Early Agenian rhinocerotids from Wischberg (Canton Bern, Switzerland) and clarification of the systematics of the genus Diaceratherium.

BACKGROUND: Wischberg is a Swiss locality in Bern Canton which has yielded numerous vertebrates remains from the earliest Miocene (= MN1). It has a very rich faunal diversity, one of the richest in Switzerland for this age. Among all the mammals reported in the original faunal list 70 years ago, three rhinocerotid species were identified. The material consists of two fragmentary skulls, cranial fragments, several mandibles, teeth and postcranial bones, in a rather good state of preservation. RESULTS: After reexamination of the material from this locality (curated in three different Swiss museums) and comparison with holotype specimens, we show that all rhinocerotid specimens from Wischberg can be referred to two species only. Most of the material can be attributed to the large-sized teleoceratine Diaceratherium lemanense, while only a few specimens, including a skull and mandible, belong to the much smaller sized Pleuroceros pleuroceros. We describe and illustrate for the first time most of these fossil remains. However, the systematics of the genus Diaceratherium is currently controversial, and based on our new observations we consider seven species as valid, though a large-scale phylogenetic study should be done in the future to resolve it. The rhinocerotid association found in Wischberg is nonetheless typical of the MN1 biozone, which results from a faunal renewal occurring just before the end of the Oligocene.

New data on Amynodontidae (Mammalia, Perissodactyla) from Eastern Europe: Phylogenetic and palaeobiogeographic implications around the Eocene-Oligocene transition.

Amynodontidae is a family of Rhinocerotoidea (Mammalia, Perissodactyla) known from the late Early Eocene to the latest Oligocene, in North America and Eurasia. European Amynodontidae are very rare, and all remains belong almost exclusively to a single post-Grande Coupure genus from the Oligocene, Cadurcotherium. The "Grande Coupure" defines an extinctions and dispersal-generated originations event in Europe that is nearly contemporaneous with the Eocene-Oligocene transition. Perissodactyls are one of the major groups affected by this event: Palaeotheriidae went almost extinct during this crisis, whereas Rhinocerotidae appeared for the first time in Europe. Study of fossiliferous Eastern-European localities from this age is crucial for the understanding of this crisis. We report here three new localities of Amynodontidae in Eastern Europe. Two of them are dated from the Eocene (Morlaca, Romania; Dorog, Hungary), whereas the other is either Late Eocene or Early Oligocene (Dobârca, Romania). The skull from this latter locality belongs unexpectedly to the same individual as a previously described mandible attributed to "Cadurcodon" zimborensis. As a result, this specimen can be allocated to its proper locality, Dobârca, and is assigned to a new genus, Sellamynodon gen. nov. It is characterised by an extraordinary growth of the nuchal crest, a unique character among amynodontids. Along with this remarkable material from Dobârca, two specimens from another Romanian locality, Morlaca, have been recently discovered and are dated from the Late Eocene. They belong, as well as new material from Dorog (Middle Eocene, Hungary), to the genus Amynodontopsis, also found in North America. The new Hungarian material represents the earliest occurrence of Amynodontidae in Europe. New phylogenetic hypotheses of Rhinocerotoidea are proposed, including the new material presented here, and show that Amynodontidae may be closer to the polyphyletic family 'Hyracodontidae' than to Rhinocerotidae. Amynodontidae, with their deep preorbital fossa and extremely reduced premolars, display in fact a very derived condition, compared to rhinocerotids.

The record of Aplodontidae (Rodentia, Mammalia) in the Oligocene and Miocene of the Valley of Lakes (Central Mongolia) with some comments on the morphologic variability.

The present publication reports new discoveries of Oligocene and early Miocene aplodontid rodents from the Taatsiin Gol area (Valley of Lakes) in Mongolia. The fossil aplodontids recovered in this area are mainly composed of dental remains, some fragmentary jaws plus one partially preserved skull. Aplodontid rodents have been found from the early Oligocene (local biozone A) to the early middle Miocene (local biozone D). Altogether, eight taxa belonging to five genera have been identified in the investigated deposits: Ninamys arboraptus, Ninamys kazimierzi, Promeniscomys cf. sinensis, Prosciurus? mongoliensis and Prosciurus? sp. nov. in the early Oligocene; N. arboraptus, Proansomys badamae sp. nov. and Ansomyinae indet. in the late Oligocene; and Ansomys sp.1 in the early Miocene. In addition, although outside of the topic of the present special issue, one additional taxon, Ansomys sp.2, is reported from the ?middle Miocene. The material of aplodontids is usually relatively scarce in Asian localities. For the first time, with a sample size of 81 specimens, the material from Central Mongolia (mainly from the Oligocene) now allows a more accurate description of the morphological and size variability and resolves some systematic problems. The study of these aplodontids reveals that they are more abundant and diverse in the early Oligocene and that the diversity decreases during the late Oligocene and Miocene. One hypothesis, to explain the opposite diversity trend observed previously for sciurids in the same region, is that both Sciuromorpha families might have competed for the same resources from the early Oligocene to the middle Miocene in Central Mongolia.

Cricetidae (Rodentia, Mammalia) from the Oligocene of the Valley of Lakes (Mongolia): the genera Aralocricetodon, Eocricetodon, Bagacricetodon, Witenia and Paracricetodon.

We describe the remains of Aralocricetodon Bendukidze, 1993; Bagacricetodon Gomes Rodrigues et al., 2012; Eocricetodon Wang, 2007; Witenia de Bruijn et al., 2003 and Paracricetodon Schaub, 1925 (Cricetidae, Rodentia, Mammalia) from the Taatsiin Gol and Taatsiin Tsagaan Nuur areas (Mongolia). The studied material (comprising 128 teeth) stems from 9 localities and 23 fossil layers spanning a time interval from ∼33 to ∼24 Ma (early to late Oligocene) and covering the biozones from A to C1. The general dental pattern between the species from the early and late Oligocene differed. The occlusal pattern of the molars was more complicated and the crowns were lower during the early versus late Oligocene. This indicates a change in diet towards more abrasive plants. Several of the studied species were common in both Europe and Asia Minor during the early Oligocene. The species collected from the late Oligocene have also been recorded in Kazakhstan and China. This indicates an interesting biogeographical pattern that merits future study.

Oligocene stratigraphy across the Eocene and Miocene boundaries in the Valley of Lakes (Mongolia).

Cenozoic sediments of the Taatsiin Gol and TaatsiinTsagaan Nuur area are rich in fossils that provide unique evidence of mammal evolution in Mongolia. The strata are intercalated with basalt flows. 40Ar/39Ar data of the basalts frame the time of sediment deposition and mammal evolution and enable a composite age chronology for the studied area. We investigated 20 geological sections and 6 fossil localities of Oligocene and early Miocene deposits from this region. Seventy fossil beds yielded more than 19,000 mammal fossils. This huge collection encompasses 175 mammal species: 50% Rodentia, 13% Eulipotyphla and Didelphomorphia, and 12% Lagomorpha. The remaining 25% of species are distributed among herbivorous and carnivorous large mammals. The representation of lower vertebrates and gastropods is comparatively poor. Several hundred SEM images illustrate the diversity of Marsupialia, Eulipotyphla, and Rodentia dentition and give insight into small mammal evolution in Mongolia during the Oligocene and early Miocene. This dataset, the radiometric ages of basalt I (∼31.5 Ma) and basalt II (∼27 Ma), and the magnetostratigraphic data provide ages of mammal assemblages and time ranges of the Mongolian biozones: letter zone A ranges from ∼33 to ∼31.5 Ma, letter zone B from ∼31.5 to ∼28 Ma, letter zone C from ∼28 to 25.6 Ma, letter zone C1 from 25.6 to 24 Ma, letter zone C1-D from 24 to ∼23 Ma, and letter zone D from ∼23 to ∼21 Ma.

Oligocene and early Miocene mammal biostratigraphy of the Valley of Lakes in Mongolia.

The Taatsiin Gol Basin in Mongolia is a key area for understanding the evolution and dispersal of Central Asian mammal faunas during the Oligocene and early Miocene. After two decades of intense fieldwork, the area is extraordinarily well sampled and taxonomically well studied, yielding a large dataset of 19,042 specimens from 60 samples. The specimens represent 176 species-level and 99 genus-level taxa comprising 135 small mammal species and 47 large mammals. A detailed lithostratigraphy and new magnetostratigraphic and radiometric datings provide an excellent frame for these biotic data. Therefore, we test and evaluate the informal biozonation scheme that has been traditionally used for biostratigraphic correlations within the basin. Based on the analysis of the huge dataset, a formalised biostratigraphic scheme is proposed. It comprises the Cricetops dormitor Taxon Range Zone (Rupelian), subdivided into the Allosminthus khandae Taxon Range Subzone and the Huangomys frequens Abundance Subzone, the Amphechinus taatsiingolensis Abundance Zone (early Chattian), the Amphechinus major Taxon Range Zone (late Chattian), subdivided into the Yindirtemys deflexus Abundance Subzone and the Upper Amphechinus major T. R. Z., and the Tachyoryctoides kokonorensis Taxon Range Zone (Aquitanian). In statistical analyses, samples attributed to these biozones form distinct clusters, indicating that each biozone was also characterised by a distinct faunal type.

Evolution of the genus Eucricetodon (Rodentia, Mammalia) from the Valley of Lakes (Mongolia): a taxonomical description and update on the stratigraphical distribution.

The Oligocene fossil deposits from Valley of Lakes in Central Mongolia have provided a wealth of rodent fossils. Among these, cricetids are a very important part. To date, only the Miocene genera have been described in detail. Here, we focus on the Oligocene genus Eucricetodon from this region. Eucricetodontinae are the most abundant fossils in the Oligocene Valley of Lakes faunas. The present study consists of the description of five species of cricetid rodents from 43 localities ranging in age from the early Oligocene to the early-late Oligocene. In addition to Eucricetodon asiaticus described in Mongolia in 1923, we have found Eucricetodon bagus and Eucricetodon jilantaiensis that were described from Nei Mongol and Eucricetodon occidentalis discovered in Kazakhstan. This taxonomical study provides new information regarding the evolution of the Cricetidae in Central and Eastern Asia during the Oligocene and, more particularly, regarding their phylogenetic relationships and the evolutionary trends.

A new species of Argyromys (Rodentia, Mammalia) from the Oligocene of the Valley of Lakes (Mongolia): Its importance for palaeobiogeographical homogeneity across Mongolia, China and Kazakhstan.

We describe a new species of Rodentia (Mammalia), Argyromys cicigei sp. nov. from Toglorhoi (fossil bed TGW-A/2a) in Mongolia and Ulantatal (fossil beds UTL 1 and UTL 7) in China. Its tooth morphology differs from the type species Argyromys aralensis from Akespe in Kazakhstan by smaller size and simpler structures. Argyromys has been assigned in different families of Muroidea, such as Tachyoryctoididae and Spalacidae. However, the presence of common characters indicates a closer relationship of Argyromys with the genera of Cricetidae s.l. (subfamilies Eucricetodontinae; Cricetopinae; Cricetodontinae and Gobicricetodontinae among others) from Asia than with the earliest representatives of Spalacidae or the endemic Tachyoryctoididae. Argyromys cicigei sp. nov. possesses a simple anterocone and anteroconid in the upper and lower first molars, respectively, which is characteristic for Cricetidae s.l. It has a flat occlusal surface in worn specimens; weakly-developed posterolophs; an oblique protolophule and metaloph on the upper molars and it lacks a labial anterolophid on the m1. These traits are also typical of the Oligocene genera Aralocricetodon and Plesiodipus, included in the subfamilies Cricetodontinae and Gobicricetodontinae respectively. The cladistic analysis performed here supports this hypothesis. The clade formed by Argyromys species is grouped with other cricetid taxa (s.l). Spalacids, however, form a different clade, as do the tachyoryctoids. Previous authors state that the Aral Formation (Kazakhstan) should be dated to the Oligocene instead of the Miocene, based on the presence of several taxa. The finds of Argyromys in both regions supports the statement that they are closer in age than previously thought. The occurrence of Argyromys in Kazakhstan, Mongolia and China evidences the biogeographic unity of the Central Asian bioprovince during the Oligocene.

Stepwise onset of the Icehouse world and its impact on Oligo-Miocene Central Asian mammals.

Central Asia is a key area to study the impact of Cenozoic climate cooling on continental ecosystems. One of the best places to search for rather continuous paleontological records is the Valley of Lakes in Mongolia with its outstandingly fossil-rich Oligocene and Miocene terrestrial sediments. Here, we investigate the response by mammal communities during the early stage of Earth's icehouse climate in Central Asia. Based on statistical analyses of occurrence and abundance data of 18608 specimens representing 175 mammal species and geochemical (carbon isotopes) and geophysical (magnetic susceptibility) data we link shifts in diversities with major climatic variations. Our data document for the first time that the post-Eocene aridification of Central Asia happened in several steps, was interrupted by short episodes of increased precipitation, and was not a gradual process. We show that the timing of the major turnovers in Oligocene mammal communities is tightly linked with global climate events rather than slow tectonics processes. The most severe decline of up 48% of total diversity is related to aridification during the maximum of the Late Oligocene Warming at 25 Ma. Its magnitude was distinctly larger than the community turnover linked to the mid-Oligocene Glacial Maximum.

Testing for Depéret's Rule (Body Size Increase) in Mammals using Combined Extinct and Extant Data.

Whether or not evolutionary lineages in general show a tendency to increase in body size has often been discussed. This tendency has been dubbed "Cope's rule" but because Cope never hypothesized it, we suggest renaming it after Depéret, who formulated it clearly in 1907. Depéret's rule has traditionally been studied using fossil data, but more recently a number of studies have used present-day species. While several paleontological studies of Cenozoic placental mammals have found support for increasing body size, most studies of extant placentals have failed to detect such a trend. Here, we present a method to combine information from present-day species with fossil data in a Bayesian phylogenetic framework. We apply the method to body mass estimates of a large number of extant and extinct mammal species, and find strong support for Depéret's rule. The tendency for size increase appears to be driven not by evolution toward larger size in established species, but by processes related to the emergence of new species. Our analysis shows that complementary data from extant and extinct species can greatly improve inference of macroevolutionary processes.

Multi-proxy approach detects heterogeneous habitats for primates during the Miocene climatic optimum in Central Europe.

The present study attempts to characterize the environmental conditions that prevailed along the western shores of the Central Paratethys and its hinterland during the early middle Miocene at the same time t primates reached their peak in species diversity in Central Europe. Based on faunal structure (using cenograms), paleotemperature reconstruction (using cricetid diversity), and dietary reconstruction of ruminants (using molar micro-wear analyses), four faunal assemblages are used to characterize the regional environmental context. The cenograms for Göriach and Devínska Novà Ves Zapfe's fissure site support the presence of mosaic environments with open areas under rather humid conditions. This is also supported by the dental micro-wear analyses of ruminants. The species of Palaeomerycidae were most probably the only predominant browsers. Surprisingly, the three cervids, Dicrocerus, Heteroprox, and Euprox, were highly involved in grazing. Pseudoeotragus seegrabensis was likely a generalist and the two specimens assigned to the second bovid, Eotragus clavatus, were browsers. The two species of tragulids plot between fruit browsers and generalists. Moreover, paleotemperatures based on cricetid diversity estimate mean annual temperature at about 18 °C with potential high seasonal variations. These data support the predominance of mosaic landscapes along the western shores of the Central Paratethys and its hinterland during the Miocene Climatic Optimum as primates reach a peak in species diversity. This result lends credence to the hypothesis that environmental heterogeneity favours radiation among mammals, and that the specific environmental context of the Central Paratethys western border might explain the high diversity of the middle Miocene primates.

Diversity trends in Neogene European ungulates and rodents: large-scale comparisons and perspectives.

The ungulate and rodent fossil records are often used independently to understand mammalian evolutionary history. Few studies have directly compared both records over long time periods and large geographic areas. Here, we compiled two datasets of European fossil localities containing rodents and/or ungulates over 20 My (Early Miocene-Early Pliocene) and processed the data with the same methodology. We counted the raw diversity and calculated a measure of evenness (Pielou's index). After controlling for potential biases on diversity estimators, we identify the evenness index as a more reliable estimator bringing interesting insights into the way both mammal groups are structured by biotic or abiotic factors. In this study, we consider that an uneven distribution of the species richness among families, when only some families successfully diversify within the "continental-scale community", represents a lower adaptability of this community to the environmental context. Pielou's index is used to estimate this adaptability through time. The responses of ungulates and rodents to environmental changes are very divergent, especially facing the climatic changes known since the Middle Miocene. The general patterns suggest that rodent broad-scale assemblages are affected by all kinds of perturbations, even short biotic and abiotic events, but show a better adaptability when facing long-term abiotic changes. Unlike rodents, the ungulate assemblages show more stability in periods of relative environmental stability but show less adaptability to long-term climatic changes. Life-history traits of mammals can help explain patterns of diversity and biogeography at different spatial scales and may probably partly explain the opposite patterns evidenced here.