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.     

Caprinae (Bovidae,Artiodactyla, Mammalia) from the Neogene of Mongolia

Gobiocerus mongolicus Sokolov, 1952, Turcocerus kekemaidengensis Ye Jie et al., 1999; and Hypsodontus sp. were recorded in Miocene deposits of the Valley of Lakes (MN4-MN8) and Oioceros atropatenes (Rodler et Weithofer, 1890) occurred in the Mio-Pliocene (MN11-MN20) of northwestern Mongolia. Gobiocerus, Turcocerus, and Hypsodontus represent the same evolutionary lineage. Thus, Caprinae existed in Central Asia in the Early Miocene. They diverged from the Bovidae at the Oligocene-Miocene boundary. The presence of early bovids in this region suggests that this group could have appeared in Central Asia.     

On some spiral-horned antelopes (Mammalia: Artiodactyla: Bovidae) from the Late Miocene of Turkey, with remarks on their distribution

Examination of some collections from several late Miocene Turkish localities allows us to describe and definitely identify for the first time several species of spiral-horned antelopes in this country:Palaeoreas lindermayeri, Protragelaphus skouzesi, Prostrepsiceros zitteli andNisidorcas sp. During the first part of the late Miocene, several endemic species are restricted to central Anatolia, while more widespread taxa appear in the later part of this period, suggesting a decrease in provinciality of this area.     

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.     

New Late Miocene murids from China (Mammalia, Rodentia)

New murids of Late Miocene (medial Baodean Chinese Mammal Unit) age from Inner Mongolia, northern China, and from Yunnan Province, southern China, are described. Hansdebruijnia perpusilla nov. sp. represents the earliest known and morphologically most primitive species of the genus, which is known from the latest Miocene of Europe and western Asia. The new species suggests an eastern Palaearctic origin of Hansdebruijnia. “Progonomys” yunnanensisQIU and STORCH, 1990 from Lufeng, Yunnan Province, is referred to Linomys nov. gen. New samples from Leilao, Yuanmou County, Yunnan Province, are included in this species, although this population is somewhat more primitive than that from Lufeng. Leilaomys zhudingi nov. gen. and sp. from Leilao shows a unique combination of apomorphic and plesiomorphic characters, which suggest an early divergence from the murid stem. The new findings indicate that our knowledge of the early radiation of murids in southeast Asia is still in its infancy.     

Evolution and taxonomy of the wild species of the genus Ovis (Mammalia,Artiodactyla, Bovidae)

New insights for the systematic and evolution of the wild sheep are provided by molecular phylogenies inferred from Maximum parsimony, Bayesian, Maximum likelihood, and Neighbor-Joining methods. The phylogeny of the wild sheep was based on cytochrome b sequences of 290 samples representative of most of the sub-species described in the genus Ovis. The result was confirmed by a combined tree based on cytochrome b and nuclear sequences for 79 Ovis samples representative of the robust clades established with mitochondrial data. Urial and mouflon, which are either considered as a single or two separate species, form two monophyletic groups (O. orientalis and O. vignei). Their hybrids appear in one or the other group, independently from their geographic origin. The European mouflon O. musimon is clearly in the O. orientalis clade. The others species, O. dalli, O. canadensis, O. nivicola, and O. ammon are monophyletic. The results support an Asiatic origin of the genus Ovis, followed by a migration to North America through North-Eastern Asia and the Bering Strait and a diversification of the genus in Eurasia less than 3 million years ago. Our results show that the evolution of the genus Ovis is a striking example of successive speciation events occurring along the migration routes propagating from the ancestral area.     

A new species of Megacerini (Cervidae, Artiodactyla) from the Late Miocene of Taralyk-Cher, Tuva (Russia), and remarks on the relationships of the group

A new species of a megacerine deer, Praesinomegaceros venustus, from the Taralyk-Cher locality in the Tuva region on the south of Eastern Siberia, Russia, is described. P. venustus nov. sp., the oldest known megacerine, occurred in Asia in the Late Turolian, about 7 Ma. The early history of megacerines is discussed. The existence of the phyletic lineage Cervavitus-Praesinomegaceros-Sinomegaceros in Asia is confirmed.     

Les Lagomerycidae (Artiodactyla, Mammalia) de FranceThe French Lagomerycidae (Cervoidea, Artiodactyla, Mammalia)

The Lagomerycidae, a small family of the Cervoidea, is known in France by the genera Lagomeryx, Ligeromeryx and Stephanocemas. Lagomeryx contains three species, the other two genera one species only. This family is known from MN3 to MN5.     

A new Late Miocene ovibovine-like bovid (Bovidae,Mammalia) from the Kassandra Peninsula (Chalkidiki,Northern Greece) and implications to the phylogeography of the group

A new Late Miocene bovid, Urmiatherium kassandriensis sp. nov., from Northern Greece is described. The material comes from the Fourka locality in the Kassandra Peninsula (Chalkidiki), and the included fauna is estimated to be of Vallesian age. The two preserved crania represent a medium-sized taxon with short, conical horn cores, a flat cranial roof (consisting of the posterior part of the frontals, parietal and occipital), thick and porous frontals and pneumatized short parietals, an extremely thick basioccipital with voluminous posterior tuberosities and accessory articular facets for the atlas. The specialized atlanto-occipital joint recalls Pleistocene and extant ovibovines, but the braincase structure as a whole and the horn core features closely match Late Miocene ovibovine-like taxa, especially Plesiaddax and even more Urmiatherium. Nevertheless, the Kassandra bovid differs from representatives of both genera in the simpler horn core morphology and external brain anatomy. Urmiatherium is known to appear first in China and Iran at about 7.8 Ma, whereas its westernmost appearance on Samos Island (Greece) is dated much later. The presence of Urmiatherium kassandriensis sp. nov. in N. Greece suggests a farther west and earlier (Vallesian at least) first appearance of the genus. This would justify a basic geographic and phylogenetic split of Urmiatherium into two main Turolian lineages: a central-eastern Asian one leading to the sister species U. polaki and U. intermedium and a western one leading to U. rugosifrons.     

A new hipparion (Mammalia,Perissodactyla) from the Vallesian (Late Miocene) of Greece

A small hipparion from the Vallesian (early Late Miocene) of Northern Greece is studied. It is coming from the locality “Ravin de la Pluie” of the lower Axios valley, near Thessaloniki. Its characteristics as the small size, the short symphysis, the long snout, the high enamel plication, the well developed protostylid, the large m₃ and the elongated m₁-m₃ series distinguish it from the other known eurasiatic small hipparions, allowed us to derive a new species,Hipparion macedonicum. It lived during Vallesian and can be used as a stratigraphic species for this period.     

Fossil Tragelaphini (Artiodactyla: Bovidae) from the Late Pliocene Hadar Formation,Afar Regional State,Ethiopia

The fossil tragelaphins from the late Pliocene of Hadar are described. These are Tragelaphus lockwoodi, new species, and Tragelaphus aff. T. nakuae. Tragelaphus lockwoodi bears long horns that define one complete spiral and that are mediolaterally compressed at the base. In these and other morphological characteristics it approaches the greater kudu, T. strepsiceros, and makes a good ancestral candidate for this living species. The Hadar T. aff. T. nakuae is similar to other specimens of this species from sites >2.8 Ma in East Africa and demonstrates well the major differences between the earlier and later representatives of this taxon. The sizes and morphological variation in the large Hadar T. aff. T. nakuae sample supports the idea that female individuals of this species were horned as is the case today in the elands and the bongo. Tragelaphus lockwoodi is present only in the lower beds of the Hadar Formation, and in small numbers, while T. aff. T. nakuae is recovered in relative abundance from throughout the ca. 3.4-ca. 2.9 Ma sequence.     

Occurrence of the anthracotheriid Brachyodus (Artiodactyla,Mammalia) in the early Middle Miocene of Thailand

Remains of Brachyodus cf. onoideus from the Ban Na Sai Basin (Thailand) are described. They correspond to the first occurrence of that genus in South Asia, and might document the latest record of the genus in the Old World. Sihongotherium from the Middle Miocene of eastern China is likely a representative of Brachyodus that was misinterpreted. To cite this article: S. Ducrocq et al., C. R. Palevol 2 (2003).     

Late Miocene Microstonyx remains (Suidae, Mammalia) from Northern China

Several large suid cranial remains attributed to Microstonyx major are part of a new Hipparion Fauna collection from the Hezheng area, Northern China. The new material confirms the presence of Microstonyx in the late Miocene of the area. The Chinese form belongs to a small-sized eastern population with reduced premolar row and clear sexual bimodality. Statistical comparison shows that Microstonyx major was a polymorphic species and reinforces recognition of Hippopotamodon as a separate genus, defined by relatively stout premolars resulting from a different underlying pattern of allometric growth. The presence of Microstonyx in North China and the distinct suid assemblage that lived there suggest biogeographic connections between Northern China and Western Eurasia in contrast to isolation from Southern China and the Indian subcontinent. The suid fauna of the late Miocene of Northern China seems to have been restricted to the later, more humid phase represented by the Red Clay faunas.     

Phylogeny of the Bovidae (Artiodactyla, Mammalia), based on mitochondrial ribosomal DNA sequences.

Portions of the 12S and 16S mitochondrial ribosomal genes for 16 species representing nine tribes in the mammal family Bovidae were compared with six previously published orthologous sequences. Phylogenetic analysis of variable nucleotide positions under different constraints and weighting schemes revealed no robust groupings among tribes. Consensus trees support previous hypotheses of monophyly for four clades, including the traditional subfamily Bovinae. However, the basal diversification of bovid tribes, which was largely unresolved by morphological, immunodiffusion, allozyme, and protein sequence data, remains unresolved with the addition of DNA sequence data. The intractability of this systematic problem is consistent with a rapid radiation of the major bovid groups. Several analyses of our data show that monophyly of the Bovidae, which was weakly supported by previous morphological and molecular work, is questionable.     

The fossil Bovidae (Artiodactyla, Mammalia) from Gesher Benot Ya'aqov, Israel: Out of Africa during the Early-Middle Pleistocene transition

We report the study of the collection of fossil bovid specimens from the Early-Middle Pleistocene Acheulian site of Gesher Benot Ya‘aqov. This locality, situated in the Levantine Corridor (the bottleneck that connects Africa and Eurasia) is a key site to explain the faunal and human dispersals out of Africa during the Matuyama/Brunhes boundary around 0.8 Ma. Two species of bovine (Bos sp., and Bovini gen. et sp. indet. cf. Bison sp.), one antelope (Gazella sp. cf. G. Gazella), and another indeterminate Bovidae gen. et sp. indet., have been recorded. The largest species, Bos sp., is an African immigrant related to the species from the Eritrean site of Buia, Bos buiaensis, which evolved from the buffalo of Olduvai Pelorovis oldowayensis, and colonized the Eurasian continent in parallel with the dispersal of the Acheulian culture into the northern continent. Numerous important species first recorded in several localities of Early-Middle Pleistocene transition from Eurasia are included in this dispersal out of Africa, including the megaherbivore, Palaeoloxodon antiquus, and the carnivores Crocuta crocuta, and later, Panthera leo and Panthera pardus. This faunal turnover is coincident with the change to colder climates that dominated the Middle Pleistocene.     

Sexual Dimorphism of the Larynx of the Mongolian Gazelle (Procapra gutturosa Pallas, 1777) (Mammalia, Artiodactyla, Bovidae)

The larynges (except for the epiglottis) of two adult Mongolian gazelles, one male and one female, were dissected. This species is characterized by a pronounced sexual dimorphism of the larynx. Dimorphism with regard to the size of the entire larynx and of the thyroid cartilage is about 2:1 whereas the difference of mean body mass is about 1.3:1 between males and females. Unexpectedly, and in contrast to other bovids, the larynx of the male Mongolian gazelle has a paired lateral laryngeal ventricle. However, in contrast to horse, dog, pig and many primate species also possessing such a paired ventricle, its rostral opening in the Mongolian gazelle is situated lateral to the corniculate process of the arytenoid cartilage. The neck of the laryngeal ventricle is embraced by the bifurcated cuneiform process of the epiglottis. Despite the enlarged laryngeal cartilages, the vocal process of the male arytenoid cartilage is relatively shorter than that of the female. The male thyroarytenoid muscle is clearly separated into a rostral ventricular muscle and a caudal vocal muscle whereas the female's, as in other bovids, is almost uniform. The lateral sac of the two-chambered laryngeal ventricle in the male projects laterally between the ventricular and the vocal muscle. As in the domestic bovids and in many other artiodactyls the larynx of the male Mongolian gazelle is lacking any rostrally directed membraneous portion of the vocal fold. Instead, the thick and tough bow-like vocal fold projects caudally into the infraglottic cavity and is supported by a peculiar pan-like fibroelastic pad. This resilient element, situated medial to the bipartite thyroarytenoid muscle, might be a homologue of the vocal ligament, eventually including lateral portions of the elastic cone. A fibroelastic pad is absent in the female. The resilient floor of the laryngeal vestibulum, ventral to the fibroelastic pad, is rostrally and caudally subducted by tube-like spaces. Evolutionary enlargement of the male larynx, including the vocal folds, and of the caudal portions of the vocal tract may have shifted the fundamental and formant frequencies to a lower register. The paired lateral laryngeal ventricle might produce an amplitude increase of the vocalizations assisted by differential action of the bipartite thyroarytenoid muscle. In addition, the peculiar shape, size and tough consistency of the male vocal folds may, as in roaring felids, assist in producing high amplitude and low frequency vocalizations. Perhaps the biological role of the enlarged male larynx of Procapra gutturosa has evolved in relation to its mating system. In the rutting season, dominant males establish individual territories and maintain harems. During prolonged courtship prior to mating, these males perform an acoustic display uttering loud and guttural bellows. In addition, the bulging ventral neck region of males may serve as an optical attractant for the females. Thus, the evolution of the enlarged larynx of the male Mongolian gazelle may have been favoured by sexual selection.     

Palaeogeographic significance of the giraffid remains (Mammalia,Arctiodactyla) from Cessaniti (Late Miocene,Southern Italy)

The late Tortonian – early Messinian shallow marine sands of Cessaniti area (Monte Poro, Vibo Valentia, Southern Italy) yield marine and continental vertebrates. The best represented taxon is the Sirenian Metaxytherium serresii, while the terrestrial mammal assemblage includes a boselafine bovid, an hexaprotodontid hippopotamus, the giraffids Samotherium cf. boissieri and Bohlinia cf. attica, a rhino and the elephantid Stegotetrabelodon syrticus. Until now, the latter was a species with an exclusive Afro-Arabian distribution and the record of Cessaniti is the first outside Afro-Arabia. Our attention is here focused on the occurrence of Samotherium cf. boissieri and Bohlinia cf. attica, both being species well represented in the Pikermian Biome. Although evidences of the distribution of the genus Samotherium in Late Miocene African assemblages are weak, it is reported at several sites, while a new species of Bohlinia reported in Chad is still debated. At Cessaniti, the co-occurrence of two giraffid taxa typical for the Pikermian biome together with a frankly Afro-Arabic species (S. syrticus), further marks the existence of a land connection between the Cessaniti area and North Africa as well as the evidence of a phase of expansion of the Pikermian Biome into the African continent.     

New material of Carnivora (Mammalia) from the Late Miocene of Axios Valley,Macedonia, Greece

A new collection of carnivores from the Late Miocene deposits of Axios Valley, Macedonia, Greece was collected in 2004–2009 at the localities named Xirochori 1 (XIR) and Ravin de la Pluie (RPl), dated to the Late Vallesian (MN 10), as well as at Ravin des Zouaves 5 (RZO), dated to the Early Turolian (MN 11). The studied material is described and compared morphologically and biometrically with other materials from Greece and its neighboring area. The following species are determined: XIR: Dinocrocuta sp.; RPl: Eomellivora wimani, Adcrocuta eximia leptoryncha, ?Hyaenictis sp., Metailurus parvulus; RZO: Plioviverrops orbignyi, Machairodus giganteus. This collection is interesting because it includes the upper deciduous dentition of A. eximia and Dinocrocuta, which are only known from limited material until now, especially the latter taxon. E. wimani is recognized for the first time in Greece and the Balkans, while M. parvulus is identified for the first time in the Vallesian of Greece. The possible presence of Hyaenictis in RPl is the first indication of its presence in the Vallesian. The guild structure diagrams of the carnivore fauna from the three studied localities provide some indications of their palaeoenvironment, which fits quite well with the results of previous studies on the palaeoenvironments of these localities and of the wider Eastern Mediterranean region.