Evolution of the Family Equidae, Subfamily Equinae, in North, Central and South America, Eurasia and Africa during the Plio-Pleistocene.

Studies of horse evolution arose during the middle of the 19th century, and several hypotheses have been proposed for their taxonomy, paleobiogeography, paleoecology and evolution. The present contribution represents a collaboration of 19 multinational experts with the goal of providing an updated summary of Pliocene and Pleistocene North, Central and South American, Eurasian and African horses. At the present time, we recognize 114 valid species across these continents, plus 4 North African species in need of further investigation. Our biochronology and biogeography sections integrate Equinae taxonomic records with their chronologic and geographic ranges recognizing regional biochronologic frameworks. The paleoecology section provides insights into paleobotany and diet utilizing both the mesowear and light microscopic methods, along with calculation of body masses. We provide a temporal sequence of maps that render paleoclimatic conditions across these continents integrated with Equinae occurrences. These records reveal a succession of extinctions of primitive lineages and the rise and diversification of more modern taxa. Two recent morphological-based cladistic analyses are presented here as competing hypotheses, with reference to molecular-based phylogenies. Our contribution represents a state-of-the art understanding of Plio-Pleistocene Equus evolution, their biochronologic and biogeographic background and paleoecological and paleoclimatic contexts.

MicroWeaR: A new R package for dental microwear analysis.

Mastication of dietary items with different mechanical properties leaves distinctive microscopic marks on the surface of tooth enamel. The inspection of such marks (dental microwear analysis) is informative about the dietary habitus in fossil as well as in modern species. Dental microwear analysis relies on the morphology, abundance, direction, and distribution of these microscopic marks. We present a new freely available software implementation, MicroWeaR, that, compared to traditional dental microwear tools, allows more rapid, observer error free, and inexpensive quantification and classification of all the microscopic marks (also including for the first time different subtypes of scars). Classification parameters and graphical rendering of the output are fully settable by the user. MicroWeaR includes functions to (a) sample the marks, (b) classify features into categories as pits or scratches and then into their respective subcategories (large pits, coarse scratches, etc.), (c) generate an output table with summary information, and (d) obtain a visual surface-map where marks are highlighted. We provide a tutorial to reproduce the steps required to perform microwear analysis and to test tool functionalities. Then, we present two case studies to illustrate how MicroWeaR works. The first regards a Miocene great ape obtained from through environmental scanning electron microscope, and other a Pleistocene cervid acquired by a stereomicroscope.

Stable carbon isotope reconstructions of diet and paleoenvironment from the late Middle Pleistocene Snake Cave in Northeastern Thailand.

Thailand's geographical location in the tropics and almost complete, relatively uninterrupted forest cover makes it valuable for paleodiet and paleoclimate research. We present the first dietary and environmental reconstructions in Northeastern Thailand, using stable isotope abundances in mammalian tooth enamel from the late Middle Pleistocene locality, Tham Wiman Nakin (Snake Cave), which reflect a much higher (over 70%) than modern (13%) occurrence of C4 plants. Bovids and cervids appear to have had almost entirely a C4 plant diet. Carnivores consumed a mixture of C3 (suids) and C4 (bovids, cervids) consumers. Rhinoceroses and orangutan appear to have maintained their preference through time for forested or open C3 environment, respectively. (13)C/(12)C from bone bioapatite, horn and hair of modern Southeast Asian mammals almost exclusively demonstrate C3 vegetation dominance. C4 consumption is rare in analysed modern species and it could be related to anthropogenic influences such as ingestion of domestic crops or livestock. Interesting implications emerge in the C4 vegetation distribution in southern Eurasian ecosystems, indicating that Southeast Asia, south of the Tibet, could be part of the global C4 vegetation spread, which occurred around 7 Ma. However, the C4 percentage in ecosystems varied geographically. Despite modern reversal towards C3 habitats due to factors such as increasing CO(2), we think that anthropological influences may be responsible for habitat and dietary changes in extant species. Bovids demonstrate the most significant shift in diet and habitat through time, from C4-dominated open habitats to C3-dominated habitats indicative of dense forest understory.

The shape of contention: adaptation, history, and contingency in ungulate mandibles.

Mandibles and teeth of ungulates have been extensively studied to discern the functional significance of their design. Grazing ungulates have deeper mandibles, longer coronoid processes, flatter incisor arcades, and more hypsodont molars in comparison to browsers. If the functional significance of both mandible and teeth shapes is well-established, it remains uncertain to what extent mandible shapes are really adapted to grazing, meaning that they evolved either to serve their current biological function or just as a structural requirement to accommodate higher crowned molars. Here, we address this question by studying the contribution of phylogeny, hypsodonty, and body size to mandibular shape variation. The mandible shape appeared to be significantly influenced by hypsodonty but not by body size. Interestingly, hypsodonty-related changes influenced the tooth row in artiodactyls and perissodactyls significantly but in the opposite directions, which is ultimately related to their different digestive strategies. Yet, we obtained a strong phylogenetic effect in perissodactyls, suggesting that their mandible shape should be strongly inherited. The strength of this effect was not significant within artiodactyls (where hypsodonty explained much more variance in mandible shape). Digestive strategy is deemed to interplay with hypsodonty to produce different paths of adaptation to particular diets in ungulates.

Human influence on distribution and extinctions of the late Pleistocene Eurasian megafauna.

Late Pleistocene extinctions are of interest to paleontological and anthropological research. In North America and Australia, human occupation occurred during a short period of time and overexploitation may have led to the extinction of mammalian megafauna. In northern Eurasia megafaunal extinctions are believed to have occurred over a relatively longer period of time, perhaps as a result of changing environmental conditions, but the picture is much less clear. To consider megafaunal extinction in Eurasia, we compare differences in the geographical distribution and commonness of extinct and extant species between paleontological and archaeological localities from the late middle Pleistocene to Holocene. Purely paleontological localities, as well as most extinct species, were distributed north of archaeological sites and of the extant species, suggesting that apart from possible differences in adaptations between humans and other species, humans could also have a detrimental effect on large mammal distribution. However, evidence for human overexploitation applies only to the extinct steppe bison Bison priscus. Other human-preferred species survive into the Holocene, including Rangifer tarandus, Equus ferus, Capreolus capreolus, Cervus elaphus, Equus hemionus, Saiga tatarica, and Sus scrofa. Mammuthus primigenius and Megaloceros giganteus were rare in archaeological sites. Carnivores appear little influenced by human presence, although they become rarer in Holocene archaeological sites. Overall, the data are consistent with the conclusion that humans acted as efficient hunters selecting for the most abundant species. Our study supports the idea that the late Pleistocene extinctions were environmentally driven by climatic changes that triggered habitat fragmentation, species range reduction, and population decrease, after which human interference either by direct hunting or via indirect activities probably became critical.