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The name “Cetartiodactyla” was proposed in 1997 to reflect the molecular data that suggested that Cetacea is closely related to Artiodactyla. Since then, that taxon has spread in popularity, even outside the scientific literature. However, the implications of the name are confusing, because Cetacea and Artiodactyla are not sister-taxa. Instead, the evidence clearly shows that cetaceans are a group embedded within Artiodactyla, not a sister-taxon of equal rank. It has long been accepted practice that systematists do not modify the names of higher groups when new subgroups are added to them. For example, Owen’s original concept of Artiodactyla did not change its name when more and more disparate taxa were added to it. Dinosauria did not become “Avedinosauria” when it became clear that birds are a subgroup of dinosaurs, nor did Reptilia become “Avereptilia”. In the interests of taxonomic priority and stability, and especially because the name is inherently misleading, we recommend that the name “Cetartiodactyla” be abandoned. If one wishes to make a reference to the fact that whales are now considered to be a subgroup of artiodactyls, they could be referred to informally as “whales and other artiodactyls” or “whales and terrestrial artiodactyls” without using a formal taxonomic name that is confusing and misleading.

Few pathological cases have been identified in fossils of small mammals. Here we report the most ancient paleopathological evidence identified in a lagomorph (Ochotonidae, middle Miocene). The tibiofibular bone was macro- and microscopically (μCT) inspected to provide a diagnosis, an etiology, and its possible relationship with the individual's cause of death. Osteogenesis (reactive bone growth) and osteolysis, processes related with neoplasms and infections, are identified in the abnormal bony region. Its location (juxta-articular) and morphology allow us to identify it as a joint infection (septic arthritis) consequential of a violent mechanism, such as a bite. Both the origin of bone accumulation (avian pellets) and the poor vital state of the specimen (with a joint infection) point to predation as the most probable cause of death. Up to now, lagomorph paleopathologies had only been described in insular populations, and the present one is the first evidence in a mainland specimen.

During recent years, knowledge about crocodyliform diversity of the uppermost Cretaceous from Europe has been substantially improved. Palaeontological efforts have also been focused on microvertebrate diversity and its palaeoecological implications. Isolated crocodylomorph teeth are, by far, one of the most frequently recovered elements in microvertebrate samples. In the present paper, morphological features of crocodylomorph teeth collected throughout the complete Maastrichtian series of the southern Pyrenean basin (north-eastern Spain), together with several mandibular remains, are described and analysed. Teeth were grouped in morphotypes and their taxonomic significance is discussed. The results highlight a diverse crocodylomorph assemblage in this area throughout the Maastrichtian. In addition, feeding habits and environmental preferences are inferred for the identified taxa according to dental features, occurrences and taphonomy.

The Miocene tragulid species from Siwalik sediments in Asia have been repeatedly studied since Lydekker erected them in 1876. Here, we describe and interpret new dental material of Tragulidae colle...

We would like to thank all the curators, collection manager, and scientists who helped us allowed us to study and provided access to material: C. Argot and G. Billet (Muséum national d’Histoire naturelle, Paris), D. Geraads (Museum national d’Histoire Naturelle) for providing the Sivatherium specimen, E. Robert (Université Claude Bernard, Lyon 1), D. Berthet (Musée des Confluences Lyon), M. Orliac (Université Montpellier 2) for providing the bony labyrinth of Bachitherium, F. Duranthon and Y. Laurent (Muséum d’histoire naturelle, Toulouse), S. Legal, P. Coster, C. Balm, O. Maridet, O. Lapauze and J. Tissier (Parc Naturel Régional du Luberon and excavation team Murs Project), A. de Perthuie for accessing his private collection, Christiane Zeitler, R. Ziegler and E. Heizmann (Staatliches Museum für Naturkunde Stuttgart), P. Brewer, A. Garbout, and F. Ahmed (Natural History Museum, London), U. Göhlich and G. Daxner-Höck (Naturhistorisches Museum Wien), A. Van der Geer (Netherland Centre for Biodiversity Leiden), R. C. Hulbert Jr. (University of Florida, Gainesville), A. M. García Forner and P. Montoya (Museu de Geologia de la Universitat de València, Burjassot) for providing the petrosals of Birgerbohlinia, M. Pina (University of Manchester) for scanning support, and J. Morales (Museo Nacional de Ciencias Naturales, Madrid), J. Galkin, R. O’Leary, M. Hill Chase, C. Grohé and A. Gishlick (AMNH New York, USA), M. Celik (University of Queensland), and M. Scheidegger. We are also grateful to all the persons and institutions who scanned for us. Tandra Fairbanks (NMB) is thanked for her help with the English. B.M. and L.C. are grateful to the Swiss National Science Foundation for supporting this research through the projects 200021_178853 and 200021_159854/1 on the ear region evolution in ruminants. B.M. P300P2_161065 and P3P3P2_161066 on the evolution of the early ruminants. F.B. acknowledges support from a Gerstner Scholarship at the American Museum of Natural History. G.R. thanks the German Research Foundation project RO 1197/3-1. G.M., B.M., and L.C. want to thank the Museum National d’Histoire Naturelle de Paris for financing the Ast-RX-2013-051 Project. D.D.M. acknowledges R+D+I project ref. PID2020-116220GB-I00 from the Ministerio de Ciencia e Innovación/Agencia Estatal de Investigación/10.13039/501100011033/. I.S. acknowledges support by the Spanish Ministry of Science and Innovation (projects ref. PID2020-117289GB-I00 and PID2020-116220GB-I00), and the Generalitat de Catalunya (CERCA Programme). M.K. is funded by Japan Society for the Promotion of Science (KAKENHI Grant No. 19K04060). M.R. acknowledges MINECO project CGL2011-25754 for providing funding for the analysis. S.W. thanks the Strategic Priority Research Program of Chinese Academy of Sciences (XDB26000000) and the National Natural Science Foundation of China (41872001). G.S. and B.Mü. acknowledge financial support from the Swiss National Science Foundation in the frame of the R’equip initiative (316030_133802). Publisher Copyright: © 2022, The Author(s). Extrinsic and intrinsic factors impact diversity. On deep-time scales, the extrinsic impact of climate and geology are crucial, but poorly understood. Here, we use the inner ear morphology of ruminant artiodactyls to test for a deep-time correlation between a low adaptive anatomical structure and both extrinsic and intrinsic variables. We apply geometric morphometric analyses in a phylogenetic frame to X-ray computed tomographic data from 191 ruminant species. Contrasting results across ruminant clades show that neutral evolutionary processes over time may strongly influence the evolution of inner ear morphology. Extant, ecologically diversified clades increase their evolutionary rate with decreasing Cenozoic global temperatures. Evolutionary rate peaks with the colonization of new continents. Simultaneously, ecologically restricted clades show declining or unchanged rates. These results suggest that both climate and paleogeography produced heterogeneous environments, which likely facilitated Cervidae and Bovidae diversification and exemplifies the effect of extrinsic and intrinsic factors on evolution in ruminants. publishersversion published

In Ruminantia, the lacrimal bone forms a considerable part of the facial skeleton, and the morphology of its facial facet is highly variable when compared to other mammals. In this study, we quantify the species-specific variability in size and shape of the lacrimal facial facet in species of Cervidae (deer) and relate it to systematics and various aspects of their ecology and behavior. We sampled 143 skull specimens from 10 genera; 12 Moschus and 3 Tragulus specimens were used as outgroups. We find that size and shape of the lacrimal facial facet allow differentiating most species analyzed here, except for Mazama gouazoubira and Capreolus capreolus. Size and shape of the lacrimal facial facet vary widely across Cervidae regardless of their systematic relationships, ecology or behavior. Thus, we could not detect a unique signature of adaptational criteria in lacrimal morphology. Our data indicate that the lacrimal facial facet scales allometrically with skull size, in particular, the lacrimojugal length scales positively and the lacrimomaxillar length scales negatively. However, correlation analyses did not reveal any differences in the integration of the lacrimal bone with any specific skull module in any of the species compared. Lastly, we could not ascertain any correlation between the size and position of the preorbital depression with the size and shape of the lacrimal facial facet. We conclude that the lacrimal facial facet is highly flexible and may rapidly adjust to its surrounding bones. Its allometric growth appears to be an example of exaptation: changes in size and shape in the context of the increase of the skull length provide lacrimal contacts, in particular, a lacrimojugal one, which may serve to reduce mechanical loads resulting from increasingly larger antlers in large cervids.

In the Pleistocene faunas of the island of Crete, Cervidae was one of the most abundant taxa. Respective species vary in body size, including dwarfs, and skeletal morphology; however, the number of species and the identity of the mainland ancestor(s) are still debated. In this paper, we morphologically and morpho­metrically describe and analyze eight skulls of Cretan deer from a so far little known fossil site near Gerani, Rethymnon, Greece. The recorded character suite allows for affiliation to dwarfed Candiacervus Kuss, 1975, Candiacervus ropalophorus de Vos, 1984 and C.reumeri van der Geer, 2018. It comprises previously unknown unique traits, some of them hinting to sexual dimorphism. Comparisons of the Candiacervus skulls presented here with those of cervids belonging to Megalocerotini Brooke, 1828, s.s. and s.l. stress certain similarities; yet more material is needed to reconstruct Candiacervus’ phylo­genetic position. The newly detected craniodental specifics allow for more insights into island adaptation of Candiacervus; at the same time, they blur the morphological heritage of their mainland ancestors.

Mouse-deer, represented by the extant genera Tragulus, Moschiola and Hyemoschus, are among the smallest living ruminants. Today these animals are exclusively distributed in the Indo-Malayan and Afrotropical regions. For the past sixty years, the taxonomic classification of mouse-deer included the species H. aquaticus, T./M. meminna, T. kanchil/T. javanicus, and T. javanicus/T. napu, and these species names were therefore used widely to label material stored in scientific collections. However, recent revisions based on broad phenomic analyses of specimens from collections in Asia, Europe, and North America indicate that there are in fact seven species in Tragulus and three (or four) in Moschiola, and thus challenge works with collection specimens named based on the older classification. This paper presents the results from an independent, multivariate morphometrical analysis of a small sample set of 43 Tragulus and Moschiola skulls kept in public collections in Basel, Munich, Stuttgart, and Vienna that was conducted to explore character diversity. Our results revealed three morphometric clusters in Tragulus and two in Moschiola that concur with some of the recently revised species concepts, i.e. T. napu ('our' Borneo, Sumatra, and Sunda Islands specimens), T. kanchil ('our' Sumatra and Thailand specimens), T. javanicus ('our' Java specimens), M. indica ('our' India specimens) and M. meminna/M. kathygre ('our' Sri Lanka specimens). To assist keepers of osteological collections with the identification of tragulids based on the most recent taxonomy, we have assembled a chart providing revised taxonomic assessments of all 43 specimens included in our analysis. In addition, we have quantified a hitherto undocumented difference in the neurocranium geometry among Asian tragulids. Individuals with a globose neurocranium are shown to belong to T. javanicus and T. kanchil, while those possessing a non-globose neurocranium are in Moschiola spp. and T. napu.

The Faluns Auger quarry (Contres, France) is famous for its Langhian mammal fauna (MN5) and tragulid remains have been identified as Dorcatherium sp. New excavations provided a fragmented skull of a tragulid, dental remains, and a metapodium which we describe here. Based on morphological and morphometrical characters, these specimens are attributed to Dorcatherium crassum. CT-scans provide insight in the petrosal bone and bony labyrinth. We prove intra-population variability of the p4 and intra-specific variability of the bony labyrinth. Nevertheless, we can demonstrate that the bony labyrinth is useful for systematics. Thus, we confirm the presence of D. crassum in the Pontlevoy–Thenay and Savigne-sur-Lathan Faluns Basins as reported already prior to our study. Yet, we cannot confirm evidence for Dorcatherium naui. D. naui specimens reported from these basins may belong to another species. D. naui should be absent from the Faluns before MN7, such as in the rest of Europe.

The identification and interpretation of paleopathological findings in the dentition of mammals have the potential to shed light on disturbed development and stress conditions. In combination with knowledge about the timing of dental development, dental pathologies can provide a unique opportunity to reconstruct specific phases in the life history of fossil taxa, as well as animal-environment interactions. The dental ontogeny of the extinct rhinoceros Prosantorhinus germanicus from the Miocene is well-known from an exceptionally preserved sample of juvenile dentaries. Two of the represented juvenile individuals each revealed a dental anomaly, which were analyzed macroscopically and via X-ray tomography in this study. One specimen documents decreased dental wear of deciduous cheek teeth, mandibular bone resorption as well as abnormal cemental deposition. At an early age, the extinct juvenile rhinoceros probably was affected by a gingival infection resulting from an accumulation of bacterial plaque. This led to a chronic gingivitis and eventually may have caused periodontitis. Furthermore, it suffered from inflammation-induced hypercementosis. Another specimen revealed an abnormal reduction in thickness of enamel in the deciduous cheek tooth. The enamel hypoplasia indicates stress experienced during the prenatal period. During a phase of environmental stress (e.g. increased fluoride exposure) or physiological stress (e.g. malnutrition) the function of the secretory ameloblasts was disrupted in the developing embryo. The present work shows that the analysis of dental paleopathologies is a useful means for retrospective assessment of specific phases in the life history of extinct animals. This adds to the picture of the living conditions of the extinct rhinoceros P.germanicus.

Número especial llamado: "Progress in Echinoderm Paleobiology", Moroccodiscus smithi represents a new cyclocystoid genus and species based on moldic specimens from the Middle Ordovician Taddrist Formation (Darriwilian) of SE Morocco. This represents the earliest articulated member of the Cyclocystoidea and is the first complete cyclocystoid described from the Ordovician of Gondwana, as well as the first cyclocystoid ever recorded from Africa. The anatomy and morphology of this new species were studied using a combination of conventional paleontological methods and nondestructive X-ray computed tomography. Because Moroccodiscus differs from other cyclocystoids, in particular by lacking cupules attached to the marginal ossicles, it is assigned to the new family Moroccodiscidae. This new taxon illustrates the relatively poorly known early diversification of these enigmatic extinct echinoderms and sheds light on the mode of life of cyclocystoids, including injuries to plate circlets during early ontogeny and folding of these disk-like specimens at the time of death. The overall thecal shape was very similar in cyclocystoids and many domal edrioasteroids, probably because they were both sessile or attached, benthic, suspension feeders. However, many oral surface, ambulacral, and marginal ring features had become very different, indicating that these two groups had either converged because of similar life modes or were only distantly related sister groups., Bavarian State Collection of Palaeontology and Geology, Alemania, Department of Earth and Environmental Sciences, Division of Palaeontology and Geobiology, Ludwig-Maximilians-Universität München, Alemania, GeoBio-Center, Alemania, Department of Geological Sciences, Jackson School of Geosciences, University of Texas, Estados Unidos, Laboratoire de Géologie de Lyon: Terre, Planète, Environnement, Unitè Mixte de Recherche, Université Lyon 1, 2, Francia, Laboratoire de Géologie de Lyon: Terre, Planète, Environnement, Unitè Mixte de Recherche, Centre National de la Recherche Scientifique, Francia, Unidad de Zaragoza, Instituto Geológico y Minero de España, España

Amphimoschus is an extinct Eurasian ruminant genus, mostly recorded in Europe, without a close living relative and, hence, an unknown systematic position. This genus is known from around 50 localities from the late early to the middle Miocene. Two species were described during 180 years, but since their first description during the late 19th century and early 20th century, hardly any detailed taxonomic work has been done on the genus. Over the years, extensive collecting and excavating activities have enriched collections with more and more complete material of this still rare and enigmatic animal. Most interestingly, a number of skull remains have been unearthed and are promising in terms of providing phylogenetic information. In the present paper, we describe cranial material, the bony labyrinth, the dentition through 780 teeth and five skulls from different ontogenetic stages. We cannot find a clear morphometric distinction between the supposedly smaller and older species Amphimoschus artenensis and the supposedly younger and larger species A. ponteleviensis. Accordingly, we have no reason to retain the two species and propose, following the principle of priority (ICZN chapter 6 article 23), that only A. ponteleviensis Bourgeois, 1873 is valid. Our studies on the ontogenetic variation of Amphimoschus does reveal that the sagittal crest may increase in size and a supraorbital ridge may appear with age. Despite the abundant material, the family affiliation is still uncertain.

Trilobites, extinct arthropods that dominated the faunas of the Palaeozoic, since their appearance c 523 million years ago, were equipped with elaborate compound eyes. While most of them possessed apposition compound eyes (in trilobites called holochroal eyes), comparable to the compound eyes of many diurnal crustaceans and insects living today, trilobites of the suborder Phacopina developed atypical large eyes with wide lenses and wide interspaces in between (schizochroal eyes). Here, we show that these compound eyes are highly sophisticated systems—hyper-compound eyes hiding an individual compound eye below each of the big lenses. Thus, each of the phacopid compound eyes comprises several tens, in cases even hundreds of small compound eye systems composing a single visual surface. We discuss their development, phylogenetic position of this hyper-compound eye, and its neuronal infrastructure. A hyper-compound eye in this form is unique in the animal realm.

Deer (Cervidae) have a long evolutionary history dating back to the Early Miocene, around 19 million years ago. The best known fossils to document this history belong to European taxa, which all bear cranial appendages more or less similar to today's deer antlers. Despite the good fossil record, relationships of the earliest stem deer and earliest crown deer are much debated. This hampers precise calibration against the independent evidence of the fossil record in molecular clock analyses. While much has been written on the Early and Middle Miocene deer, only two phylogenetic analyses have been performed on these taxa to date mostly based on cranial appendage characters. Because the petrosal bone and bony labyrinth have been shown to be relevant for phylogeny in ruminants, we describe for the first time these elements for four iconic early cervids from Europe (Procervulus dichotomus, Heteroprox larteti, Dicrocerus elegans and Euprox furcatus) and include them in a phylogenetic analysis based on the ear region exclusively. The analysis recovered E. furcatus in a sister position to the living red deer (Cervus elaphus). Further, it placed D. elegans in a sister position to Euprox+Cervus and a clade Procervulinae that includes P. dichotomus and H. larteti, in sister position to all other deer. The inclusion of E. furcatus in crown Cervidae, which was previously suggested based on antler morphology, cannot be ruled out here but needs a more comprehensive comparison to other crown deer to be confirmed. J. Morphol. 277:1329-1338, 2016. (c) 2016 Wiley-Blackwell.

Deer (Cervidae) have a long evolutionary history dating back to the Early Miocene, around 19 million years ago. The best known fossils to document this history belong to European taxa, which all bear cranial appendages more or less similar to today's deer antlers. Despite the good fossil record, relationships of the earliest stem deer and earliest crown deer are much debated. This hampers precise calibration against the independent evidence of the fossil record in molecular clock analyses. While much has been written on the Early and Middle Miocene deer, only two phylogenetic analyses have been performed on these taxa to date mostly based on cranial appendage characters. Because the petrosal bone and bony labyrinth have been shown to be relevant for phylogeny in ruminants, we describe for the first time these elements for four iconic early cervids from Europe (Procervulus dichotomus, Heteroprox larteti, Dicrocerus elegans and Euprox furcatus) and include them in a phylogenetic analysis based on the ear region exclusively. The analysis recovered E. furcatus in a sister position to the living red deer (Cervus elaphus). Further, it placed D. elegans in a sister position to Euprox + Cervus and a clade Procervulinae that includes P. dichotomus and H. larteti, in sister position to all other deer. The inclusion of E. furcatus in crown Cervidae, which was previously suggested based on antler morphology, cannot be ruled out here but needs a more comprehensive comparison to other crown deer to be confirmed. J. Morphol. 277:1329-1338, 2016. © 2016 Wiley Periodicals, Inc.

Fossil evidence of complete sequences of dental ontogeny in extinct mammals is rare but contains valuable information on the animal’s physiology, life history, and individual age. Here, we analyzed an exceptionally high number of juvenile dentaries at different developmental stages including highly fragile tooth germs of the extinct rhinoceros Prosantorhinus germanicus from the Miocene fossil lagerstatte Sandelzhausen in Germany. We used dental wear stages, eruption stages, and tooth germ development in order to reconstruct the tooth replacement pattern for P. germanicus. The results allow for the distinction of 11 dental eruption stages and document a tooth eruption sequence of (d2, d3), (d1, d4), m1, m2, p2, p3, p4, m3; a pattern identical to that reported for the extant African rhinoceros, Diceros bicornis. Moreover, our findings indicate that P. germanicus falls into the life history category of slow-growing, long-living mammals. The dental eruption stages of the fossil rhinoceros were correlated with data of living rhinoceroses in order to gain insight into the age-at-death distribution of P. germanicus at Sandelzhausen. The juvenile mortality profile of P. germanicus shows a trend of selective mortality at an inferred age range of about 3 months to 3 years. As this age range represents a life phase of increased natural risk of mortality, our findings indicate a gradual accumulation of corpses (attritional fossil assemblage). This result supports the interpretation of a taphocenosis found at the Sandelzhausen fossil site.

Deer are an iconic group of large mammals that originated in the Early Miocene of Eurasia (ca. 19 Ma). While there is some consensus on key relationships among their members, on the basis of molecular- or morphology-based analyses, or combined approaches, many questions remain, and the bony labyrinth has shown considerable potential for the phylogenetics of this and other groups. Here we examine its shape in 29 species of living and fossil deer using 3D geometric morphometrics and cladistics. We clarify several issues of the origin and evolution of cervids. Our results give new age estimates at different nodes of the tree and provide for the first time a clear distinction of stem and crown Cervidae. We unambiguously attribute the fossil Euprox furcatus (13.8 Ma) to crown Cervidae, pushing back the origin of crown deer to (at least) 4 Ma. Furthermore, we show that Capreolinae are more variable in bony labyrinth shape than Cervinae and confirm for the first time the monophyly of the Old World Capreolinae (including the Chinese water deer Hydropotes) based on morphological characters only. Finally, we provide evidence to support the sister group relationship of Megaloceros giganteus with the fallow deer Dama.

New materials of the ornithischian dinosaur Manidens condorensis highlight a strong heterodonty between the upper and lower dentitions and reveal a novel occlusion type previously unreported in herbivorous dinosaurs. The diamond-shaped maxillary teeth have prominent cingular entolophs in a V- to Z-shaped configuration that are absent in dentary teeth. These cingular entolophs bear denticles and serrations with vertical wear that is coplanar with the apical wear facets, supporting their involvement in chewing. The separated apical and basal wear in dentary teeth is consistent with the apical and cingular wear in maxillary teeth, indicating an alternate occlusion, an orthal jaw motion, and shearing interactions between marginal and cingular edges in a double occlusion. Measurements of the length and wear area along the marginal and cingular edges indicate that the latter are functionally equivalent to adding eight teeth to a maxillary toothrow of ten, almost doubling the lengths of cutting edges and the degree of intraoral processing, while maintaining a plesiomorphic skull anatomy, an adaptation to herbivory unique in Ornithischia. Fil: Becerra, Marcos Gabriel. Museo Paleontológico Egidio Feruglio; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Pol, Diego. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Museo Paleontológico Egidio Feruglio; Argentina Fil: Rössner, Gertrud E.. Universidad de Múnich; Alemania. Bayerische Staatssammlung für Paläontologie und Geologie; Alemania Fil: Rauhut, Oliver W. M.. Bayerische Staatssammlung für Paläontologie und Geologie; Alemania. Universidad de Múnich; Alemania

Over the last decade, important progress has been made in the study of the mammal successions from Hungary and Romania. A critical review of the taxonomy of the published small mammals is provided herein, as well as some new data and an overview of the accompanying vertebrate fauna (excluding fishes) in their stratigraphic context. In addition, the first data regarding the ruminants from Matraszőlős and Felsőtarkany are presented. This contribution aims to characterize the middle to earliest late Miocene fossil record from these countries, and provide important data for the chrono(bio)stratigraphic and palaeoenvironmental studies at the European scale.

The omasum of pecoran ruminants (which is absent in tragulids) and shorter gestation periods in non-giraffid crown pecorans (as opposed to giraffids) could represent cases of key innovations that caused disparity in species diversity in extant ruminants. Literature suggests that the different ruminant groups inhabited similar niche spectra at different times, supporting the ‘increased fitness’ interpretation where a key innovation does not mainly open new niches, but allows more efficient use of existing ones. In this respect, we explored data on fossil species diversity of Afro-Eurasian ruminants from the Neogene and Quaternary. Tragulid and giraffid diversity first increased during the Early/Middle Miocene with subsequent declines, whereas bovid and cervid diversity increased distinctively. Our resulting narrative, combining digestive physiology, life history and the fossil record, thus provides an explanation for the sequence of diversity patterns in Old-World ruminants.

One of the rare records of a rich ruminant fauna of late Middle Miocene age (Sarmatian sensu stricto; 12.2–12.0 Ma) was discovered at the Gratkorn locality (Styria, Austria). It comprises, besides Micromeryx flourensianus, ?Hispanomeryx sp., Euprox furcatus, Palaeomerycidae gen. et sp. indet., and Tethytragus sp., one of the oldest records of Dorcatherium naui. Gratkorn specimens of the latter species are in metric and morphologic accordance (e.g. selenodont teeth, bicuspid p2, non-fusion of malleolus lateralis and tibia) with type material from Eppelsheim (Germany) and conspecific material from Atzelsdorf (Austria), and do not show an intermediate morphology between Late Miocene Dorcatherium naui and Middle Miocene Dorcatherium crassum, thus enforcing the clear separation of the two species. It furthermore confirms the assignation of Dorcatherium naui to a selenodont lineage (together with Dorcatherium guntianum) distinct from a bunoselenodont lineage (including Dorcatherium crassum). The record of ?Hispanomeryx sp. is the first of this genus in Central Europe. While Tethytragus sp. could also be a new bovid representative for the Sarmatian of Central Europe, Micromeryx flourensianus and Euprox furcatus are well-known taxa in the Middle Miocene of Central Europe, but comprise their first records from Styria. Morphological data from this work in combination with isotopic measurements (δ18OCO3, δ13C; Aiglstorfer et al. 2014a, this issue) indicate a niche partitioning for the ruminants from Gratkorn with subcanopy browsing (Euprox furcatus), top canopy browsing (Tethytragus sp.) and even a certain amount of frugivory (Dorcatherium naui and Micromeryx flourensianus).

Seven hitherto unpublished Dorcatherium guntianum teeth from the Early Miocene of Germany are described. Morphology and size of the teeth are documented in detail and taxonomic affiliation is assessed based on comparisons to type materials of European tragulids. The fossils represent one of the earliest European Dorcatherium records and the oldest unequivocal evidence of D. guntianum from Germany. A review of the European Dorcatherium occurrence pattern suggests that a bunoselenodont and a selenodont lineage immigrated into Europe simultaneously from the East or Southeast, and corroborates the interpretation of the genus as being diphyletic. Moreover, our data add support to immigration scenarios that have been linked with the emerging freshwater wetlands, which formed as a result of the regression in the Western Paratethys.

Gazella is one of the most species-rich genera within horned ruminants. Despite overall similarity in body size and morphology, gazelles show variability in coloration and horn morphology. Unfortunately, however, species differentiation based on these characters, or on discrete skull characters, is very difficult due to high intraspecific variability. Furthermore, most species have fragmented and allopatric distributions, so that species boundaries were hard to define in the past. Mitochondrial DNA sequences have proven useful for investigating gazelle taxonomy in recent years, but especially for old museum material, i.e. type specimens, destructive sampling is often impossible. We provide a comprehensive morphometric framework for the genus Gazella based on linear skull measurements reconciled with results from molecular phylogenetic analysis based on the largest dataset available so far. In particular for males, the skull morphology shows interspecific differences concurrent with DNA data and provides a reliable tool for species identification. Based on morphometric data we synonymize G. karamii with G. marica, and confirm the identification of the G. arabica and G. a. rueppelli type skulls from analyses of mitochondrial DNA sequences. © 2013 The Linnean Society of London

Antilopini (gazelles and their allies) are one of the most diverse but phylogenetically controversial groups of bovids. Here we provide a molecular phylogeny of this poorly understood taxon using combined analyses of mitochondrial (CYTB, COIII, 12S, 16S) and nuclear (KCAS, SPTBN1, PRKCI, MC1R, THYR) genes. We explore the influence of data partitioning and different analytical methods, including Bayesian inference, maximum likelihood and maximum parsimony, on the inferred relationships within Antilopini. We achieve increased resolution and support compared to previous analyses especially in the two most problematic parts of their tree. First, taxa commonly referred to as "gazelles" are recovered as paraphyletic, as the genus Gazella appears more closely related to the Indian blackbuck (Antilope cervicapra) than to the other two gazelle genera (Nanger and Eudorcas). Second, we recovered a strongly supported sister relationship between one of the dwarf antelopes (Ourebia) and the Antilopini subgroup Antilopina (Saiga, Gerenuk, Springbok, Blackbuck and gazelles). The assessment of the influence of taxon sampling, outgroup rooting, and data partitioning in Bayesian analyses helps explain the contradictory results of previous studies.

Gazella arabica Lichtenstein, 1827, a gazelle species currently classified as “data deficient” on the IUCN Red List, has puzzled researchers for more than a century. The type specimens have repeatedly been classified as subspecies of G. gazella, G. dorcas or a distinct species since their first description about 180 years ago. Especially the skull is causing problems, as no similar gazelle individual has ever been found. We unravel the identity of G. arabica by sequencing two mitochondrial markers from the skull and skin which constitute the G. arabica lectotype and by performing a phylogenetic analysis of the genus. The results show that the lectotype skull and skin are not derived from the same animal. They belong to two individuals representing the two monophyletic lineages within the Mountain gazelle clade, Gazella gazella. By tracing the taxonomic history of G. arabica and following the rules of the ICZN we are able to resolve the hypodigm of G. arabica.

Recently, many new (extant) mammal species have been named, mostly by raising subspecies to species rank. This is primarily a consequence of the phylogenetic species concept (PSC) that has become very popular over the last few decades. We highlight several cases of splitting and argue that much of this taxonomic inflation is artificial due to shortcomings of the PSC and unjustified reliance on insufficient morphological and/or genetic data. We particularly discourage species splitting based on gene trees inferred from mitochondrial DNA only and phenetic analyses aimed at diagnosability. Uncritical acceptance of new species creates an unnecessary burden on the conservation of biodiversity.

Cervid phylogenetics has been puzzling researchers for over 150 years. In recent decades, molecular systematics has provided new input for both the support and revision of the previous results from comparative anatomy but has led to only partial consensus. Despite all of the efforts to reach taxon-wide species sampling over the last two decades, a number of cervid species still lack molecular data because they are difficult to access in the wild. By extracting ancient DNA from museum specimens, in this study, we obtained partial mitochondrial cytochrome b gene sequences forMazama bricenii,Mazama chunyi,Muntiacus atherodes,Pudu mephistophiles, andRusa marianna, including three holotypes. These new sequences were used to enrich the existing mitochondrial DNA alignments and yielded the most taxonomically complete data set for cervids to date. Phylogenetic analyses provide new insights into the evolutionary history of these five species. However, systematic uncertainties withinMuntiacuspersist and resolving phylogenetic relationships withinPuduandMazamaremain challenging.

Dental characters have always been important in mammal taxonomy and systematics. This is primarily due to the fact that the mammalian fossil record largely consists of teeth, which show tremendous morphological diversity throughout the group and sufficient variability in size and shape to be used as major classifying traits in subordinate taxa. To describe and compare tooth crown morphologies, their individual elements are named by specified nomenclatures (see, e.g. Thenius 1989 ). These usually comprise terms for the basic cusps and cuspids referring to the descriptive tritubercular terminology (protoconus, paraconus, etc.) of Cope, 1874 , Cope, 1883 and Osborn (1888) and continuative work. They are complemented with additional terms for apomorphic elements of the individual mammalian clades. In Ruminantia a variety of nomenclatures were applied in the past to describe dental features. This paper aims to clarify the meaning and correspondence of the array of previously used terms, focusing not only on permanent cheek teeth but also on deciduous premolars. Therefore, we synonymize the nomenclature used by various authors in table form to facilitate their identification and eventually their further use in a comparative way (e.g. in a phylogenetic analysis). Moreover, we propose a set of terms, based on these previous studies, which we consider most useful for future work on ruminant teeth.

In more than 20 years of excavation at the fossil lagerstatte Sandelzhausen (Early/Middle Miocene boundary, Burdigalian/Langhian boundary, early middle MN5) a substantial amount of fossil remains of ruminants have been recovered. Currently, it is the largest recorded assemblage of ruminants from the Miocene Northern Alpine Foreland Basin. More than 1,000 teeth, almost 70 antler remains and one skull enable the identification of five ruminants, namely the tragulid Dorcatherium crassum, the palaeomerycid Germanomeryx n. g. fahlbuschi n. sp., and the cervids Lagomeryx parvulus, Lagomeryx pumilio, and Heteroprox eggeri n. sp. Lagomeryx parvulus and L. pumilio have the most extensive record yet known for these species, opening up a much more complete view of them 120 years after the discovery of the type materials. The newly established G. n. g. fahlbuschi n. sp. and H. eggeri n. sp. enlarge our knowledge on the taxonomic composition of Miocene European Ruminantia. Because of the exceptionally large number of specimens, nearly all tooth positions of all five species are documented, thereby completing hitherto partially known character sets. The investigation comprises extensive taxonomic descriptions of all species represented and an interpretation of the palaeoecology based on an analysis of the community structure. This clearly suggests a humid closed canopy forest interspersed with temporary and perennial waters and accompanying open areas. Moreover, a comparison with other, stratigraphically close Molasse Basin communities emphasizes the various peculiarities of the Sandelzhausen community (low species number, cervid-dominance, dominance of L. pumilio over L. parvulus, non-dominance of very small-sized ruminants, comparably high portion of palaeomerycids, all species being browsers, no Eotragus and no Amphimoschus). The investigation also clarifies the similarity with the communities from Undorf and Viehhausen (Germany, MN5). The deduced dynamics in community structure of the late Early and early Middle Miocene Northern Alpine Foreland Basin provides further support for the current hypothesis of a vast wetland envi- ronment under the strong influence of alternating dry and flood seasons.

The Miocene fossil lagerstatte Sandelzhausen in southern Germany ranks among the most important Neogene fossil terrestrial localities in Europe, due to its exceptional record of both macroand microvertebrates as well as invertebrates. From its discovery in 1959 until its final closure in 2001, the site has been the subject of two long-term, systematic digging campaigns (1969–1975 and 1994–2001) as well as occasional collecting by the Ludwig-MaximiliansUniversitat (LMU) together with the Bayerische Staatssammlung fur Palaontologie und Geologie (BSPG; both Munich). In addition to a few plant remains a fossil fauna comprising more than 200 taxa and 50,000 identifiable specimens (Fahlbusch 2003, and this paper Table 1) has been collected including ostracods, molluscs, and, especially, vertebrates of all groups, with an emphasis on mammals (all material housed in BSPG, Munich). Though the vertebrate remains are almost exclusively disarticulated, they represent a variety of both autochthonous and allochthonous environments and they document extensive series of ontogenetic stages and within-species variability. Scientific results derived from Sandelzhausen have been published since 1970 and, ultimately, 26 specialists involved in Sandelzhausen research came together in September 2005 at the Sandelzhausen Symposium in Mainburg to discuss the latest results on all faunistic, ecological, and genetic aspects of the fossil site (Fig. 5a, b; Anonymous 2005, Rossner 2006a, Rossner and Fahlbusch 2006). The present volume resulted from that meeting and presents new contributions to the fossil fauna of Sandelzhausen. The purpose of this paper is to give a short overview and a summary of previous work on the Sandelzhausen fossil site, to document the fieldwork and the geological context, and to discuss the age of the fauna. A German-language summary article including the most recent faunal list, details of the excavation, and the complete publication list up to 2002, including popular accounts (e.g., Fahlbusch and Liebreich 1996), can be found in Fahlbusch (2003). A partly synthetic paper on the genesis and ecology of the site compiling all available research results and new evidence about the fossil lagerstatte will be published elsewhere (in preparation).

Currently the only members of the family Suidae known from the Miocene of Myanmar and Thailand belong to the subfamilies Tetraconodontinae with the genera Tetraconodon and Sivachoerus and Suinae with the genus Propotamochoerus recorded by few specimens. Here new records of the genus Propotamochoerus from the Lower Irrawaddy Formation at Tebingan in Myanmar are documented with additional material of P. “hysudricus” and for the first time with P. wui. Though the species P. “hysudricus” from the Indian Subcontinent needs revision, its presence in Myanmar indicates faunal affinities between both areas. Propotamochoerus wui is further known from Lufeng, southern China, and indicates faunal affinities between SE Asia and southern China. The finds are consistent with a Late Miocene age of the deposits.

Ruminants made up a major component in Early and Middle Miocene European land mammal faunas. Hypotheses are tested which concern the diversity of feeding niches in Miocene ruminants and their resource partitioning. Because teeth are the basis of taxonomic work in the biota investigated here, the tooth-based mesowear method of dietary evaluation is applied. Dental wear equilibria as evaluated using mesowear analysis represent an immediate taxon-independent habitat interface, which bridges extant and Neogene communities. Two adjacent palaeoenvironments have been selected as models for testing our hypotheses, because ruminants are well recorded, the temporal window represents a long lasting and stable period before the onset of a global and regional environmental change that led to an increase of grass-dominated environments and because both environments represent fundamentally different habitat conditions but are spatially connected and represent the same climatic zone (i.e., the Molasse Basin wetland and the adjacent karstified Jurassic limestone plateau in Southern Germany). The ruminant communities of both biomes are brachydont throughout and a prevalence of browsing ruminants is found in both biomes, which is in accordance with isotope data. Both habitats, however, did also support strategies equalling those of extant mixed feeders. Among these mixed feeders, there are more grass-dominated mixed feeders in the karst habitat than there are in the wetland. The ruminant communities are thus found to apparently support a range of feeding traits, not evidenced by other ecomorphological approaches such as tooth crown height evaluation nor taxonomic uniformitarianism. The presence of these feeding types suggests that niche segregation in Lower and Middle Miocene brachydont ruminants was already fairly pronounced and reflects the hypothesised niche availability in the two palaeobiomes. The comparatively dry karstified landscape of the Jurassic limestone plateau is reflected by a higher diversity in abrasive feeders compared to the Molasse Basin wetland. As a second approach, two cervid species that co-occur in both biomes, Lagomeryx parvulus and L. pumilio have been tested in terms of differences in the abrasiveness of their diet in the two biomes. L. pumilio has a large dietary spectrum, and in the karst biome, it shows a signal equalling that of extant grass-dominated mixed feeders, while in the wetland biome it behaves as a browser as many other taxa also do. L. parvulus is shown to have a much less pronounced flexibility, as its diet consists of browse in both biomes. The coincidence of both, community structure and interspecific variability in dietary traits of the Lagomeryx species is a strong argument to consider the central hypothesis as supported, that drier environments result in more abrasion dominated dietary signals of flexible feeders, and that there is a general relationship between habitat structure, food availability, the abrasiveness of the forage, and tooth wear. The dietary variability of a species therefore seems to be a highly indicative characteristic in any ecological analysis of extant and fossil biomes and the presented approach adds rarely gained information on certain fossil members of a palaeocommunity and their potential role in the ecosystem.

Background Body size variation within clades of mammals is widespread, but the developmental and life-history mechanisms by which this variation is achieved are poorly understood, especially in extinct forms. An illustrative case study is that of the dwarfed morphotypes of Candiacervus from the Pleistocene of Crete versus the giant deer Megaloceros giganteus, both in a clade together with Dama dama among extant species. Histological analyses of long bones and teeth in a phylogenetic context have been shown to provide reliable estimates of growth and life history patterns in extant and extinct mammals. Results Similarity of bone tissue types across the eight species examined indicates a comparable mode of growth in deer, with long bones mainly possessing primary plexiform fibrolamellar bone. Low absolute growth rates characterize dwarf Candiacervus sp. II and C. ropalophorus compared to Megaloceros giganteus displaying high rates, whereas Dama dama is characterized by intermediate to low growth rates. The lowest recorded rates are those of the Miocene small stem cervid Procervulus praelucidus. Skeletal maturity estimates indicate late attainment in sampled Candiacervus and Procervulus praelucidus. Tooth cementum analysis of first molars of two senile Megaloceros giganteus specimens revealed ages of 16 and 19 years whereas two old dwarf Candiacervus specimens gave ages of 12 and 18 years. Conclusions There is a rich histological record of growth across deer species recorded in long bones and teeth, which can be used to understand ontogenetic patterns within species and phylogenetic ones across species. Growth rates sensu Sander & Tückmantel plotted against the anteroposterior bone diameter as a proxy for body mass indicate three groups: one with high growth rates including Megaloceros, Cervus, Alces, and Dama; an intermediate group with Capreolus and Muntiacus; and a group showing low growth rates, including dwarf Candiacervus and Procervulus. Dwarf Candiacervus, in an allometric context, show an extended lifespan compared to other deer of similar body size such as Mazama which has a maximum longevity of 12 years in the wild. Comparison with other clades of mammals reveals that changes in size and life history in evolution have occurred in parallel, with various modes of skeletal tissue modification. Electronic supplementary material The online version of this article (doi:10.1186/s12862-015-0295-3) contains supplementary material, which is available to authorized users.

Hambach 6C, a recently discovered middle Miocene vertebrate locality in northwestern Germany, produced the fourth occurrence of the extremely rare ruminant Orygotherium escheri. The new record is an isolated M3. In Central Europe, O. escheri is known only from lignitic deposits of zone MN5, and may have been restricted to swampy habitats. Figures and measurements of the lecto- and paralectotypes are published for the first time. The phylogenetic position of Orygotherium within the Ruminantia remains uncertain despite 160 years of research.

Dental and cranial appendage remains of Cervidae from the fossil site of Dorn-Dürkheim are studied in detail. The material mainly includes isolated teeth, isolated pedicles and antler pieces. Neither tooth rows nor complete appendages are recorded. Comparative morphology and statistics of morphometrics (principal component analysis and discriminant analysis) allow for the classification of small and large dentitions, small cranial appendages, two morphotypes of large pedicles and two morphotypes of large antlers. Possible combinations of the classified units document the sympatric occurrence of three species, namely, Procapreolus sp., Muntiacinae gen. and sp. indet., cf. Cervavitulus mimus, but the fragmentary condition of the material leads to ambiguity regarding their composition and, consequently, to a certain extent regarding the taxonomic identification. However, these remains indicate the contemporaneous occurrence of early Turolian members of the crown cervids Muntiacinae and Capreolinae and close a previous spatiotemporal gap in the European cervid record. In addition, their presence proves the progressive turnover from dichotomous-antlered muntiacines to early monopodial-antlered crown cervids from NE to SW Europe in the late Miocene. The taxonomical assignment challenges the recent hypothesis on the origin of crown Cervidae around the middle/late Miocene border since Dorn-Dürkheim cervids provide further evidence for the successive achievement of derived characters in cranial appendages of crown cervids (mediopostorbital position and backwards orientation of pedicles, coronet development, shaft development/elongation, beam development and increase in number of antler tines) in the lineage of crown cervids, which originated during the middle Miocene. Fil: Azanza, Beatriz. Universidad de Zaragoza. Facultad de Ciencias; España Fil: Rössner, Gertrud E.. Bayerische Staatssammlung Fur Palaontologie Und Geologie; Alemania Fil: Ortiz Jaureguizar, Edgardo. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina