The Miocene differentiation of old world Pecora (Mammalia)

Dental and horn core/antler morphology is discussed for some representative lower and middle Miocene pecorans in a review of the phylogeny and early zoogeography of the modern Old World families. Giraffes and bovids are more closely related than either is to deer. Andegameryx, Walangania and Teruelia are related to giraffoids plus bovoids together; Amphimoschus and Hispanomeryx are a probable sister group of bovoids; Propalaeoryx is the sister group of the giraffids plus climacoceratids. Procervulus and Lagomeryx may be closer to modern deer than are Stephanocemas and Dicrocerus but all are included in the Cervidae. The horn structure of Palaeomeryx has resemblances to giraffids, but its teeth and postcranial bones are more like cervids and it is placed in the Cervoidea. Pecorans seem to have evolved in Eurasia and there gave rise to deer. By the early Miocene some pecorans had entered Arabo‐Africa where they gave rise to giraffids and probably bovids. The relationships and zoogeography of the early bovids Eotragus, Homoiodorcas and Caprotragoides are likely to be important for later bovid history.     

The petrosal bone and bony labyrinth of early to middle Miocene European deer (Mammalia,Cervidae) reveal their phylogeny

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.     

Molecular phylogeny of musk deer: a genomic view with mitochondrial 16S rRNA and cytochrome b gene

The phylogenetic status of the infra order Pecora is controversial, even though it is supported by paleontological, morphological, and molecular evidence. We analyzed two mitochondrial genes (i.e., 16S rRNA and cytochrome b) to resolve the phylogenetic position of pecoran species, i.e., the Bovidae, Cervidae, and Moschidae endemic to the Indian subcontinent. We used phylogenetic analysis based on different algorithms, including neighbor joining, maximum parsimony, Bayesian inference, maximum likelihood, minimum evolution, median joining network, along with multidimensional scaling, and DNA word analysis. Our results established the basal position of Tragulidae and the monophyly of the infra order Pecora within the Suborder Ruminantia. Our results also demonstrated that Bovidae, Cervidae, and Moschidae are allied with the placement of musk deer as more closely related to bovids than to cervids. Molecular dating based on sequence analysis shows that the radiation of Pecora occurred during the early Oligocene and that the majority of the pecoran families radiated and dispersed rapidly during the Oligocene/Miocene transition.     

The earliest musk deer of the genus <Emphasis Type="Italic">Moschus</Emphasis> and their significance in clarifying of evolution and relationships of the family Moschidae

Moschus grandaevus Schlosser, the most ancient musk deer, is recorded from two Late Miocene localities in the south of Eastern Siberia, Olkhon Island (Lake Baikal) and Taralyk-Cher near Kyzyl (Tuva). The morphological study of the species elucidates the origin, evolution, and relationships of the genus Moschus and the entire family Moschidae. A new classification of the Moschidae is proposed.     

Prostrepsiceros and Protragelaphus (Artiodactyla,Mammalia) from the Late Miocene locality of Chomateri (Attica,Greece)

The Late Miocene fossiliferous locality of Chomateri is located close to the classic locality of Pikermi, but unfortunately its faunal context is not sufficiently known. Some fossil remains of spiral-horned antelopes from Chomateri are assigned to Prostrepsiceros rotundicornis and Protragelaphus skouzesi. The co-existence of P. rotundicornis and P. skouzesi is relatively rare in the Late Miocene, but well-documented in Pikermi. Comparisons of the Chomateri material support a middle Turolian age for the Chomateri locality and suggest a close chronological relation to the Pikermi fauna.     

First identification of Decennatherium Crusafont, 1952 (Mammalia,Ruminantia, Pecora) in the Siwaliks of Pakistan

Previously undescribed remains of a new large giraffid have been identified from the late Miocene Siwaliks Hills of the Potwar Plateau in Pakistan. This taxon is very intimately related to the late Miocene giraffid genus Decennatherium, previously only certainly identified in the Iberian Peninsula, and with some possible remains assigned to Decennatherium crusafonti from Aliäbäd (Iran). The new material collected in the Siwaliks Hills shows a high morphological similarity with the Spanish remains of the genus, especially the early Vallesian Decennatherium pachecoi. Two previously undescribed ossicone fragments from the same area are also described and identified as cf. Decennatherium. These findings represent the easternmost occurrence of the genus Decennatherium and show a late Miocene migration of the genus throughout the Iberian Peninsula and Southern Asia.     

Phylogeny and systematics of the Orycteropodidae (Mammalia,Tubulidentata)

The systematics of the order Tubulidentata is poorly known. Its phylogeny has never been thoroughly analysed and only a single review has ever been performed, which was over 30 years ago. This situation has hampered palaeoecological and palaeobiogeographical studies of these Neogene mammals. The present revision of the Orycteropodidae deals with the phylogeny and systematics of all African and Eurasian species over the last 20 Myr. The first comprehensive cladistic analysis of the family is presented here. The results of this analysis, based on 39 coded morphological characters, supplemented by non‐coded features taken from all over the skeleton, was used to reconstruct the phylogeny of the order Tubulidentata. Two distinct lineages within the genus Orycteropus are recognized and characterized. The new genus Amphiorycteropus is subsequently created, in order to harmonize taxonomy and phylogeny. The fossil genera Leptorycteropus and Myorycteropus are validated, bringing the number of genera in the order Tubulidentata to four. Moreover, within the family Orycteropodidae, the number of confirmed species is now 14. The outcome of this study allows us to propose a consistent palaeobiogeographical scenario for aardvarks. Finally, this revision represents the most comprehensive work on the evolutionary history of the order Tubulidentata to date, and provides a new framework for future studies. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 649–702.     

Palaeoreas,Majoreas, and Stryfnotherium gen. nov. (Mammalia: Artiodactyla: Bovidae) from the Late Miocene of Greece

A new comparative study of the two crania, LGPUT RZO-03 and LGPUT RZO-68 attributed to Palaeoreas zouavei Bouvrain, 1980 from the Late Miocene of Axios Valley, Greece reveals great morphological and metric differences between them, which require the exclusion of LGPUT RZO-68 from this species and even from Palaeoreas Gaudry, 1861. This decision leaves unsupported many diagnostic features of P. zouavei applied by previous authors and along with recent data from Bulgaria put in doubt the validity of P. zouavei as a whole. As a result, the genus Palaeoreas is re-defined as monotypic and the cranial morphometric changes of P. lindermayeri (Wagner, 1848) are discussed on a temporal basis. An updated taxonomic analysis confirms the presence of P. lindermayeri in the Late Turolian mammal assemblage of Dytiko-1 (Axios Valley, Greece) but rejects its occurrence at Dytiko-2. Instead, a re-examination of old cranial specimens and a study of new ones suggest the presence in both sites of Majoreas Kostopoulos, 2004 a Late Miocene taxon previously mostly known from the eastern banks of the Aegean Sea. A reassessment of the LGPUT RZO-68 cranium, and its comparison with Palaeoreas and Criotherium Forsyth-Major, 1891, suggests that it deserves taxonomic distinction at genus and species levels: Stryfnotherium exophthalmon.     

Boselaphines (Artiodactyla, Ruminantia, Bovidae) from the Middle Siwaliks of Hasnot, Pakistan

In this paper, boselaphine material from several localities in the area of the Hasnot Pakistan, is described, identified, and discussed. Four species that belong to three different genera of the tribe Boselaphini have been found: Selenoportax vexillarius, S. lydekkeri, Pachyportax latidens and Eotragus sp. Eotragus sp. is reported for the first time from the Hasnot and consequently from other Upper Middle Siwalik sediments of Pakistan and equivalent strata of the world, extending the range of the genus from the Lower to the Middle Siwaliks. Reviewing the Siwaliks’ Selenoportax species, S. dhokpathanensis Akhtar and S. tatrotensis Akhtar are synonymized with S. lydekkeri and S. vexillarius, respectively.     

Molecular phylogeny of caprines (Bovidae, Antilopinae): the question of their origin and diversification during the Miocene

Caprines include all bovids related to sheep and goat. The composition of the group is controversial and inter-generic relationships have been widely debated. Here, we analysed 2469 characters draw from three distinct molecular markers, i.e. two mitochondrial genes (cytochrome b and 12S rRNA) and one nuclear fragment (exon 4 of the κ -casein gene). The taxonomic sampling includes all genera putatively described as caprines, as well as several other bovid genera in order to elucidate the position of caprines within the family Bovidae, and to determine the exact composition of the group. Phylogenetic analyses confirm firstly that Pseudoryx and Saiga do not belong to caprines, and secondly, that all tribes classically defined in the literature are not monophyletic, supporting the inclusion of all caprine species into a unique enlarged tribe Caprini sensu lato . Our results are in contradiction with previous investigations suggesting a sister-group relationship between Ovis (sheep and mouflons) and Budorcas (takins). By using a molecular calibration point at 18.5 Mya for the first appearance of bovids, we estimated divergence times with our molecular data. We also performed biogeographic inferences to better understand the origin and diversification of caprines during the Neogene. Our analyses suggest that caprines shared a common ancestor with Alcelaphini and Hippotragini in the middle-late Miocene (13.37 ± 0.70 Mya). Our results also indicate that the extant generic diversity of caprines resulted from a rapid adaptive radiation during the late Miocene, at 10.96 ± 0.73 Mya. We propose that this adaptive radiation resulted from the acquisition of reduced metacarpals, a key innovation which occurred during the late Miocene as a consequence of insularity isolation in the mountainous mega-archipelago between Mediterranean and Paratethys Seas.     

New skull of Schizodelphis sulcatus Gervais, 1861 (Mammalia, Odontoceti, Eurhinodelphinidae) from the Lower Miocene of Pignan (Hérault, France) and its implications for systematics of Eurhinodelphinidae

Schizodelphis sulcatus is the type species of the Eurhinodelphinid genus Schizodelphis and was until now only known from a rather fragmentary specimen, lacking some important parts of the skull (vertex, ear bones and apex of rostrum). The study of a well-preserved skull of S. sulcatus Gervais, discovered in the lower Miocene of Hérault (France), helps to improve the diagnosis of this enigmatic species. Diagnostic features include the low height at the base of the rostrum, the lower position of the paroccipital process compared to the height of the postglenoid process in lateral view, the ventrally deflected postglenoid process and the ovoid-shaped nasals. It also confirms that the type species of Schizodelphis is indeed distinct. For the first time in Schizodelphis, the vertex is preserved and the ear bones are found in anatomic connection with the basicranium. A cladistic analysis of cranial characters of Eurhinodelphinidae, including ear bones, is performed and reveals a close phylogenetic relationship between S. sulcatus and the North American species Schizodelphis barnesi. This analysis also highlights some apomorphies of the genus as the smoothly laterally deflected anterior spine of the bulla in ventral view and the reestablishment of the contact between the falciform process of the squamosal and the lateral lamina of the pterygoid. Conversely, the attribution of Schizodelphis morckhoviensis to the genus Schizodelphis is questioned by this analysis.     

Molecular phylogeny and biogeography of Oriental voles: genus Eothenomys (Muridae, Mammalia)

Oriental voles of the genus Eothenomys are predominantly distributed along the Southeastern shoulder of the Qinghai-Tibetan Plateau. Based on phylogenetic analyses of the mitochondrial cytochrome b gene (1143 bp) obtained from 23 specimens (eight species) of Oriental voles collected from this area, together with nucleotide sequences from six specimens (two species) of Japanese red-backed voles (Eothenomys andersoni and Eothenomys smithii) and five species of the closely related genus Clethrionomys, we revised the systematic status of Eothenomys. We also tested if vicariance could explain the observed high species diversity in this area by correlating estimated divergence times to species distribution patterns and corresponding paleo-geographic events. Our results suggest that: (1) the eight species of Oriental voles form a monophyletic group with two distinct clades, and that these two clades should be considered as valid subgenera--Eothenomys and Anteliomys; (2) Eothenomys eleusis and Eothenomys miletus are not independent species; (3) Japanese red-backed voles are more closely related to the genus Clethrionomys than to continental Asian Eothenomys taxa; and (4) the genus Clethrionomys, as presently defined, is paraphyletic. In addition, the process of speciation of Oriental voles appears to be related to the Trans-Himalayan formation via three recent uplift events of the Qinghai-Tibetan Plateau within the last 3.6 million years, as well as to the effects of the mid-Quaternary ice age.     

The phylogeny and taxonomy of Hippopotamidae (Mammalia: Artiodactyla): a review based on morphology and cladistic analysis

The phylogeny and taxonomy of the whole family Hippopotamidae is in need of reconsideration, the present confusion obstructing palaeoecology and palaeobiogeography studies of these Neogene mammals. The revision of the Hippopotamidae initiated here deals with the last 8 Myr of African and Asian species. The first thorough cladistic analysis of the family is presented here. The outcome of this analysis, including 37 morphological characters coded for 15 extant and fossil taxa, as well as non-coded features of mandibular morphology, was used to reconstruct broad outlines of hippo phylogeny. Distinct lineages within the paraphyletic genus Hexaprotodon are recognized and characterized. In order to harmonize taxonomy and phylogeny, two new genera are created. The genus name Choeropsis is re-validated for the extant Liberian hippo. The nomen Hexaprotodon is restricted to the fossil lineage mostly known in Asia, but also including at least one African species. The genus Hippopotamus is confirmed. These changes represent substantial advances for understanding the evolutionary history of the Hippopotamidae, and provide a new framework for future studies.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 1–26.     

Molecular genetic diversity of musk deer Moschus moschiferus L., 1758 (Ruminantia, Artiodactyla) from the northern subspecies group

Genetic diversity within the northern subspecies group of musk deer Moschus moschiferus L., 1758 was examined based on the mtDNA control region hypervariable fragment (300-bp) sequence polymorphism. Nucleotide diversity, constituting 2.6% for the whole sample (n = 34), varied in the range from 0.6 to 1.9% for individual subspecies. Maximum values of this index were observed for Siberian subspecies (M. m. moschiferus), which had the widest range. Genetic similarity between the haplotypes of the musk deer from the Far East (Russia) and Sakhalin Island, which grouped in one cluster in a phylogenetic tree, was demonstrated. The data obtained indicate that the distribution of musk deer along the territory of Russia occurred from Eastern Siberia to the Far East, and from there to the Sakhalin Island. A currently observed decrease of the musk deer population number along with the increased habitat fragmentation can result in a decrease of the total genetic diversity and in inbreeding depression in the local isolated groups.     

The phylogeny of the Rhinocerotoidea (Mammalia, Perissodactyla)

A new phylogeny of the Superfamily Rhinocerotoidea is proposed, based upon an analysis of shared derived characters of the skull, teeth and skeleton. Hyrachyus is considered the primitive sister-taxon of the three rhinocerotoid families (Amynodontidae, Hyracodontidae, Rhinocerotidae), and the amynodonts appear to be the sister-group of hyracodonts and rhinocerotids. The relationships of primitive hyracodonts and rhinocerotids are clarified. Menoceras and Pleuroceras (new Subfamily Menoceratinae) are removed from the Diceratheriinae, since they appear to be more closely related to higher rhinoceroses than they are to Diceratherium. Of the three major monophyletic groups of higher rhinocerotids (aceratherines, teleoceratines and rhinocerotines), the last two groups are more closely related to each other than either is to aceratherines. Toxotherium and Schizotheroides are tentatively removed from the Rhinocerotoidea and placed in the Lophiodontidae.     

The beaver (Mammalia,Castoridae) from the Miocene of Sandelzhausen (southern Germany)

The Early/Middle Miocene boundary locality Sandelzhausen (MN5, southern Germany) yielded three beaver teeth: one lower incisor fragment and two strongly worn upper premolars, a right and a left one. The latter are so similar in wear, height, size, and enamel pattern that they probably represent one individual. Although the teeth have been referred to Chalicomys jaegeri Kaup, 1832 (e.g., Hugueney 1999, Fahlbusch 2003), it seems more likely that they belong to Steneofiber depereti Mayet, 1908. However, the assignment of the present material to either one of these species is difficult, because in the advanced wear stage some diagnostic characteristics cannot be assessed. As to ecological implications for the locality Sandelzhausen the taxonomic identity of these two beaver species may not be important. Both are known from riverine, lacustrine or swamp sediments, and it can be assumed that they usually lived in close vicinity to water. Thus, the presence of beaver remains supports the presence of water at Sandelzhausen. However, the teeth might have an allochthonous origin from creeks of the rising Alps.