Phenetic Analysis of Aboriginal and Introduced Populations of Sable (Martes zibellina) in Russia

Using standard and multivariate statistic methods, an epigenetic character—foramina in fossa condyloidei inferior(FFCI)—was studied in sable populations. This character was shown to be most frequent in southeastern populations of the species (Primorye and the Baikal region) while its distribution in the remaining part of the range was polyclinal. The expression of FFCI was directly associated with coat color and longitude, and inversely associated with skull size. This trend was broken by some western populations formed in the 1950s by introduction, which exhibited stable morphological differences with adjacent aboriginal sable populations. Most populations of the species exhibit differences in the manifestation of the studied characters. Frequency of the FFCI manifestation can be used as an additional population characteristic, an associative diagnostic character that shows high discriminating capability in detecting phenogenetic relationships of intraspecific groups.     

Breeding of the Russian sable: Stages of industrial domestication and genetic variability

Creating farms for sable breeding was associated with the commercial destruction of natural populations and, consequently, the overall decline in the species number. The gene pool of the first farm-bred sable population in Russia, established in the vicinity of Moscow (“Pushkinskiy” fur farm), was formed by crossing of animals removed from nine natural populations. In the first eight years of farm operation, approximately one thousand animals were used for sable breeding; some of these animals were able to adapt to the farm management and, subsequently, to the selection for a number of quantitative traits in the period of industrial domestication. It took about ten years for breeders to work out the breeding and selection technologies, which became successfully employed in the established affiliated sable breeding farms. The main achievement in sable breeding over the 85-year historical period of breeding in Russia is the creation of two unique breeds, black sable (1969) and Saltykovskaya 1 (2007). In general, industrial domestication in fur farming and the subsequent breeding works made the fur of many species (mink, fox, Arctic fox) obtained from natural populations uncompetitive, which undoubtedly reduced the hunting interest in the animals living in the wild. Consequently, hunting for fur-bearing animals of most species decreased and has only local importance. Owing to the specific features of sable biology, the fur of farm-bred animals cannot yet completely replace the furs obtained by hunting; however, the farm-bred sable population is constantly growing. This review presents the results of the analysis of the level of genetic variability in natural and farm populations at nuclear and mitochondrial loci. The comparative analysis makes it possible to estimate the loss of genetic diversity upon the species adaptation to the new conditions of existence.     

A comparative analysis of craniological variability of Eurasian (sable and pine marten) and North American (American marten) species of the <Emphasis Type="Italic">Martes</Emphasis> genus

This paper studies the craniological variability of closely related species—the pine marten Martes martes, the American marten M. americana, and the sable M. zibellina. The study included the data on 22 nonmetric characters of skulls in 2344 individuals. The studied Martes species were revealed to have an evident species-specific manifestation of the investigated craniological characters. Sables stand apart most of all in the interspecific comparison of populations. The degree of epigenetic variability is somewhat higher in the sable than in the pine marten. The American marten significantly differs from Eurasian species in the manifestation of the nonmetric skull characters, which is noted when analyzing even a small number of samples.     

Microsatellite analysis of two captive populations of sable (Martes zibellina L.)

The high value of sable (Martes zibellina L.) fur and stable demand for it over the centuries have led to suboptimal hunting patterns and, as a result, considerable fluctuations in the sizes of natural populations of this species. To maintain the traditional export of sable fur, efforts towards commercial domestication of sable have been made in Russia. The first farm population of sable consisted of animal from eight natural populations in 1929. After the problems related to breeding in captivity were solved, directional selection began. Eighty years of breeding have resulted in sable herds with homogeneous quantitative characters. Prospects for further breeding depend on the current level of genetic diversity in the captive populations of sables formed during the first stages of domestication. The sable populations of the Pushkinsky and Saltykovsky fur farms located in Moscow oblast, which were the objects of this study, are the progenitors of the existing captive populations. The first estimation of genetic variation of this species by means of a panel of microsatellite markers was developed for this study. Two captive sable populations were analyzed using ten microsatellite loci; a total of 75 alleles were found in both populations. Population-specific alleles were identified (6 and 13 in the Pushkinsky and Saltykovsky populations, respectively). The populations studied were found to be differentiated with respect to four microsatellite loci.     

Microsatellite analysis of two captive populations of sable (<Emphasis Type="Italic">Martes zibellina</Emphasis> L.)

The high value of sable (Martes zibellina L.) fur and stable demand for it over the centuries have led to suboptimal hunting patterns and, as a result, considerable fluctuations in the sizes of natural populations of this species. To maintain the traditional export of sable fur, efforts towards commercial domestication of sable have been made in Russia. The first farm population of sable consisted of animal from eight natural populations was founded in 1929. After the problems related to breeding in captivity were solved, directional selection began. Eighty years of breeding have resulted in sable herds with homogeneous quantitative characters. Prospects for further breeding depend on the current level of genetic diversity in the captive populations of sables formed during the first stages of domestication. The sable populations of the Pushkinsky and Saltykovsky fur farms located in Moscow oblast, which were the objects of this study, are the progenitors of the existing captive populations. The first estimation of genetic variation of this species by means of a panel of micro-satellite markers was developed for this study. Two captive sable populations were analyzed using ten micro-satellite loci; a total of 75 alleles were found in both populations. Population-specific alleles were identified (6 and 13 in the Pushkinsky and Saltykovsky populations, respectively). The populations studied were found to be differentiated with respect to four microsatellite loci.     

Genetic Effects of Sable (<Emphasis Type="Italic">Martes zibellina</Emphasis> L.) Reintroduction in Western Siberia

In the middle of the 20th century, massive introductions of sables were performed to recover the area of this valuable fur species. In this work, genetic variation of a naturalized sable population from the Vakh River basin (Nizhnevartovskiy district, Khanty-Mansi Autonomous Okrug) was investigated. This population developed as a result of sable introduction from Cisbaikalia in 1952?1957. The naturalized sable population of the Vakh basin occupies an intermediate position between two autochthonous sable populations from the Ob River area and Cisbaikalia, as assessed by variation of five microsatellite loci. Apparently, the genetic structure of the modern sable population from the Vakh basin was formed by mixing the gene pools of the original Cisbaikalian population and the neighboring autochthonous populations from the Ob River area which recovered their numbers in a natural way. Data on genetic variation in the naturalized sable population agree with the results of previous morphological studies and can be employed for long-term monitoring of the outcome of sable introduction.     

The Results of Sable (<Emphasis Type="Italic">Martes zibellina</Emphasis>) Reintroduction Demonstrate the Founder Effect

The relative abundance of intrapopulation groups with different parameters of skull size, coat color, and expression of an epigenetic cranial trait was compared in autochthonous, reintroduced, and donor populations of sable. Recovery of the species resources and broad variability of the phenotypic trait complex in the newly fomned populations were observed. A large proportion of the animals had the phenotype that included large size, dark coat color, and pronounced expression of a specific phene trait (foramen in the condylar fossa) and was not characteristic of the neighboring autochthonous populations. It is reasonable to attribute the presence of individuals with an unusual morphology in the newly formed populations of animals to a manifestation of the founder principle, because the effect of this principle was promoted by spatial isolation of the primary foci of translocated animals.     

Distributional niche of relatively rare sable antelope in a South African savanna: habitat versus biotic relationships

The geographic distribution of a species is governed by climatic conditions, topography, resources and habitat structure determining the fundamental niche, while the local distribution expressed via home range occupation may be compressed by biotic interactions with competitors and predators, restricting the realised niche. Biotic influences could be especially important for relatively rare species. We investigated how rainfall, geology, land type and abundance of other ungulate species serving as competitors or prey for predators contributed to the patchy distribution of sable antelope herds within Kruger National Park. Data were provided by annual aerial surveys of ungulate populations conducted between 1978 and 1988. Sable herds were more commonly present on granitic and sandstone substrates than on more fertile basalt. They occurred both in the moist south‐west and dry north of the park. They were most abundant in sour bushveld and mopane savanna woodland, and mostly absent from knob thorn‐marula parkland. The presence of sable was negatively associated with high concentrations of impala and wildebeest, less consistently related to the abundance of zebra, and positively associated with the occurrence of buffalo herds. Best supported models included the separate effects of the most abundant grazers along with land type. Interspecific relationships seemed more consistent with vulnerability to predation as the underlying mechanism restricting the distribution of sable herds than with competitive displacement. Sable favoured land types distinct from those where wildebeest, the most preferred prey of lions, and impala, numerically the most important resident prey species, were most abundant. Hence the risk of predation, associated with habitat conditions where abundant prey species are most concentrated, can exert an overriding influence on the distribution of rarer species in terms of their home range occupation.     

Mitochondrial DNA population structure of roan and sable antelope: implications for the translocation and conservation of the species

Mitochondrial DNA (mtDNA) control region sequences were analysed to determine the geographical genetic structure of Hippotragus niger (sable antelope) and H. equinus (roan antelope). Analyses by AMOVA, minimum evolution networks, parsimony, neighbour joining and maximum likelihood show that the roan antelope populations are geographically partitioned and that their phylogeography corresponds well with subspecific boundaries. In sharp contrast, however, our sequence data group all the sable antelope into two well-delineated maternal clades. One comprising specimens from the strictly east African H.n. roosevelti, and a second geographically diverse group which includes H.n. niger (South Africa, Zimbabwe), H.n. variani (Angola), and H.n. Kirkii (Zambia, Malawi). The differences in the genetic population structure suggest that distinct extrinsic and intrinsic factors have shaped the phylogeography of these two closely related antelope species. In the absence of obvious extrinsic barriers to gene flow, we suggest that behavioural differences may be responsible, at least in part, for the contrasting spatial patterns detected among geographical localities. Although our data reflect only the maternal phylogeny of these important game species, they nonetheless suggest that different management strategies should be applied. We recommend that roan antelope be translocated only within the currently defined subspecies boundaries, while in the case of sable antelope, conservation measures should focus on preserving the genetic integrity of the east African and southern African sable populations. Implicit in this is that translocations of animals between populations comprising these two distinct maternal genetic clades should be actively discouraged.     

Genetic structure of sable (Martes zibellina L.) in Eurasia—analysis of the mitochondrial lineages distribution

The phylogeography of the sable, which is a commercially valuable species, is extremely complicated and poorly investigated. Specifically, the effects of factors such as the range dynamics of the sable during the Pleistocene Epoch, the localization of glacial refugia, species distribution pattern in Holocene, and recent dramatic population decline, along with massive reacclimatization measures, on the species phylogeography remain unclear. Based on the sequence analysis of the control region of mitochondrial DNA from sables that inhabit different parts of the species range, a suggestion was made of the considerably high Pleistocene genetic diversity in sable, which was subsequently lost. The initial diversity of mitochondrial lineages is mostly preserved in the Urals, while in the eastern part of the range, it seems to have been depleted as early as before the last glacial maximum. On the other hand, the even greater depletion of the mitochondrial lineages observed in some populations of central Siberia can be associated with the dramatic population decline at the turn of the 20th century.     

基于MaxEnt模型的北极村国家级自然保护区紫貂栖息地适宜性评价

在人类支配的景观中,生境退化已经导致多个物种种群数量不断减少,分布范围不断缩减。紫貂（Martes zibellina）为国家Ⅰ级重点保护动物,种群数量稀少,开展栖息地适宜性研究工作对制定科学的栖息地保护计划至关重要。于2021年1月-2022年8月在黑龙江省北极村国家级自然保护区采用样线调查法、足迹链跟踪、远红外相机监测综合收集到紫貂和猎物（雪兔）活动点信息。利用最大熵（MaxEnt）栖息地建模分析方法,首次在多个分辨率尺度背景下对紫貂种群的栖息地适宜性进行评价,研究结果表明：（1）利用ArcGIS 10.4重采样后在6个分辨率尺度（30m、60m、120m、240m、480m、960m）进行栖息地建模分析,基于主要栖息地变量因子对模型的贡献率及稳定性影响,并综合考虑研究区域面积,最终选定30m分辨率尺度作为紫貂栖息地最佳分析建模尺度,在30m分辨率尺度栖息地预测模型的曲线下面积（AUC）值为0.881;（2）研究发现猎物资源、植被类型和地形变量是影响紫貂栖息地适宜性的主要变量因子：雪兔出现概率较高、距草地与河流较近、海拔约400-600m、距常绿针叶林1.5km、距落叶针叶林约200m、坡向为50-250°的区域为紫貂的适宜栖息地;（3）栖息地适宜性分析表明,北极村国家级自然保护区紫貂适宜栖息地和次适宜栖息地面积共计23.66km²,约占保护区的17.2%,主要集中在保护区中部,而东部和西北部区域,栖息地破碎化较严重。基于模型结果与野外调查,提出了三条建议：（1）应严格控制人为活动,避免因人类干扰造成不适宜栖息地面积的持续扩大;（2）建立生态廊道促进保护区西部与中部紫貂种群进行个体交流,降低紫貂种群局部区域灭绝概率;（3）对东部地区破碎化的栖息地进行修复,扩大东部适宜栖息地面积,使破碎化的栖息地连接为整体。为分布于我国最北端的紫貂种群恢复创造条件,这对于构建该地区相对稳定的生物多样性保护空间格局有着重要意义。     

Changes in correlations between commercially valuable characters of the sable Martes zibellina L. during artificial selection

A farm population of the sable Martes zibellina L. has been selected for darker coat color during 40 years. Correlations between fitness characters and correlations of these characters with the selected character have been monitored. Correlation analysis has shown that the female fertility in the first year of reproduction is a promising predictor of how valuable the female will be for further breeding. Artificial directional selection has been shown to change the correlations that have been formed in natural populations of the sable. The relationship of this phenomenon with a decrease in the overall fitness during selection that has been observed in the sable population is discussed.     

Genetic analysis of sable (<Emphasis Type="Italic">Martes zibellina</Emphasis>) and pine marten (<Emphasis Type="Italic">M. martes</Emphasis>) populations in sympatric part of distribution area in the northern Urals

Analysis of nucleotide sequences of the mitochondrial DNA (mtDNA) control region (495 bp) of sables (Martes zibellina) and pine martens (M. martes) from allopatric parts of the species ranges has shown a considerable interspecific genetic distance (>3%). In sympatric populations of these species in the northern Urals, differences between two species-specific mtDNA lineages are still large; however, classification of each individual nucleotide sequence with one of the two lineages is not correlated with whether the given animal is phenotypically a sable, a pine marten, or a potential hybrid (the so-called “kidas”). This indicates a high degree of reciprocal introgression of the sable and pine marten mtDNA in the northern Urals and suggests that their interspecific hybridization is common in the sympatric zone.     

Resource use and the nutritional status of sable antelope in the Okavango Delta region of northern Botswana

The resource‐use patterns and nutritional status of sable antelope herds were investigated in the Okavango Delta region of northern Botswana for comparison with those documented for declining sable antelope populations elsewhere in southern Africa. GPS collars recorded the relative use of floodplain, upland and wooded grassland habitats by the sable herds while VHF beacons facilitated locating the herds for direct observations on feeding. Surprisingly, the sable herds made greatest use of upland grasslands, rather than the floodplain grasslands exposed after floodwater had receded, during the dry season months. In the upland grasslands, they exploited tall, fibrous grass species that retained quite high levels of greenness through the dry season. This ability, together with partial use of the floodplain and some browsing on new leaves and flowers, helped maintain dietary nitrogen and phosphorus levels, as indicated by faecal nutrient levels, above maintenance thresholds through the dry season. Hence, the sable herds in the study region did not seem to be limited nutritionally under the rainfall and flooding conditions prevailing during the study.     

Genetic variability of sable Martes zibellina L., pine marten M. martes L., and their hybrids in Western Siberia: Protein and DNA polymorphism

Using four types of markers, the genetic variability of sable and pine marten inhabiting Western Siberia was examined. Izoenzyme and restriction endonuclease analysis of the mtDNA cytochrome b gene fragment, as well as the ISSR-PCR and analysis of microsatellite variation, revealed a low differentiation level of sable and pine marten and confirmed the hybrid origin of atypical representatives of these species. The hybrids were characterized by an increased heterozygosity level and were genetically closer to sable than to pine marten. In atypical martens, the presence of mtDNA haplotypes of eastern sable was identified. This could be the consequence of the reintroduction of the Barguzin sable in the 20th century. In Western Siberia, the introgression of genes between sable and pine marten was massive and symmetrical. It apparently occurred in the past and continues in the present.     

Genetic individualization of sable (Martes zibellina L. 1758) using microsatellites

Genetic individualization based on non-invasive sampling is crucial for estimating the numbers of individuals in endangered mammalian populations. In sable (Martes zibellina)-poaching cases, identifying the number of animals involved is critical for determining the penalty. In addition, investigating animal numbers for wild sable populations requires genetic individualization when collecting several samples in neighboring regions. Microsatellites have been demonstrated to be reliable markers for individual identification. Thirty-three microsatellite loci derived from Mustelidae were selected to develop a genetic individualization method for sable. Three reference populations containing 54 unrelated sables were used to calculate allele number, allelic frequencies, and the polymorphic information content of each locus. The data were subsequently used to assess the validity of a combination of twelve loci for sable individualization. We defined twelve polymorphic loci that were easy to be amplified and genotyped. Four significant deviations from Hardy-Weinberg equilibrium were observed among the 12?loci in the three populations. The match probability of an individual from the reference populations with a random individual based on the 12?loci was 1.37?×?10^?13. Using the combination of the twelve loci provides sufficient power to individualize sables considering the levels of microsatellite polymorphism observed. These loci were successfully applied to a case of sable poaching and provided valid evidence to determine the penalty. The genetic individualization of sable based on these loci might also be useful to investigate the numbers of animals in wild populations.     

Comparisons among selected neonatal biomedical parameters of four species of semi-free ranging Hippotragini: Addax (Addax nasomaculatus), scimitar-horned oryx (Oryx dammah), Arabian oryx (Oryx leucoryx), and sable antelope (Hippotragus niger)

Basic biomedical data from 164 neonates of four species of the tribe Hippotragini, addax (Addax nasomaculatus), scimitar-horned oryx (Oryx dammah), Arabian oryx (Oryx leucoryx), and sable antelope (Hippotragus niger), were compared at one zoological institution over a 9-year period. Measured biomedical parameters included body weight, temperature, pulse and respiratory rates, packed cell volume (PCV), total plasma protein, glucose, IgG assessment via zinc sulfate turbidity, and white blood cell count with differential. All species were maintained in a semi-free ranging setting with the same diet, available shelter, and opportunity for social interaction. Based on clinical and field observations, all neonates used in the study were believed to be at least 24 hr old, to have bonded with the dam, and to have no obvious physical abnormalities. Median body weights were similar only in the addax and Arabian oryx with sable antelope having the largest median body weight. No significant differences in rectal temperatures or pulse rates were found among species. Median respiratory rates were similar between certain groups. Arabian oryx and scimitar-horned oryx shared the highest packed cell volumes while the sable antelope had the lowest. Sable antelope had the highest median total plasma protein with no significant differences among the other species. Sable were also significantly lower in median blood glucose than the three other Hippotraginae. Zinc sulfate turbidities in all species were similar. Addax had higher median total white blood cell counts than sable. No significant differences in the median numbers of segmented neutrophils, band neutrophils, and eosinophils were detected among species. Basophils were only found in the scimitar-horned oryx in one animal. Addax had higher median lymphocyte counts than sable and Arabian oryx as well as higher median monocyte counts than sable. All four species exhibited higher median counts of neutrophils compared with lymphocytes. The biomedical differences observed highlight the importance of having an accurate database of clinical normal values against which to evaluate neonatal health. Zoo Biol 20:47-54, 2001. Copyright 2001 Wiley-Liss, Inc.     

An exceptional case of historical outbreeding in African sable antelope populations

Empirical investigations of intraspecific outbreeding and subsequent introgressive hybridization in natural populations are rare, particularly among conspecific populations of large mammals. Using mitochondrial DNA data [partial control region (496 basepairs - bp) and cytochrome b gene (343 bp) sequences analysed from 95 individuals representing 17 sampling locations scattered through the African miombo (Brachystegia) woodland ecosystem] and phylogeographical statistical procedures (gene genealogy, nested cladistic and admixture proportion analyses), we (i) give a detailed dissection of the geographical genetic structure of Hippotragus niger; (ii) infer the processes and events potentially involved in the population history; and (iii) trace extensive introgressive hybridization in the species. The present-day sable antelope population shows a tripartite pattern of genetic subdivision representing West Tanzanian, Kenya/East Tanzanian and Southern Africa locations. Nested clade analysis revealed that past allopatric fragmentation, caused probably by habitat discontinuities associated with the East African Rift Valley system, together with intermediary episodic long-distance colonization and restricted, recurrent gene flow have played an predominant role in shaping the extent of maternal genetic diversity (10.4%) and population structure. An extensive (average rate of admixture = 20.0%), but geographically circumscribed and unidirectional hybridization event in the past was inferred, resulting in an extreme (the highest discovered so far in mammals) intraspecific difference of 18.2% among morphologically monotypic sable antelopes from West Tanzania. The results are used to provide an evolutionary framework within which taxonomic implications and conservation decisions can be evaluated.     

Character shifts in the defensive armor of sympatric sticklebacks

Natural enemies may contribute to the morphological divergence of sympatric species, yet their role has received little attention to date. We tested for character shifts in defensive armor of sympatric threespine sticklebacks (Gasterosteus aculeatus complex) previously shown to exhibit ecological character displacement in traits related to resource use. We scored five defensive armor traits in sympatric benthic and limnetic stickleback species from southwestern British Columbia and compared them with the same traits in nearby allopatric populations in the presence of the same predatory fish (Oncorhynchus sp.). This approach is analogous to tests of ecological character displacement that compare trophic traits of sympatric and allopatric species in the presence of the same community of resource types. Three patterns consistent with character displacement in defensive armor were found. First, limnetics in different lakes had consistently more armor than sympatric benthics. Second, the average amount of armor, averaged over both species, was reduced in sympatry compared to allopatric populations. This reduction was almost entirely the result of shifts by benthic species, whereas armor in limnetics was more similar to that in allopatric populations. Third, differences between sympatric benthics and limnetics in total armor were greater than expected from comparisons with allopatric populations. We interpret these patterns as the result of differences in habitat-specific predation regimes accompanying ecological character displacement and indirect interactions between sympatric stickleback species mediated by their top predators. These results suggest that predation may facilitate, rather than hinder, the process of divergence in sympatry.     

Phylogeography of sable (Martes zibellina L. 1758) in the southeast portion of its range based on mitochondrial DNA variation: highlighting the evolutionary history of the sable

To reveal phylogeographic features of sable (Martes zibellina) in the southeast part of its range, we analyzed variability of the mitochondrial DNA (mtDNA) cytochrome b gene, tRNA (Pro), tRNA (Thr) and control region (D-loop) sequences from 78 specimens in populations of the Russian Far East, northeast China, and Mongolia. Our results revealed the presence of 49 different haplotypes split into two major phylogenetic groups—clades A and B, the latter separated into two clades, B1 and B2. Comparative analysis of D-loop haplotypes in populations originating from the southeast (Russian Far East, China and Mongolia) and the west (northern Urals) portions of sable range indicated that all three mtDNA clades were present in different regional groups. However, highest diversity of clade B1 in northeast China and its nearly complete absence from the Urals suggest that the southeast sable range, being a refuge during Pleistocene glacial periods, can be considered the center of genetic diversification and possibly origin of this species. All divergence estimates fall within the Pleistocene suggesting that Quarternary glaciations played an important role in phylogeographic differentiation of sable.