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.     

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.     

On habitat selection by the goral Nemorhaedus goral bedfordi (Bovidae, Artiodactyla)

Habitat selection, diet, and escape behaviour of grey goral ( Nemorhaedus goral bedfordi ) were studied in Majhatal Harsang Wildlife Sanctuary (MHWS) in the Indian Himalaya. Five trails covering a total length of 8.5 km were monitored in the morning hours for availability of habitat components and their use by goral. Habitat components quantified were: vegetation type, tree density, shrub understorey, shrub height, and slope angle. Diet composition of goral was determined through faecal analysis. Information on escape strategy was gathered by recording the escape areas used by disturbed animals.
Goral fed almost entirely on grass (92.2% in the cold season and 98.3% in the warm season). They preferred open vegetation communities with good grass cover and avoided shrub-rich patches, especially areas where the shrub height exceeded their shoulder height. Goral selected patches with fresh grass sprout and were partial to steep (>30°) slopes.
In the morning, goral avoided most forested vegetation types in the area owing to extensive understorey. However, there was high use of forest cover with little understorey 'for escape' (32%, n = 200). Other escape terrain used were cliffs (31%), open gullies (16%), and rocky areas (13%).     

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).     

Centric fusion differences among Oryx dammah, O. gazella, and O. leucoryx (Artiodactyla, Bovidae)

G- and C-banded karyotypes of the genus Oryx were compared using the standard karyotype of Bos taurus. Chromosomal complements were 2n = 56 in O. gazella gazella, 2n = 58 in O. g. beisa and O. g. callotis, 2n = 56-58 in O. dammah, and 2n = 57-58 in O. leucoryx. The number of autosomal arms in all karyotypes was 58. Nearly all variation in diploid number was the result of three independent centric fusions, but one 2n = 57 specimen of O. g. gazella deviated from the normal complement of 2n = 56 due to XXY aneuploidy. A 2;17 centric fusion was fixed in O. g. gazella, whereas O. g. beisa and O. g. callotis lacked this fusion and had indistinguishable karyotypes. Oryx dammah was polymorphic for a 2;15 centric fusion, and O. leucoryx was polymorphic for an 18;19 centric fusion. The five Oryx taxa shared a fixed 1;25 centric fusion; the small acrocentric element involved in the 1;25 fusion was identified by fluorescence in situ hybridization using a cosmid specific to Bos chromosome 25. The X and Y chromosomes were also conserved among the five taxa. Oryx g. gazella differed from the other Oryx species because of the fixed 2;17 centric fusion. This difference reflects an apparently longer period of geographic isolation between O. g. gazella and other populations of Oryx, and it is consistent with the classification of O. gazella and O. beisa as distinct species (see Kingdon, 1997). The lack of monobrachial relationships among the Oryx taxa indicates that sterility barriers between species have not developed. Viability of hybrid offspring constitutes a threat to captive breeding programs designed for endangered species conservation; in the case of Oryx, the 2;15, 2;17, and 18;19 metacentrics could serve as marker chromosomes for assessing hybridization between certain Oryx taxa.     

Large sequence divergence of mitochondrial DNA genotypes of the control region within populations of the african antelope, kob (Kobus kob)

The genetic differentiation among kob populations (Kobus kob) representing two recognized subspecies was examined using mitochondrial control region sequences. Two distinct lineages (estimated sequence divergence of 9.8%) exhibited different geographical distributions and do not coincide with previously recognized ranges of subspecies. The presence of the two lineages was further supported with sequences of mitochondrial cytochrome b gene. One lineage was predominant in the west and southern ranges of the populations studied and the other was commonly found in a more northern distribution (Murchison populations) in Uganda. Murchison and the geographically intermediate Toro populations (Uganda) represented the area of overlap. The existence of the two lineages in the area of overlap is hypothesized to have resulted from a range expansion and secondary contact of the two lineages of kob that evolved in allopatry. The existence of the kob during the Pleistocene offers a plausible explanation for the observed biogeographic pattern. Our mitochondrial data reveal two examples of discordance between a gene tree and presumed species tree as: (i) the two lineages co-occur in the kob subspecies, Kobus kob thomasi (Uganda kob); and (ii) the puku, which was included in the analysis because of its controversial taxonomic status (currently recognized as a distinct species from the kob), is paraphyletic with respect to the kob. Significant degrees of heterogeneity were detected between populations. Relatively high genetic variation was observed in the populations, however, the inclusion of distinct lineages influences the population structure and nucleotide diversity of the kob populations.     

Phylogeography, hybridization and Pleistocene refugia of the kob antelope (Kobus kob)

Mitochondrial DNA control region sequences and seven microsatellites were used to estimate the genetic structuring, evolutionary history and historic migration patterns of the kob antelope (Kobus kob). Ten populations were analysed, representing the three recognized K. kob subspecies: K. k. kob in west Africa, K. k. thomasi in Uganda and K. k. leucotis in Sudan and Ethiopia. Despite being classified as K. k. thomasi and being phenotypically identical to the kob in Queen Elizabeth National Park (NP), the Murchison Falls population in Uganda showed high genetic similarity with the phenotypically distinct K. k. leucotis populations in Sudan and Ethiopia. This was regardless of marker type. Pairwise comparisons and genetic distances between populations grouped Murchison with K. k. leucotis, as did the Bayesian analysis, which failed to find any genetic structuring within the group. We propose that the divergent phenotype and life-history adaptations of K. k. leucotis reflect the isolation of kob populations in refugia in west and east Africa during the Pleistocene. Subsequent dispersal has led to secondary contact and hybridization in northern Uganda between lineages, which was supported by high levels of genetic diversity in Murchison. The reduced variability observed in Queen Elizabeth NP reflects a small founder population from west Africa and in part the decimation of Uganda's wildlife during the country's political turmoil in the 1970s. Due to similarities in phenotype and ecology, and the joint evolutionary history of their mtDNA sequences, the taxonomic status of K. k. kob and K. k. thomasi as separate subspecies is called into question.     

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.     

Mating system and ecology of black Iechwe (Kobus: Bovidae) in Zambia

The mating system and ecology of black lechwe Kobus leche smithemani was studied in the Bangweulu Basin, northern Zambia. Black lechwe were migratory and concentrated at high density during the wet season on shallow water floodplain and peripheral grassland. Female lechwe were evenly distributed in small groups within the two major habitat types, and density was related to water depth and the quality of vegetation. A proportion of the adult male population defended small, contiguous, resource-based territories of approximately 1–2 ha in size. Particularly high density concentrations of male and female lechwe occurred on small raised areas, which remained dry in comparison to the inundated floodplain. These areas superficially resembled the leks of other reduncine antelope. However, the presence of vegetational resources and the comparatively low levels of agonistic and sexual behaviour suggested that raised areas were clusters of resource-based territories. We suggest that the absence of lek-breeding in this high density population of lechwe is related to the homogeneity of vegetational resources and the corresponding even distribution of females. This may have led to a reduction in male harassment of oestrous females and the increased ability of males to retain oestrous females within single resource territories.     

Early Pleistocene Spiral-Horned Antelopes (Artiodactyla,Bovidae) from the Taurida Cave (Crimea,Russia)

Paleontological Journal - The article describes fossil remains of two spiral-horned antelope species Gazellospira torticornis (Aymard, 1854) and Pontoceros ambiguus Vereschagin, Alexejeva, David et...     

Conservation of the puku antelope (Kobus vardoni, Livingstone) in the Kilombero Valley, Tanzania

Although there are populations of puku antelope Kobusvardoni (Livingstone) scattered throughout eastern and centralAfrica, it is estimated that 75% of the total population is now restricted tothe Kilombero Valley, Tanzania. The Kilombero Valley is an area rich inagricultural potential and natural resources; wildlife populations are also highand the inner valley is a Game Controlled Area, although this only provideslimited protection. Aerial surveys during the 1989, 1994 and 1998 dry seasonsshowed the puku population to be stable at around50000–60000 animals. Livestock populations fluctuated, butincreased from 17309 ± 6487 to 54047 ±17247 over the same period. Signs of human activity (e.g. huts, fieldsand livestock) were highest around the edge of the Game Controlled Area,indicating intense pressure on 'boundary-zone' habitats at thefloodplain–woodland interface. Puku use 'boundary-zone'habitats during the wet season when large areas of grassland are flooded.Potential threats to the puku population are therefore likely due to habitatdegradation through over-grazing by domestic herbivores, agriculturalencroachment, and the expansion of human settlements. Licensed trophyhunting probably has a negligible impact on puku because of very low off-take, but illegalhunting represents a serious threat near human settlements during thewet season and in accessible parts of the floodplain during the dry season.     

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。     

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.