclear genes from individuals covering most of its distribution range. Our analysis revealed the existence of at least five distinct genetic lineages with only marginal overlaps among their distributional ranges. Morphological comparisons, however, showed large overlaps in external body measurements and only a weak differentiation in skull form. Therefore, we suggest maintaining M. triton as a single taxon with pronounced intraspecific genetic structure. Divergence dating analysis placed the most recent common ancestor of the extant lineages of M. triton to the early Pleistocene (about 2.0 Ma). The phylogeographic structure of the species was likely shaped by Pleistocene climatic oscillations and the highly diverse topography of eastern Africa.

Reconstructing evolution at the community level: A case study on Mediterranean amphibians.

Reconstructing reliable timescales for species evolution is an important and indispensable goal of modern biogeography. However, many factors influence the estimation of divergence times, and uncertainty in the inferred time trees remains a major issue that is often insufficiently acknowledged. We here focus on a fundamental problem of time tree analysis: the combination of slow-evolving (nuclear DNA) and fast-evolving (mitochondrial DNA) markers in a single time tree. Both markers differ in their suitability to infer divergences at different time scales (the 'genome-timescale-dilemma'). However, strategies to infer shallow and deep divergences in a single time tree have rarely been compared empirically. Using Mediterranean amphibians as model system that is exceptional in its geographic and taxonomic completeness of available genetic information, we analyze 202 lineages of western Palearctic amphibians across the entire Mediterranean region. We compiled data of four nuclear and five mitochondrial genes and used twelve fossil calibration points widely acknowledged for amphibian evolution. We reconstruct time trees for an extensive lineage-level data set and compare the performances of the different trees: the first tree is based on primary fossil calibration and mitochondrial DNA, while the second tree is based on a combination of primary fossil and on secondary calibrations taken from a nuclear tree using mitochondrial DNA (two-step protocol). Focusing on a set of nodes that are most likely explained by vicariance, we statistically compare the reconstructed alternative time trees by applying a biogeographical plausibility test. Our two-step protocol outperformed the alternative approach in terms of spatial and temporal plausibility. It allows us to infer scenarios for Mediterranean amphibian evolution in eight geographic provinces. We identified several tectonic and climatic events explaining the majority of Mediterranean amphibian divergences, with Plio-Pleistocene climatic fluctuations being the dominant driver for intrageneric evolution. However, often more than one event could be invoked for a specific split. We give recommendations for the use of secondary calibrations in future molecular clock analyses at the community level.

Plio-Pleistocene diversification and biogeographic barriers in southern Australia reflected in the phylogeography of a widespread and common lizard species.

Palaeoclimatic events and biogeographical processes since the mid-Tertiary have played an important role in shaping the evolution and distribution of Australian fauna. However, their impacts on fauna in southern and arid zone regions of Australia are not well understood. Here we investigate the phylogeography of an Australian scincid lizard, Tiliqua rugosa, across southern Australia using mitochondrial DNA (mtDNA) and 11 nuclear DNA markers (nuDNA), including nine anonymous nuclear loci. Phylogenetic analyses revealed three major mtDNA lineages within T. rugosa, geographically localised north and south of the Murray River in southern Australia, and west of the Nullarbor Plain. Molecular variance and population analyses of both mtDNA and nuDNA haplotypes revealed significant variation among the three populations, although potential introgression of nuDNA markers was also detected for the Northern and Southern population. Coalescent times for major mtDNA lineages coincide with an aridification phase, which commenced after the early Pliocene and increased in intensity during the Late Pliocene-Pleistocene. Species distribution modelling and a phylogeographic diffusion model suggest that the range of T. rugosa may have contracted during the Last Glacial Maximum and the locations of optimal habitat appear to coincide with the geographic origin of several distinct mtDNA lineages. Overall, our analyses suggest that Plio-Pleistocene climatic changes and biogeographic barriers associated with the Nullarbor Plain and Murray River have played a key role in shaping the present-day distribution of genetic diversity in T. rugosa and many additional ground-dwelling animals distributed across southern Australia.

The inner ear of the Paranthropus specimen DNH 22 from Drimolen, South Africa.

OBJECTIVES: Morphological variation within the southern African hypodigm of Paranthropus has been the focus of major interest since the earliest discoveries in the "Cradle of Humankind." Given the relevance of the bony labyrinth for investigating fossil primate paleobiodiversity, this article aims to provide additional evidence for assessing the degree of regional variation within Paranthropus through the comparative analysis of the inner ear of DNH 22. MATERIALS AND METHODS: As comparative material, 18 southern African hominin specimens from Sterkfontein, Swartkrans, and Makapansgat (plus published data from Kromdraai B), attributed to Australopithecus, early Homo or Paranthropus, as well as 10 extant human and 10 extant common chimpanzee specimens are investigated. A landmark-based geometric morphometric method is applied for quantitatively assessing labyrinthine morphology. Additionally, cochlear parameters and oval window area are measured. RESULTS: In terms of semicircular canal and cochlear shape, DNH 22 most resembles the Paranthropus specimen SKW 18 from Swartkrans. Both specimens differ from the other Paranthropus specimens investigated in this study by an anteroposteriorly large posterior semicircular canal and a cochlea with loose turns in the apical portion. Conversely, the oval window area in DNH 22 closely fits the range observed in Paranthropus from Swartkrans and Kromdraai B. DISCUSSION: The inner ear of the DNH 22 specimen represents a unique opportunity to provide further insight into the early hominin labyrinthine variation pattern. In particular, the description of DNH 22 raises critical questions on the diversity of the vestibular system and evolutionary pattern of the auditory apparatus in Paranthropus.

Molecular systematics and biogeography of lowland antpittas (Aves, Grallariidae): The role of vicariance and dispersal in the diversification of a widespread Neotropical lineage.

We infer phylogenetic relationships, divergence times, and the diversification history of the avian Neotropical antpitta genera Hylopezus and Myrmothera (Grallariidae), based on sequence data (3,139 base pairs) from two mitochondrial (ND2 and ND3) and three nuclear nuclear introns (TGFB2, MUSK and FGB-I5) from 142 individuals of the 12 currently recognized species in Hylopezus and Myrmothera and 5 outgroup species. Phylogenetic analyses recovered 19 lineages clustered into two major clades, both distributed in Central and South America. Hylopezus nattereri, previously considered a subspecies of H. ochroleucus, was consistently recovered as the most divergent lineage within the Grallaricula/Hylopezus/Myrmothera clade. Ancestral range estimation suggested that modern lowland antpittas probably originated in the Amazonian Sedimentary basin during the middle Miocene, and that most lineages within the Hylopezus/Myrmothera clade appeared in the Plio-Pleistocene. However, the rate of diversification in the Hylopezus/Myrmothera clade appeared to have remained constant through time, with no major shifts over the 20 million years. Although the timing when most modern lineages of the Hylopezus/Myrmothera clade coincides with a period of intense landscape changes in the Neotropics (Plio-Pleistocene), the absence of any significant shifts in diversification rates over the last 20 million years challenges the view that there is a strict causal relationship between intensification of landscape changes and cladogenesis. The relative old age of the Hylopezus/Myrmothera clade coupled with an important role ascribed to dispersal for its diversification, favor an alternative scenario whereby long-term persistence and dispersal across an ever-changing landscape might explain constant rates of cladogenesis through time.

Ethiopian highlands as a cradle of the African fossorial root-rats (genus Tachyoryctes), the genetic evidence.

Root-rats of the genus Tachyoryctes (Spalacidae) are subterranean herbivores occupying open humid habitats in the highlands of Eastern Africa. There is strong disagreement about species diversity of the genus, because some authors accept two species, while others more than ten. Species with relatively high surface activity, the giant root-rat Tachyoryctes macrocephalus, which is by far largest member of the genus, and the more fossorial African root-rat Tachyoryctes splendens, which eventually has been divided up to 12-13 species, represent two major morphological forms within the genus. In our study, we carried out a multilocus analysis of root-rats' genetic diversity based on samples from 41 localities representing most of Tachyoryctes geographic distribution. Using two mitochondrial and three nuclear genes, we found six main genetic clades possibly representing separate species. These clades were organised into three basal groups whose branching is not well resolved, probably due to fast radiation in the late Pliocene and early Pleistocene. Climatic changes in that time, i.e. fast and repeated changes between extremely dry and humid conditions, which both limited root-rat dispersal, probably stimulated their initial genetic diversification. Contrary to expectation based on the largest root-rat diversity in Kenya (up to eight species by some authors), we found the highest diversity in the Ethiopian highlands, because all but one putative species occur there. All individuals outside of Ethiopia belong to a single recently diverged and expanded clade. This species should bear the name T. annectens (Thomas, 1891), and all other names of taxa described from outside of Ethiopia should be considered its junior synonyms. However, to solve taxonomic issues, future detailed morphological analyses should be conducted on all main clades together with genetic analysis of material from areas of their supposed contact. One of the most interesting findings of the study is the internal position of T. macrocephalus in T. splendens sensu lato. This demonstrates the intriguing phenomenon of accelerated morphological evolution of rodents occupying the Afroalpine zone in Ethiopia. Finally, we discuss how the distribution of Tachyoryctes is influenced by competition with another group of subterranean herbivores on the continent, the African mole-rats. We assume that both groups do not compete directly as previously expected, but specialisation to different subterranean niches is the main factor responsible for their spatial segregation.

Evolution of the modern baboon (Papio hamadryas): A reassessment of the African Plio-Pleistocene record.

Baboons (Papio hamadryas) are among the most successful extant primates, with a minimum of six distinctive forms throughout Sub-Saharan Africa. However, their presence in the fossil record is unclear. Three early fossil taxa are generally recognized, all from South Africa: Papio izodi, Papio robinsoni and Papio angusticeps. Because of their derived appearance, P. angusticeps and P. robinsoni have sometimes been considered subspecies of P. hamadryas and have been used as biochronological markers for the Plio-Pleistocene hominin sites where they are found. We reexamined fossil Papio forms from across Africa with an emphasis on their distinguishing features and distribution. We find that P. robinsoni and P. angusticeps are distinct from each other in several cranial features, but overlap extensively in dental size. Contrary to previous assessments, no diagnostic cranio-mandibular material suggests these two forms co-occur, and dental variation at each site is comparable to that within P. h. ursinus, suggesting that only one form is present in each case. P izodi, however, may co-occur with P. robinsoni, or another Papio form, at Sterkfontein Member 4. P izodi appears more primitive than P. robinsoni and P. angusticeps. P. robinsoni is slightly distinct from P. hamadryas subspecies in its combination of features while P. angusticeps might be included within one of the modern P. hamadryas varieties (i.e., P. h. angusticeps). No definitive Papio fossils are currently documented in eastern Africa until the Middle Pleistocene, pointing to southern Africa as the geographic place of origin for the genus. These results have implications for Plio-Pleistocene biochronology and baboon evolution.

The manual pressures of stone tool behaviors and their implications for the evolution of the human hand.

It is widely agreed that biomechanical stresses imposed by stone tool behaviors influenced the evolution of the human hand. Though archaeological evidence suggests that early hominins participated in a variety of tool behaviors, it is unlikely that all behaviors equally influenced modern human hand anatomy. It is more probable that a behavior's likelihood of exerting a selective pressure was a weighted function of the magnitude of stresses associated with that behavior, the benefits received from it, and the amount of time spent performing it. Based on this premise, we focused on the first part of that equation and evaluated magnitudes of stresses associated with stone tool behaviors thought to have been commonly practiced by early hominins, to determine which placed the greatest loads on the digits. Manual pressure data were gathered from 39 human subjects using a Novel Pliance® manual pressure system while they participated in multiple Plio-Pleistocene tool behaviors: nut-cracking, marrow acquisition with a hammerstone, flake production with a hammerstone, and handaxe and flake use. Manual pressure distributions varied significantly according to behavior, though there was a tendency for regions of the hand subject to the lowest pressures (e.g., proximal phalanges) to be affected less by behavior type. Hammerstone use during marrow acquisition and flake production consistently placed the greatest loads on the digits collectively, on each digit and on each phalanx. Our results suggest that, based solely on the magnitudes of stresses, hammerstone use during marrow acquisition and flake production are the most likely of the assessed behaviors to have influenced the anatomical and functional evolution of the human hand.

Cranial vault thickness variation and inner structural organization in the StW 578 hominin cranium from Jacovec Cavern, South Africa.

The Sterkfontein Caves site is one of the richest early hominin fossil localities in Africa. More specifically, the fossiliferous deposits within the lower-lying Jacovec Cavern have yielded valuable hominin remains; prominent among them is the Australopithecus partial cranium StW 578. Due to the fragmentary nature of the braincase, the specimen has not yet been formally assigned to a species. In this context, we employ microtomography to quantify cranial thickness and composition of StW 578 in order to assess its taxonomic affinity. As comparative material, we investigate 10 South African hominin cranial specimens from Sterkfontein (StW 505, Sts 5, Sts 25, Sts 71), Swartkrans (SK 46, SK 48, SK 49) and Makapansgat (MLD 1, MLD 10, MLD 37/38), attributed to either Australopithecus or Paranthropus, as well as 10 extant human and 10 extant chimpanzee crania. Thickness variation in and structural arrangement of the inner and outer cortical tables and the diploë are automatically assessed at regular intervals along one parasagittal and one coronal section. Additionally, topographic cranial vault thickness distribution is visualized using color maps. Comparisons highlight an absolutely and relatively thickened condition of the StW 578 cranial vault versus those of other South African Plio-Pleistocene hominins. Moreover, in StW 578, as well as in the Australopithecus specimens Sts 5 and Sts 71 from Sterkfontein, the diploic layer contributes substantially to cumulative vault thickness (i.e., >60%). Within the comparative sample investigated here, StW 505 and Sts 71 from Sterkfontein Member 4, both attributed to Australopithecus, most closely resemble StW 578 in terms of cranial vault thickness values, tissue proportions, and two- and three-dimensional distributions. Including additional Plio-Pleistocene Australopithecus and Paranthropus crania from South and East Africa in future studies would further help establish morphological variability in these hominin taxa.