Talking heads: disentangling the shape changes of the large extant hippopotamus during its ontogenetic development.

Using a 3D geometric morphometrics approach, we shed light on the major changes during the ontogenetic development of Hippopotamus amphibius. The main shift in the shape variability of cranial elements in juveniles can probably be related to the interruption of suckling. A less stressed shift in the cranial variability can probably be related to the attainment of sexual maturity. Semilandmarks and landmarks are powerful tools to discriminate between extant species and, therefore, this technique will be particularly useful in the future to study fossil ones as well.

Increase on environmental seasonality through the European Early Pleistocene inferred from dental enamel hypoplasia.

An in-depth study of the Early Pleistocene European remains of Hippopotamus has allowed the first detailed description of the incidence and types of dental alterations related to palaeopathologies and potentially linked to climatic and environmental factors. The results of a long-term qualitative and quantitative assessment highlight the importance of nutrient deficiencies on the development of dental enamel hypoplasia in Hippopotamus. Glacial cyclicity and the resulting changes in humidity and plant community structure conditioned the local environments critical for the survival of this taxon. Two main intervals of putative constrained nutritionally restrictions were detected at ca. 1.8 Ma and ca. 0.86 Ma (i.e., MIS63 and MIS21, respectively). Statistical comparisons show an increase in the frequency of dental hypoplasia between these two chronological periods, thus reinforcing the idea of increased seasonality in the circum-Mediterranean environments during the Early Pleistocene.

Paleoenvironmental inferences on the Late Miocene hominoid-bearing site of Can Llobateres (NE Iberian Peninsula): An ecometric approach based on functional dental traits.

Hispanopithecus laietanus from the Late Miocene (9.8 Ma) of Can Llobateres 1 (CLL1; Vallès-Penedès Basin, NE Iberian Peninsula) represents one of the latest occurrences of fossil apes in Western mainland Europe, where they are last recorded at ∼9.5 Ma. The paleoenvironment of CLL1 is thus relevant for understanding the extinction of European hominoids. To refine paleoenvironmental inferences for CLL1, we apply ecometric models based on functional crown type (FCT) variables-a scoring scheme devised to capture macroscopic functional traits of occlusal shape and wear surfaces of herbivorous large mammal molars. Paleotemperature and paleoprecipitation estimates for CLL1 are provided based on published regional regression models linking average FCT of large herbivorous mammal communities to climatic conditions. A mapping to Whittaker's present-day biome classification is also attempted based on these estimates, as well as a case-based reasoning via canonical variate analysis of FCT variables from five relevant biomes. Estimates of mean annual temperature (25 °C) and mean annual precipitation (881 mm) classify CLL1 as a tropical seasonal forest/savanna, only in partial agreement with the canonical variate analysis results, which classify CLL1 as a tropical rainforest with a higher probability. The former biome agrees better with previous inferences derived from fossil plants and mammals, as well as preliminary isotopic data. The misclassification of CLL1 as a tropical forest is attributed to the mixture of forest-adapted taxa with others adapted to more open environments, given that faunal and plant composition indicates the presence of a dense wetland/riparian forest with more open woodlands nearby. The tested FCT ecometric approaches do not provide unambiguous biome classification for CLL1. Nevertheless, our results are consistent with those from other approaches, thus suggesting that FCT variables are potentially useful to investigate paleoenvironmental changes through time and space-including those that led to the extinction of European Miocene apes.

The earliest evidence of deep-sea vertebrates.

Vertebrate macroevolution has been punctuated by fundamental habitat transitions from shallow marine origins to terrestrial, freshwater, and aerial environments. Invasion of the deep sea is a less well-known ecological shift because of low fossilization potential and continual loss of abyssal fossil record by ocean floor subduction. Therefore, there has been a lack of convincing evidence of bottom-living vertebrates from pre-Paleogene deep seas. Here, we describe trace fossils from abyssal plain turbidites of the Tethys Ocean, which, combined with nannofossil dating, indicate that fishes have occupied the deep seafloor since at least the Early Cretaceous (Hauterivian-Barremian). These structures are identical to those produced by modern demersal fishes that feed by either scratching the substrate or expose their prey by water flow generated by suction or jetting. The trace fossils suggest activity of at least three fish species exploiting a productive abyssal invertebrate sediment fauna. These observations are consistent with Early Cretaceous vertebrate transition to the deep sea triggered by the availability of new food sources. Our results anticipate the appearance of deep-seafloor fishes in the fossil record by over 80 My while reassessing the mode of vertebrate colonization of the deep sea.

Trace Elements in Soil and Urban Groundwater in an Area Impacted by Metallurgical Activity: Health Risk Assessment in the Historical Barga Municipality (Tuscany, Italy).

Trace elements were measured in soil and groundwater collected within the Fornaci di Barga urban area (Serchio River Valley, Tuscany, Italy), a territory that integrates natural assets with touristic vocation, impacted by long-lasting metallurgical activity. Epidemiological studies highlighted that the area surrounding the industrial plants is characterized by a persistent excess of diseases, attributed to heavy metal pollution. Soils were taken in school gardens, public parks, sport grounds and roadsides. The results indicate that Cu, Zn and Cd represent the main contaminants in surface soil, likely originated by deposition of airborne particulate matter from metallurgical activity. Risk assessment considering soil ingestion and dermal contact exposure routes revealed that the cadmium Hazard Quotient approaches unity for children, and the cadmium risk-based concentration obtained by combining exposure information with toxicity data is only slightly lower compared with the cadmium maximum concentration actually measured in soil. Groundwater does not show evidence of trace metal contamination, suggesting that the migration of contaminants from soil to subsurface is a slow process. However, assessment of the possible interconnections between shallow and deep-seated aquifers requires monitoring to be continued. The obtained results highlight the possible link between space clusters of diseases and metal concentration in soil.

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.

A comparative study of the Early Pleistocene carnivore guild from Dmanisi (Georgia).

The carnivore guild of the Early Pleistocene site of Dmanisi is among the most diverse of the Early Pleistocene of the entire Old World. It includes 14 carnivoran taxa: Homotherium latidens, Megantereon whitei, Panthera onca georgica, Acinonyx pardinensis, Lynx issiodorensis; Pachycrocuta brevirostris; Canis (Xenocyon) lycaonoides, Canis borjgali, Vulpes alopecoides; Ursus etruscus; Lutra sp., Martes sp., Meles sp., and Pannonictis sp. The analysis of this rich carnivore guild was carried out under different methodological approaches to compare the assemblage with other chronological coeval European, Asian, and African sites from a paleobiological perspective. To achieve the goal, we used a permutational hierarchical method called boostrapping cluster analysis based on taxonomic absence/presence matrices (at both generic and specific level) and on ecological matrices (considering dietary preferences/hunting strategies of each carnivoran) and carried out Mantels tests assessing magnitude of time, space, ecology, and taxonomy as source of difference between guilds. Our results suggest a close similarity among the Dmanisi carnivore assemblage and other guilds recorded from European late Villafranchian sites such as Pirro Nord, Venta Micena, and Apollonia 1 and, in a lesser extent, to European Epivillafranchian sites as Vallonnet, Untermassfeld, or the Vallparadís Section. Early to Middle Pleistocene Asian carnivore assemblages display several similarities with the Dmanisi guild mainly in the record and diversity of felid and the canid ecomorphotypes. Eastern African sites such as Olduvai and Omo, as well as South African sites, display a lower similarity with the studied sample, basically for the most diverse hyenid taphocoenoses. To sum up, the present study suggests a close similarity between the Dmanisi carnivore guild and other European Late Early Pleistocene assemblages without close parallels with African or Asian assemblages.

Zoogeographic significance of Dmanisi large mammal assemblage.

We undertake a comparative mammalian zoogeographic analysis with the aim of revealing the extent to which the Dmanisi Early Pleistocene large mammal assemblage resembles, at the genus level, African, Arabian, and Eurasian localities of similar age. The inclusion of Old World Pliocene and Pleistocene mammalian faunas provides us with insights into the provincial origins of specific mammalian taxa and permits us to assess the relative affiliation of the Dmanisi mammalian faunas to other faunas in the Old World. Our analysis also allows us to consider hypotheses about the timing and direction of zoogeographic connections between western Eurasia and Africa during the Early Pleistocene. We utilize multiple zoogeographic analytical tools as a cross-comparison of Dmanisi with 42 other Eurasian and African mammalian-bearing localities between 2.7 and 0.7 Ma. Overall, we find that Dmanisi compares most closely with a subgroup of Greek, Italian, and Spanish localities that are slightly younger than Dmanisi itself. This could suggest a progressive dispersal from East to West of the large mammal communities during the late Early Pleistocene and the first occurrence at Dmanisi, and then later in Western Europe, of some taxa such as Stephanorhinus ex gr. etruscus-hundsheimensis, Equus altidens, Bison georgicus, Soergelia minor, Megantereon whitei, Canis borjgali, Canis (Xenocyon) lycaonoides. Dmanisi's habitats included drier areas, probably of open wooded savannah and grassland and by mountainous to semiarid rocky terrain. There is evidence that Dmanisi records short intervals of increased aridity in the middle part of the succession contemporaneous with the occurrence of Homo.

Evolution of Old World Equus and origin of the zebra-ass clade.

Evolution of the genus Equus has been a matter of long debate with a multitude of hypotheses. Currently, there is no consensus on either the taxonomic content nor phylogeny of Equus. Some hypotheses segregate Equus species into three genera, Plesippus, Allohippus and Equus. Also, the evolutionary role of European Pleistocene Equus stenonis in the origin of the zebra-ass clade has been debated. Studies based on skull, mandible and dental morphology suggest an evolutionary relationship between North American Pliocene E. simplicidens and European and African Pleistocene Equus. In this contribution, we assess the validity of the genera Plesippus, Allohippus and Equus by cladistic analysis combined with morphological and morphometrical comparison of cranial anatomy. Our cladistic analysis, based on cranial and postcranial elements (30 taxa, 129 characters), supports the monophyly of Equus, denies the recognition of Plesippus and Allohippus and supports the derivation of Equus grevyi and members of the zebra-ass clade from European stenonine horses. We define the following evolutionary steps directly relevant to the phylogeny of extant zebras and asses: E. simplicidens-E. stenonis-E. koobiforensis-E. grevyi -zebra-ass clade. The North American Pliocene species Equus simplicidens represents the ancestral stock of Old World Pleistocene Equus and the zebra-ass clade. Our phylogenetic results uphold the most recent genomic outputs which indicate an age of 4.0-4.5 Ma for the origin and monophyly of Equus.

Paleoecology, biochronology, and paleobiogeography of Eurasian Rhinocerotidae during the Early Pleistocene: The contribution of the fossil material from Dmanisi (Georgia, Southern Caucasus).

Rhinocerotidae represents a common element in the Eurasian Pleistocene faunas. Origin, dispersal route, and biochronology of several species are still poorly understood due to gaps in the fossil record, in particular from central Eurasia. A remarkable collection of rhinoceros remains was recovered from the Early Pleistocene site of Dmanisi (Georgia). This collection is unique for the Early Pleistocene Rhinocerotidae records due to its abundance in remains, its age (ca 1.8 Ma) and geographic position (between Eastern and Western Eurasia). Two crania, which display some different morphological traits, are assigned to two different morphotypes and investigated by means of geometric morphometrics using landmarks and semilandmarks. Shapes in lateral and dorsal views of different Rhinocerotini species are compared with the studied crania to infer paleoecological information. The shape in the lateral view reflects ecological niche, in particular feeding type from browsing to grazing, and it also represent taxonomic discrimination. Morphotypes 1 and 2 from Dmanisi fall in two different clusters, corresponding to two different species, notably in lateral view. The results suggest a niche partitioning during the Early Pleistocene of Dmanisi between a browse-dominated and a grass-dominated mixed feeders, or possibly the presence of two ecomorphotypes of the same species. A comprehensive update of the Early Pleistocene occurrences of Eurasian Rhinocerotidae is reported in the discussion on the paleoecology of the extinct Northern Eurasian rhinocerotines.

Early Pleistocene enamel proteome from Dmanisi resolves Stephanorhinus phylogeny.

The sequencing of ancient DNA has enabled the reconstruction of speciation, migration and admixture events for extinct taxa1. However, the irreversible post-mortem degradation2 of ancient DNA has so far limited its recovery-outside permafrost areas-to specimens that are not older than approximately 0.5 million years (Myr)3. By contrast, tandem mass spectrometry has enabled the sequencing of approximately 1.5-Myr-old collagen type I4, and suggested the presence of protein residues in fossils of the Cretaceous period5-although with limited phylogenetic use6. In the absence of molecular evidence, the speciation of several extinct species of the Early and Middle Pleistocene epoch remains contentious. Here we address the phylogenetic relationships of the Eurasian Rhinocerotidae of the Pleistocene epoch7-9, using the proteome of dental enamel from a Stephanorhinus tooth that is approximately 1.77-Myr old, recovered from the archaeological site of Dmanisi (South Caucasus, Georgia)10. Molecular phylogenetic analyses place this Stephanorhinus as a sister group to the clade formed by the woolly rhinoceros (Coelodonta antiquitatis) and Merck's rhinoceros (Stephanorhinus kirchbergensis). We show that Coelodonta evolved from an early Stephanorhinus lineage, and that this latter genus includes at least two distinct evolutionary lines. The genus Stephanorhinus is therefore currently paraphyletic, and its systematic revision is needed. We demonstrate that sequencing the proteome of Early Pleistocene dental enamel overcomes the limitations of phylogenetic inference based on ancient collagen or DNA. Our approach also provides additional information about the sex and taxonomic assignment of other specimens from Dmanisi. Our findings reveal that proteomic investigation of ancient dental enamel-which is the hardest tissue in vertebrates11, and is highly abundant in the fossil record-can push the reconstruction of molecular evolution further back into the Early Pleistocene epoch, beyond the currently known limits of ancient DNA preservation.

The Aggradational Successions of the Aniene River Valley in Rome: Age Constraints to Early Neanderthal Presence in Europe.

We revise the chronostratigraphy of several sedimentary successions cropping out along a 5 km-long tract of the Aniene River Valley in Rome (Italy), which yielded six hominin remains previously attributed to proto- or archaic Neanderthal individuals, as well as a large number of lithic artefacts showing intermediate characteristics somewhere between the local Acheulean and Mousterian cultures. Through a method of correlation of aggradational successions with post-glacial sea-level rises, relying on a large set of published 40Ar/39Ar ages of interbedded volcanic deposits, we demonstrate that deposition of the sediments hosting the human remains spans the interval 295-220 ka. This is consistent with other well constrained ages for lithic industries recovered in England, displaying transitional features from Lower to Middle Paleolithic, suggesting the appearance of Mode 3 during the MIS 9-MIS 8 transition. Moreover, the six human bone fragments recovered in the Aniene Valley should be regarded as the most precisely dated and oldest hominin remains ascribable to Neanderthal-type individuals in Europe, discovered to date. The chronostratigraphic study presented here constitutes the groundwork for addressing re-analysis of these remains and of their associated lithic industries, in the light of their well-constrained chronological picture.