Saccorhytus is an early ecdysozoan and not the earliest deuterostome.

The early history of deuterostomes, the group composed of the chordates, echinoderms and hemichordates1, is still controversial, not least because of a paucity of stem representatives of these clades2-5. The early Cambrian microscopic animal Saccorhytus coronarius was interpreted as an early deuterostome on the basis of purported pharyngeal openings, providing evidence for a meiofaunal ancestry6 and an explanation for the temporal mismatch between palaeontological and molecular clock timescales of animal evolution6-8. Here we report new material of S. coronarius, which is reconstructed as a millimetric and ellipsoidal meiobenthic animal with spinose armour and a terminal mouth but no anus. Purported pharyngeal openings in support of the deuterostome hypothesis6 are shown to be taphonomic artefacts. Phylogenetic analyses indicate that S. coronarius belongs to total-group Ecdysozoa, expanding the morphological disparity and ecological diversity of early Cambrian ecdysozoans.

A new primate community from the earliest Oligocene of the Atlantic margin of Northwest Africa: Systematic, paleobiogeographic, and paleoenvironmental implications.

We report a new Paleogene primate community discovered in the uppermost part of the Samlat Formation outcropping on the continental shore of the Rio de Oro, east of the Dakhla peninsula (in the south of Morocco, near the northern border of Mauritania). Fossils consist of isolated teeth, which were extracted by wet screening of estuarine sediments (DAK C2) dating from the earliest Oligocene (ca. 33.5 Ma). These dental remains testify to the presence of at least eight primate species, documenting distinct families, four of which are among the Anthropoidea (Oligopithecidae [Catopithecus aff. browni], Propliopithecidae [?Propliopithecus sp.], Parapithecidae [Abuqatrania cf. basiodontos], and Afrotarsiidae [Afrotarsius sp.]) and four in the Strepsirrhini (a Djebelemuridae [cf. 'Anchomomys' milleri], a Galagidae [Wadilemur cf. elegans], a possible lorisiform [Orogalago saintexuperyi gen. et sp. nov.], and a strepsirrhine of indeterminate affinities [Orolemur mermozi gen. et sp. nov.]). This record of various primates at Dakhla represents the first Oligocene primate community from Northwest Africa, especially from the Atlantic margin of that landmass. Considering primates plus rodents (especially hystricognaths), the taxonomic proximity at the generic (even specific) level between DAK C2 (Dakhla) and the famous Egyptian fossil-bearing localities of the Jebel Qatrani Formation (Fayum Depression), either dating from the latest Eocene (L-41) or from the early Oligocene, suggests the existence of an east-west 'trans-North African' environmental continuum during the latest Eocene-earliest Oligocene time interval. The particularly diverse mammal fauna from DAK C2, recorded within the time window of global climate deterioration characterizing the Eocene/Oligocene transition, suggests that this tropical region of northwest Africa was seemingly less affected, if at all, by the cooling and associated paleoenvironmental and biotic changes documented at that time or at least that the effects were delayed. The expected densely forested paleoenvironment bordering the western margin of North Africa at the beginning of the early Oligocene probably offered better tropical refugia than higher latitudes or more inland areas during the cooling episode.

Paleoproteomics.

Paleoproteomics, the study of ancient proteins, is a rapidly growing field at the intersection of molecular biology, paleontology, archaeology, paleoecology, and history. Paleoproteomics research leverages the longevity and diversity of proteins to explore fundamental questions about the past. While its origins predate the characterization of DNA, it was only with the advent of soft ionization mass spectrometry that the study of ancient proteins became truly feasible. Technological gains over the past 20 years have allowed increasing opportunities to better understand preservation, degradation, and recovery of the rich bioarchive of ancient proteins found in the archaeological and paleontological records. Growing from a handful of studies in the 1990s on individual highly abundant ancient proteins, paleoproteomics today is an expanding field with diverse applications ranging from the taxonomic identification of highly fragmented bones and shells and the phylogenetic resolution of extinct species to the exploration of past cuisines from dental calculus and pottery food crusts and the characterization of past diseases. More broadly, these studies have opened new doors in understanding past human-animal interactions, the reconstruction of past environments and environmental changes, the expansion of the hominin fossil record through large scale screening of nondiagnostic bone fragments, and the phylogenetic resolution of the vertebrate fossil record. Even with these advances, much of the ancient proteomic record still remains unexplored. Here we provide an overview of the history of the field, a summary of the major methods and applications currently in use, and a critical evaluation of current challenges. We conclude by looking to the future, for which innovative solutions and emerging technology will play an important role in enabling us to access the still unexplored "dark" proteome, allowing for a fuller understanding of the role ancient proteins can play in the interpretation of the past.

The game of models: Dietary reconstruction in human evolution.

Despite substantial additions to the paleontological record and unanticipated improvements in analytical techniques since the Journal of Human Evolution was first published, consensus on the diet of early hominin species remains elusive. For instance, the notable advances in the analyses of hominin dental microwear and stable isotopes have provided a plethora of data that have in some instances clouded what was once ostensibly a clear picture of dietary differentiation between and within hominin taxa. In the present study, we explore the reasons why the retrodiction of diet in human evolution has proven vexing over the last half century from the perspective of both ecological and functional-mechanical models. Such models continue to be indispensable for paleobiological reconstructions, but they often contain rigid or unstated assumptions about how primary paleontological data, such as fossils and their geological and taphonomic contexts, allow unambiguous insight into the evolutionary processes that produced them. In theoretical discussions of paleobiology, it has long been recognized that a mapping function of morphology to adaptation is not one-to-one, in the sense that a particular trait cannot necessarily be attributed to a specific selective pressure and/or behavior. This article explores how the intrinsic variability within biological systems has often been underappreciated in paleoanthropological research. For instance, to claim that derived anatomical traits represent adaptations related to stereotypical behaviors largely ignores the importance of biological roles (i.e., how anatomical traits function in the environment), a concept that depends on behavioral flexibility for its potency. Similarly, in the paleoecological context, the underrepresentation of variability within the 'edible landscapes' our hominin ancestors occupied has inhibited an adequate appreciation of early hominin dietary flexibility. Incorporating the reality of variation at organismal and ecological scales makes the practice of paleobiological reconstruction more challenging, but in return, allows for a better appreciation of the evolutionary possibilities that were open to early hominins.

Phylogenetic Signal and Bias in Paleontology.

An unprecedented amount of evidence now illuminates the phylogeny of living mammals and birds on the Tree of Life. We use this tree to measure the phylogenetic value of data typically used in paleontology (bones and teeth) from six data sets derived from five published studies. We ask three interrelated questions: 1) Can these data adequately reconstruct known parts of the Tree of Life? 2) Is accuracy generally similar for studies using morphology, or do some morphological data sets perform better than others? 3) Does the loss of non-fossilizable data cause taxa to occur in misleadingly basal positions? Adding morphology to DNA data sets usually increases congruence of resulting topologies to the well-corroborated tree, but this varies among morphological data sets. Extant taxa with a high proportion of missing morphological characters can greatly reduce phylogenetic resolution when analyzed together with fossils. Attempts to ameliorate this by deleting extant taxa missing morphology are prone to decreased accuracy due to long-branch artifacts. We find no evidence that fossilization causes extinct taxa to incorrectly appear at or near topologically basal branches. Morphology comprises the evidence held in common by living taxa and fossils, and phylogenetic analysis of fossils greatly benefits from inclusion of molecular and morphological data sampled for living taxa, whatever methods are used for phylogeny estimation. [Concatenation; fossilization; morphology; parsimony; systematics; taphonomy; total-evidence.].

A tiny ornithodiran archosaur from the Triassic of Madagascar and the role of miniaturization in dinosaur and pterosaur ancestry.

Early members of the dinosaur-pterosaur clade Ornithodira are very rare in the fossil record, obscuring our understanding of the origins of this important group. Here, we describe an early ornithodiran (Kongonaphon kely gen. et sp. nov.) from the Mid-to-Upper Triassic of Madagascar that represents one of the smallest nonavian ornithodirans. Although dinosaurs and gigantism are practically synonymous, an analysis of body size evolution in dinosaurs and other archosaurs in the context of this taxon and related forms demonstrates that the earliest-diverging members of the group may have been smaller than previously thought, and that a profound miniaturization event occurred near the base of the avian stem lineage. In phylogenetic analysis, Kongonaphon is recovered as a member of the Triassic ornithodiran clade Lagerpetidae, expanding the range of this group into Africa and providing data on the craniodental morphology of lagerpetids. The conical teeth of Kongonaphon exhibit pitted microwear consistent with a diet of hard-shelled insects, indicating a shift in trophic ecology to insectivory associated with diminutive body size. Small ancestral body size suggests that the extreme rarity of early ornithodirans in the fossil record owes more to taphonomic artifact than true reflection of the group's evolutionary history.

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

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

A new species of Comahuesuchus Bonaparte, 1991 (Crocodyliformes: Notosuchia) from the Upper Cretaceous of Neuquén, Lake Barreales, Patagonia, Argentina.

Notosuchian crocodyliforms were major components of the South American Cretaceous biota and, for over 125 years, paleontological fieldwork in this continent recovered several well-preserved fossils of these animals. They are largely recognized for terrestrial life and specialized feeding habits, frequently presenting bizarre taxa such as Comahuesuchus. A new species, Comahuesuchus bonapartei n. sp. (MUCPv 597; cast MN), is described from geological strata of Sierra Barrosa Formation (Upper Turonian) and Portezuelo Formation (Lower Coniacian) of Lake Barreales, Patagonia, Argentina. The new fossil comprises a right dentary bone that shares important and unique anatomical features with specimens of Comahuesuchus brachybuccalis, such as the presence of a well-marked shelf on the lateral surface of the bone; a flat, low, and wide mandibular symphysis; an enlarged, labiolingually compressed caniniform tooth at caudal position in the dentary; the presence of serrated mesial and distal carinae in the caniniform with a faceted labial surface, and the absence or extremely reduction in number of the postcaniniform dentition. C. bonapartei differs from C. brachybuccalis in having individual dentary alveoli, rather a dentition set in groove. Phylogenetic analyses support a sister-relationship between both species, which are well nested within notosuchians.

Deep Time Paleoproteomics: Looking Forward.

The goal of paleoproteomics is to characterize proteins from specimens that have been subjected to the degrading and obscuring effects of time, thus obtaining biological information about tissues or organisms both unobservable in the present and unobtainable through morphological study. Although the description of sequences from Tyrannosaurus rex and Brachylophosaurus canadensis suggested that proteins may persist over tens of millions of years, the majority of paleoproteomic analyses have focused on historical, archeological, or relatively young paleontological samples that rarely exceed 1 million years in age. However, recent advances in methodology and analyses of diverse tissues types (e.g., fossil eggshell, dental enamel) have begun closing the large window of time that remains unexplored in the fossil history of the Cenozoic. In this perspective, we discuss the history and current state of deep time paleoproteomics (DTPp), here defined as paleoproteomic study of samples ∼1 million years (1 Ma) or more in age. We then discuss the future of DTPp research, including what we see as critical ways the field can expand, advancements in technology that can be utilized, and the types of questions DTPp can address if such a future is realized.

Mosaic habitats at Woranso-Mille (Ethiopia) during the Pliocene and implications for Australopithecus paleoecology and taxonomic diversity.

Many important Pliocene hominin specimens have been recovered from Woranso-Mille, a paleontological research area in the Afar region of Ethiopia, including the complete cranium of Australopithecus anamensis, a partial skeleton of Australopithecus afarensis, mandibular and maxillary elements representing a new species, Australopithecus deyiremeda, and a partial foot of an as-yet-unnamed species. Woranso-Mille is the only site, so far, to have reported the co-existence of more than one early hominin species between 3.8 and 3.3 Ma and the temporal overlap between A. anamensis and A. afarensis. Thus, the site has important implications for our understanding of the paleoecology and taxonomic diversity of early hominins and their ecological niche. This paper explores the paleohabitats of Woranso-Mille through its faunal community ecological structure and taxonomic composition using correspondence analysis and Forbes modified similarity index. The results suggest that Pliocene Woranso-Mille was a mosaic of different habitat types, with riparian woodland and floodplain grassland along rivers draining into a lake, along with less mesic habitats such as woodland, grassland, and shrubland. The apparent high level of vegetation heterogeneity may have promoted dietary specializations and niche differentiation among the different Australopithecus species at Woranso-Mille and allowed for their co-existence at the site.

A dense sample of fossil primates (Adapiformes, Notharctidae, Notharctinae) from the Early Eocene Willwood Formation, Wyoming: Documentation of gradual change in tooth area and shape through time.

OBJECTIVES: The Willwood Formation of the southern Bighorn Basin, Wyoming is a fluvial rock sequence that spans approximately 3 million years of early Eocene time. It has yielded one the largest collections of fossil mammals in the world including thousands of dentitions of extinct lemur-like primates known as notharctines. In the southern Bighorn Basin, specimens of these primates have been collected on numerous paleontological expeditions and the stratigraphic levels yielding the dentitions have been carefully recorded. Notharctine dentitions represent a rare opportunity to study morphological variation in a single anatomical system through time among closely related individuals. MATERIALS AND METHODS: Prior studies of Bighorn Basin notharctines through time produced measurements of hundreds of specimens but I report here results from measurement and comparison of the dentitions and dentaries of more than 3,000 specimens, all stratigraphically mapped. RESULTS: Variation in premolar and molar area and variation in dentary depth are apparent throughout the section. Specimens with relatively small teeth and dentaries are known from the older part of the section. In younger rocks, variation in tooth area among specimens increases. Variation in tooth area is continuous and overlaps extensively both within and between stratigraphic levels. Other dental variables examined by inspection change in a mosaic and continuous fashion through the section. These features include variation in the presence and number of paraconids on the lower fourth premolar (p4), the size and shape of the entoconid notch on the lower first and second molars, and the relative development of the pseudohypocone, mesostyle, and cingula on the upper molars. DISCUSSION: These broad patterns can be identified despite notharctine alpha taxonomy being in need of extensive revision and, importantly, simplification. Such revision is beyond the scope of this article but is essential if we are to develop a taxonomy that is both free of stratigraphic influence and useful for rapid, repeatable species assignment. Boundaries among the patterns of tokogenesis, anagenesis, and cladogenesis are blurred in this dense sample of extinct primates. While pattern of evolution, a population-level phenomenon, may be difficult to falsify in the fossil record, this notharctine sample suggests that in the rare instance such as this, when the fossil record is densely sampled, change through time is continuous and more consistent with gradual evolution.

Exploring late Paleolithic and Mesolithic diet in the Eastern Alpine region of Italy through multiple proxies.

OBJECTIVES: The analysis of prehistoric human dietary habits is key for understanding the effects of paleoenvironmental changes on the evolution of cultural and social human behaviors. In this study, we compare results from zooarchaeological, stable isotope and dental calculus analyses as well as lower second molar macrowear patterns to gain a broader understanding of the diet of three individuals who lived between the end of the Late Pleistocene and the Early Holocene (ca., 17-8 ky cal BP) in the Eastern Alpine region of Italy. MATERIALS AND METHODS: We analyze individuals buried at the sites of Riparo Tagliente (Verona), Riparo Villabruna, and Mondeval de Sora (Belluno). The three burials provide a unique dataset for diachronically exploring the influence of climatic changes on human subsistence strategies. RESULTS: Isotopic results indicate that all individuals likely relied on both terrestrial and freshwater animal proteins. Even though dental calculus analysis was, in part, hindered by the amount of mineral deposit available on the teeth, tooth macrowear study suggests that the dietary habits of the individuals included plant foods. Moreover, differences in macrowear patterns of lower second molars have been documented between Neanderthals and modern humans in the present sample, due to a prevalence of Buccal wear among the former as opposed to higher values of Lingual wear in modern human teeth. DISCUSSION: Isotopic analyses have emphasized the contribution of animal proteins in the diet of the three foragers from the Eastern Alpine region. The possible intake of carbohydrate-rich plant foods, suggested by the retrieval of plant remains in dental calculus, is supported by the signal of macrowear analysis. Moreover, the latter method indicates that the distribution of macrowear in lower second molars (M2 s) allows us to discriminate between Neanderthals and modern humans within the present reference sample. Overall, our results show these three prehistoric hunter-gatherers were well adapted to the environment in which they lived exploiting many natural resources.

An abundance of developmental anomalies and abnormalities in Pleistocene people.

Diverse developmental abnormalities and anomalous features are evident in the Pleistocene Homo fossil record, varying from minor but rare dental, vertebral, and carpal variants to exceptional systemic disorders. There are currently 75 documented anomalies or abnormalities from 66 individuals, spanning the Pleistocene but primarily from the Late Pleistocene Middle and Upper Paleolithic with their more complete skeletal remains. The expected probabilities of finding these variants or developmental disorders vary from <5% to <0.0001%, based on either recent human incidences or relevant Pleistocene sample distributions. Given the modest sample sizes available for the skeletal or dental elements in question, especially if the samples are appropriately limited in time and geography, the cumulative multiplicative probability of finding these developmental changes is vanishingly small. These data raise questions regarding social survival abilities, differing mortuary treatments of the biologically unusual, the role of ubiquitous stress among these Pleistocene foragers, and their levels of consanguinity. No single factor sufficiently accounts for the elevated level of these developmental variants or the low probability of finding them in the available paleontological record.

A New Species of Typhlomys (Platacanthomyidae, Rodentia) from the Middle Pleistocene of Northern Vietnam.

A pygmy dormouse Typhlomys stegodontis sp. nov. is described on the base of the maxillary fragment and isolated teeth from the Middle Pleistocene Tham Hai cave locality in northern Vietnam (Lang Son Province). This first finding of the fossil Platacanthomyidae in Vietnam fills the Middle Pleistocene gap in the paleontological record of the family.

Giant Hyena Pachycrocuta brevirostris (Hyaenidae, Carnivora) from the Lower Pleistocene of Taurida Cave, Crimea.

The dental remains of a giant hyena Pachycrocuta brevirostris (Gervais, 1850) from the Early Pleistocene locality of the Taurida cave (Crimea, Late Villafranchian, 1.8-1.5 Ma) are described. This species was a typical representative of the Villafranchian fauna of the Eastern Mediterranean. The Taurida cave was occasionally used by hyenas and other carnivorans as a den and retreat.

Early Oligocene chinchilloid caviomorphs from Puerto Rico and the initial rodent colonization of the West Indies.

By their past and present diversity, rodents are among the richest components of Caribbean land mammals. Many of these became extinct recently. Causes of their extirpation, their phylogenetic affinities, the timing of their arrival in the West Indies and their biogeographic history are all ongoing debated issues. Here, we report the discovery of dental remains from Lower Oligocene deposits (ca 29.5 Ma) of Puerto Rico. Their morphology attests to the presence of two distinct species of chinchilloid caviomorphs, closely related to dinomyids in a phylogenetic analysis, and thus of undisputable South American origin. These fossils represent the earliest Caribbean rodents known thus far. They could extend back to 30 Ma the lineages of some recently extinct Caribbean giant rodents (Elasmodontomys and Amblyrhiza), which are also retrieved here as chinchilloids. This new find has substantial biogeographic implications because it demonstrates an early dispersal of land mammals from South America to the West Indies, perhaps via the emergence of the Aves Ridge that occurred ca 35-33 Ma (GAARlandia hypothesis). Considering both this new palaeontological evidence and recent molecular divergence estimates, the natural colonization of the West Indies by rodents probably occurred through multiple and time-staggered dispersal events (chinchilloids, then echimyid octodontoids (spiny rats/hutias), caviids and lastly oryzomyin muroids (rice rats)).

Enamel chipping in Taï Forest cercopithecids: Implications for diet reconstruction in paleoanthropological contexts.

Antemortem enamel chipping in living and fossil primates is often interpreted as evidence of hard-object feeding (i.e., 'durophagy'). Laboratory analyses of tooth fracture have modeled the theoretical diets and loading conditions that may produce such chips. Previous chipping studies of nonhuman primates tend to combine populations into species samples, despite the fact that species can vary significantly in diet across their ranges. Chipping is yet to be analyzed across population-specific species samples for which long-term dietary data are available. Here, we test the association between enamel chipping and diet in a community of cercopithecid primates inhabiting the Taï Forest, Ivory Coast. We examined fourth premolars and first molars (n = 867) from naturally deceased specimens of Cercocebus atys, Colobus polykomos, Piliocolobus badius,Procolobus verus, and three species of Cercopithecus. We found little support for a predictive relationship between enamel chipping and diet across the entire Taï monkey community. Cercocebus atys, a dedicated hard-object feeder, exhibited the highest frequencies of (1) chipped teeth and (2) chips of large size; however, the other monkey with a significant degree of granivory, Co. polykomos, exhibited the lowest chip frequency. In addition, primates with little evidence of mechanically challenging or hard-food diets-such as Cercopithecus spp., Pi. badius, and Pr. verus-evinced higher chipping frequencies than expected. The equivocal and stochastic nature of enamel chipping in the Taï monkeys suggests nondietary factors contribute significantly to chipping. A negative association between canopy preference and chipping suggests a role of exogenous particles in chip formation, whereby taxa foraging closer to the forest floor encounter more errant particulates during feeding than species foraging in higher strata. We conclude that current enamel chipping models may provide insight into the diets of fossil primates, but only in cases of extreme durophagy. Given the role of nondietary factors in chip formation, our ability to reliably reconstruct a range of diets from a gradient of chipping in fossil taxa is likely weak.

New fossils of Mesopithecus from Hasnot, Pakistan.

Here, we report on a new collection of mostly isolated molars of a colobine monkey from near Hasnot on the Potwar Plateau of Pakistan. The specimens are from three late Miocene localities, with ages constrained to between 7.9 and 7.1 Ma. Morphological and metrical comparisons of the new Hasnot molars with those of previously recognized Mesopithecus species and living Asian colobines lead to the conclusion that the Hasnot colobine is most probably Mesopithecus, but not Mesopithecus pentelicus. The most morphologically distinctive aspect of the Hasnot specimens is the lower third molars, which exhibit large and bulbous protoconids set off by deeply incised mesial buccal and median buccal clefts and large, broad distobuccally placed hypoconulids. Colobine specimens previously recovered from the Potwar Plateau have been assigned to Mesopithecus sivalensis, but because these specimens have not yet been fully described, a detailed comparison with the new Hasnot specimens is not yet possible. For these reasons, we assign the new Hasnot colobine fossils to cf. Mesopithecus sp. Mesopithecus was one of the most widespread and successful of late Miocene primates. As a colobine equipped with features of the molar teeth, limbs, and, presumably, gut enabling it to succeed in more highly seasonal woodland environments, Mesopithecus was able to rapidly disperse into and adapt to the conditions in South Asia brought about by profound climatic and concomitant environmental change during the latest Miocene.

Inner morphological and metric characterization of the molar remains from the Montmaurin-La Niche mandible: The Neanderthal signal.

Here, we present a metric and morphological study of the molar remains from the Montmaurin-La Niche mandible by means of microcomputed tomography. According to the last analysis, based on the combination of geomorphological and paleontological data, the level bearing this human mandible probably corresponds to the marine isotope stages (MIS) 7. These data place the Montmaurin-La Niche in a chronologically intermediate position between the Neanderthals and the Middle Pleistocene fossils (e.g., Sima de los Huesos, la Caune de l'Arago). A recent study has revealed that while the mandible is more closely related to the Early and Middle Pleistocene African and Eurasian populations, the morphology of the outer enamel surfaces of its molars is typical of the Neanderthal linage. The data presented here are in line with this finding because the morphology of the enamel-dentine junction of the molars is similar to that of Neanderthals, whereas the absolute and relative enamel thickness values (2D and 3D) are closer to those exhibited by some Early Pleistocene hominins. Moreover, the pulp cavity morphology and proportions are in concordance with the Neanderthal populations. Our results strengthen the hypothesis that the settlement of Europe could be the result of several migrations, at different times, originated from a common source population. Thus, the variability in the European Middle Pleistocene populations (e.g., Montmaurin, Sima de los Huesos, Arago, Mala Balanica) could indicate different migrations at different times and/or population fragmentation, without excluding the possible hybridization between residents and new settlers.

Rodents and other terrestrial small mammals from Kanapoi, north-western Kenya.

Excavations at Kanapoi in north-western Kenya have yielded the most numerically abundant and taxonomically diverse early Pliocene (4.19 Ma) terrestrial small mammal assemblage known from Kenya. A minimum of 15 species are reported, including soricids, sengis, leporids, and rodents: all taxa are referable to extant genera, with the exception of the murine rodent, Saidomys. The majority of the terrestrial small mammals are derived from a bone bed at Nzube's Mandible Site, closely associated with the holotype mandible of Australopithecus anamensis. A smaller number were surface-collected or obtained from screening at several other sites, including the Bat Site. Most small mammals from Nzube's Mandible Site and the Bat Site likely represent prey accumulated as regurgitated pellets from owls, in particular barn owls or giant eagle owls. The small mammal fauna is dominated by the spiny mouse, Acomys: the next most commonly recovered taxa are the multimammate mouse, Mastomys, and the African gerbil, Gerbilliscus. Comparisons of the Kanapoi fauna to other eastern African late Miocene-Pliocene (and one Pleistocene) faunas at the generic level suggest the greatest similarity is to Lemudong'o, Kenya, and Omo B and Aramis, Ethiopia. Further similarities with other localities such as Laetoli, Tanzania, and Hadar, Ethiopia, suggest the existence of a corridor for dispersal along the East African Rift Valley between Ethiopia, Kenya, and Tanzania in the early Pliocene. Further comparisons of the relative abundances of individuals in different families (or subfamilies) emphasize the distinctiveness of the Kanapoi small mammal fauna. The Kanapoi fauna is likely derived from a heterogeneous but relatively arid environment.