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.     

Hippotraginae (Bovidae,Artiodactyla, Mammalia) from the Late Miocene of Tuva

Fossil remains of Late Miocene (Baode, NMU10-NMU11) horse antelopes from Tuva (Russia) are described, including Tragoreas sp., Protoryx tuvaensis sp. nov. from the Taralyk Cher locality and Quirliqnoria sp. from the Kholu locality. A new species, Protoryx tuvaensis Dmitrieva et Serdyuk, is described. These taxa compose a Late Miocene antelope assemblage of a new eastern geographical point (Russia, Tuva, Baode, NMU10-MN11).     

Geometric morphometrics of mandibles for dietary differentiation of Bovidae (Mammalia: Artiodactyla)

The mammalian family Bovidae has been widely studied in ecomorphological research, with important applications to paleoecological and paleohabitat reconstructions. Most studies of bovid craniomandibular features in relation to diet have used linear measurements. In this study, we conduct landmark-based geometric-morphometric analyses to evaluate whether different dietary groups can be distinguished by mandibular morphology. Our analysis includes data for 100 species of extant bovids, covering all bovid tribes and 2 dietary classifications. For the first classification with 3 feeding categories, we found that browsers (including frugivores), mixed feeders, and grazers are moderately well separated using mandibular shape. A finer dietary classification (frugivore, browser, browser–grazer intermediate, generalist, variable grazer, and obligate grazer) proved to be more useful for differentiating dietary extremes (frugivores and obligate grazers) but performed equally or less well for other groups. Notably, frugivorous bovids, which belong in tribe Cephalophini, have a distinct mandibular shape that is readily distinguished from all other dietary groups, yielding a 100% correct classification rate from jackknife cross-validation. The main differences in mandibular shape found among dietary groups are related to the functional needs of species during forage prehension and mastication. Compared with browsers, both frugivores and grazers have mandibles that are adapted for higher biomechanical demand of chewing. Additionally, frugivore mandibles are adapted for selective cropping. Our results call for more work on the feeding ecology and functional morphology of frugivores and offer an approach for reconstructing the diet of extinct bovids.     

牛科(哺乳纲:偶蹄目)动物与食物有关的适应形态模式

利用逐步分辨分析方法（Stepwise discriminant analysis,SCDA）检测了广义牛科动物的颅齿部结构,这些结构特征可以作为采食行为生态适应特征。在本研究中,测量了72种广义牛科动物的28个颅齿部结构。逐步分辨分析方法得出了6种采食方式适应类型：一般粗食者、新鲜禾草粗食者、开阔生境混合型采食者、精食者、郁闭生境混合型采食者、食果者。用103个标本检测了分辨指标的预测能力,所用标本为缺损标本,大多数缺少一项或多项结构。从这些标本获得的分辨函数的平均预测能力为94％,比用72种广义牛科动物标本建立的分辨函数的平均预测能力（98％）低一些。从一个颅齿部结构小样本建立的分辨函数可以用于考古发掘物中不完整标本的研究。这些指标与用颅下结构测量建立的运动能力和生境选择的指标相结合,可以推断古牛科动物的个体生态学以及古环境重建。     

Competitive resource sharing by seaweed-eating sheep Ovis aries (Mammalia: Artiodactyla)

The seaweed-eating sheep of North Ronaldsay, Orkney, were used to examine the distribution of a herbivore in relation to patchily distributed food. Seaweed occurred in discrete patches on the beach and sheep distributed themselves amongst the patches to ked. Sheep selected Rhodymenia palmata in autumn and Rhodymenia palmala and Alara esculenta in spring. There was a statistically significant agreement between the distribution of food and the distribution of sheep between patches. There was a significant correlation between the distribution of the two sexes between patches. The sex ratio in patches differed from the beach as a whole, probably due to social groupings. Social organization will constrain, and may account for, the unexplained variation in the sheep distribution.     

Molecular evidence for the polyphyly of the genus Hemitragus (Mammalia, Bovidae)

The genus Hemitragus includes three species of tahr distributed in distant geographical areas: the Himalayan tahr, H. jemlahicus, occupies the southern flanks of the Himalaya Mountains; the Nilgiri tahr, H. hylocrius, is endemic to southern India; and the Arabian tahr, H. jayakari, is unique to the mountains of south-east Arabia. All previous investigations were based on morphology, and these three species together have never been included in a molecular phylogenetic study. In this study, we constructed a molecular phylogeny of the tribe Caprini sensu lato to determine the taxonomic status of the three species of tahr. Phylogenetic analyses were carried out on a matrix including most extant species currently described in the tribe Caprini sensu lato, and 3165 nucleotide characters, coming from four different markers, i.e., an intron of the nuclear gene coding for the protein kinase C iota, and three mitochondrial genes (subunit II of the cytochrome c oxidase, cytochrome b, and 12S rRNA). The results show that the genus Hemitragus is polyphyletic, as H. jemlahicus is associated with Capra (goats), H. hylocrius is the sister-group of Ovis (sheep), and H. jayakari is allied with Ammotragus lervia (aoudad). In the light of these unexpected results, we revaluate the validity of the morphological characters originally used for defining the genus Hemitragus. At least, we propose a new taxonomy, where the three species of tahr are ranged into three monospecific genera: the genus Hemitragus is restricted to the Himalayan tahr, and two new genera are created: Arabitragus for the Arabian tahr and Nilgiritragus for the Nilgiri tahr.     

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.     

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.     

Morphology, constraints, and scaling of frontal sinuses in the hartebeest, Alcelaphus buselaphus (Mammalia: Artiodactyla, Bovidae)

The frontal sinuses of bovid mammals display a great deal of diversity, which has been attributed to both phylogenetic and functional influences. In-depth study of the hartebeest (Alcelaphus buselaphus), a large African antelope, reveals a number of previously undescribed details of frontal sinus morphology. In A. buselaphus, the frontal sinuses conform closely to the shape of the frontal bone, filling nearly the entire element. However, the horncores are never extensively pneumatized, contrasting with the condition seen in many other bovids. This evidence is inconsistent with the hypothesis that sinuses are opportunistic pneumatizing agents, suggesting that phylogenetic factors also play a role in determining sinus size. Both cranial sutures and neurovasculature appear to constrain the growth of sinuses in part. In turn, the sinus also affects the growth of the parietal; apparently this element is not truly pneumatized by the sinus in most cases, but the bone's shape changes under the influence of the sinus. Furthermore, the sinuses present relatively few struts when compared with the sinuses of some other bovids, such as Ovis. By adapting methods previously developed for measuring structural parameters of trabecular bone, it is possible to quantify certain aspects of sinus morphology. These include the number of bony struts within the sinus, the spacing of these struts, and the size of individual cavities within the sinus. Some differences in the number of struts are evident between subspecies. Similarly, significant differences occur in the relative number of struts between male and female A. buselaphus, which may be related to behavior. The volume of the sinus is strongly correlated with the size of the frontal, but less so with overall cranial size. This finding illustrates the importance of choosing variables carefully when comparing sinus sizes and growth between species.     

Phylogenetic reconstructions in the genus Capra (Bovidae, Artiodactyla) based on the mitochondrial DNA analysis

Mitochondrial genome fragments were examined in all species of the genus Capra (Bovidae, Artiodactyla). Phylogenetic analysis was carried out using 59 cytochrome b gene sequences (392 bp), and 22 sequences of the mtDNA variable fragment (402 bp). In the control region, two unique deletions were revealed. One of the deletions was found only in Capra cilindricornis (17 bp), while another one grouped C. caucasica with C. aegagrus (1 bp). The group of Caucasian wild goats splits into two clades, and furthermore, the sequences of C. caucasica demonstrate remarkable similarity to the sequences of C. aegagrus, while C. cylindricus seems to have evolved independently for a long period of time. It was demonstrated that C. pyrenaica and C. ibex were extremely close to one another. Capra sibirica formed an outer group relative to the other species, and according to our data, was the most ancient species of the genus. On the contrary, genetic distance separating C. falconeri (the most independent species of the genus related to its morphology) from the other species is small.     

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.     

Chromosomes of the antelope genus Kobus (Artiodactyla, Bovidae): karyotypic divergence by centric fusion rearrangements

G- and C-banded karyotypes of four species of the genus Kobus were compared using the standard karyotype of Bos taurus. Chromosomal complements were 2n = 50-54 in K. ellipsiprymnus, 2n = 50 in K. kob, 2n = 48 in K. leche, and 2n = 52 in K. megaceros. The number of autosomal arms in all karyotypes was 58. Fifteen autosomal pairs were conserved among these four species, including the 1;19 and 2;25 centric fusions, and autosomal differences involved eight centric fusion rearrangements. Five centric fusions were each unique to a particular taxon: 3;10 (K. leche), 3;11 and 6;29 (K. kob), and 5;17 and 7;11 (K. ellipsiprymnus). The 4;7 fusion occurred in K. leche and K. megaceros, whereas the 5;13 fusion occurred in K. kob and K. leche; the 6;18 fusion was found in three species but was absent in K. kob. Differences between the X chromosomes of the four Kobus species were attributed to heterochromatic additions or deletions, and Y-chromosome differences may have been the result of pericentric inversion. G-banded karyotypes of putative K. l. leche and K. l. kafuensis appeared identical, as did C-banded karyotypes of the two subspecies. Karyotypes of K. e. ellipsiprymnus and K. e. defassa differed as a result of the 6;18 centric fusion, which was polymorphic in K. e. defassa, and the 7;11 centric fusion, which was polymorphic in K. e. ellipsiprymnus but absent in K. e. defassa. Several centric fusions were related by monobrachial chain-IV complexes; however, records of hybridization indicate that reproductive isolation between at least certain species of Kobus is incomplete. Karyotypic differences between K. ellipsiprymnus (including K. e. ellipsiprymnus and K. e. defassa), K. kob, K. leche, and K. megaceros support the validity of these taxa, as well as the need to manage them as separate populations.     

陕西蓝田灞河组晚中新世Gazella羚羊的形态特征与分类（英文）

Gazella羚羊是"三趾马动物群"中常见成员,在晚中新世至更新世地层中广泛分布,演化速率相对较快,具有重要的生物地层学及生态指示意义,但在我国尚未有保德期之前的化石报道。本文研究的化石发现于陕西蓝田灞河组中部,磁性地层学资料显示其年代为晚中新世灞河期。化石标本包括了5个近乎完整的头骨、下颌以及颅后骨骼。根据发现的标本装架起第一个完整的Gazella羚羊骨架。形态对比与测量数据表明,蓝田标本与巴基斯坦西瓦里克发现的Gazella lydekkeri非常相近,而不同于欧洲晚中新世常见的Gazella各种以及我国发现的种类。从肢骨的形态分析与测量比例数据来看,Gazella cf.G.lydekkeri适合快速奔跑运动,可能生活在相对开阔的环境中,与灞河期动物群的生态类型以及与灞河组沉积学、同位素地球化学等研究所指示的相对干旱、半干旱的开阔草原环境相一致。     

牛科(哺乳纲:偶蹄目)动物与生境利用有关的适应形态模式

对广义牛科动物颅后骨骼的多元变量分析揭示了牛科生境利用和体型之间的骨学特征。利用逐步分辨分析方法和一个基于机器学习的决策树方法鉴别了每种生境中牛科动物颅后解剖结构的形态特征。从110个广义牛科动物测量了43个指标进行了这项分析。利用所有主要肢骨测量值和以单根肢骨测量为主的测量值获得的分辨函数和决策树可以完美地区分适应开阔生境、森林和山地的牛科动物(在所有分析中得到了100 %正确的再分类)。由于调整的函数仅涉及到很小的颅后骨骼测量集,这些函数可以应用于研究考古学和古生物学发掘物中保存的不完整标本。这些表征生境利用的生态适应函数与那些用颅齿部性状建立、用于推测牛科动物食物选择的函数结合,具有刻画已灭绝的分类类群的古个体生态学和重建古环境的潜力。我们还分析了多元回归是否较单一因子回归表现出较高的预测能力,并提出了从每一种单根主要肢骨测量的颅后形态变量得到的最好代数函数     

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.     

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.