Changes in the skull morphology of the Arctic wolf, Canis lupus arctos, during the twentieth century

Measurements of a large series of skulls of the Arctic wolf, Canis lupus arctos . have shown that since 1930 there has been an overall reduction in the size of the skulls, together with widening of the cranium, shortening of the facial region, and reduction in size of the teeth. This suggests that hybridization and subsequent introgression occurred with huskies ( Canis familiaris ) during the 1930s, which is consistent with historical accounts. Since 1950 there has been a reversion in skull morphology to a more 'wolf-like' form, suggesting that hybridization is no longer occurring.
The skull of a wolf/dog hybrid is intermediate in size between the skulls of wolves and huskies but its shape is allometrically dissimilar. Skulls of wolves from the period 1930–50 are moi-e similar to the skull of this hybrid than in the other time periods.
The skull of a male canid from a carcass collected on Ellesmere Island and presented to the Natural History Museum. London, in 1986 was at first thought to be from a wolf/dog hybrid but analyses of the measurements show that it is more likely to be from an Arctic wolf with severe abnormalities to the jaws.     

Prey specialization may influence patterns of gene flow in wolves of the Canadian Northwest

This study characterizes population genetic structure among grey wolves (Canis lupus) in northwestern Canada, and discusses potential physical and biological determinants of this structure. Four hundred and ninety-one grey wolves, from nine regions in the Yukon, Northwest Territories and British Columbia, were genotyped using nine microsatellite loci. Results indicate that wolf gene flow is reduced significantly across the Mackenzie River, most likely due to the north-south migration patterns of the barren-ground caribou herds that flank it. Furthermore, although Banks and Victoria Island wolves are genetically similar, they are distinct from mainland wolf populations across the Amundsen Gulf. However, low-level island-mainland wolf migration may occur in conjunction with the movements of the Dolphin-Union caribou herd. Whereas previous authors have examined isolation-by-distance in wolves, this study is the first to demonstrate correlations between genetic structure of wolf populations and the presence of topographical barriers between them. Perhaps most interesting is the possibility that these barriers reflect prey specialization by wolves in different regions.     

周口店直立人3号与5号头骨形态特征对比及其演化速率所反映的群体隔离

德日进认为周口店直立人保持着形态上的稳定性; 这种进化上的缓慢性表明周口店直立人进化过程中无论何时都未曾发生过外来移民的闯入, 从而打乱、干扰或推进其进化过程。本项研究采用三维结构重叠和精确测量数据的方式对比了周口店第一地点晚期代表周口店直立人(ZKD)5号头骨相对于早期代表ZKD3的形态特征演化变化, 并与南京直立人(NJ)1和2号头骨之间的差别进行了对比。结果表明, 周口店晚期标本相对于早期标本来说, 头骨尺寸在各个方向上都有所增加, 但轮廓形状基本保持一致, 与同期的南京古人类的相比, 周口店直立人的演化速率显得很慢。这很可能是由于缺乏基因交流的缘故; 周口店直立人群有可能是一个相对隔离的群体。本项研究支持德日进等关于周口店直立人群缺乏基因交流的观点。     

Ecological factors drive differentiation in wolves from British Columbia

Aim Limited population structure is predicted for vagile, generalist species, such as the grey wolf (Canis lupus L.). Our aims were to study how genetic variability of grey wolves was distributed in an area comprising different habitats that lay within the potential dispersal range of an individual and to make inferences about the impact of ecology on population structure. Location British Columbia, Canada – which is characterized by a continuum of biogeoclimatic zones across which grey wolves are distributed – and adjacent areas in both Canada and Alaska, United States. Methods We obtained mitochondrial DNA control region sequences from grey wolves from across the province and integrated our genetic results with data on phenotype, behaviour and ecology (distance, habitat and prey composition). We also compared the genetic diversity and differentiation of British Columbia grey wolves with those of other North American wolf populations. Results We found strong genetic differentiation between adjacent populations of grey wolves from coastal and inland British Columbia. We show that the most likely factor explaining this differentiation is habitat discontinuity between the coastal and interior regions of British Columbia, as opposed to geographic distance or physical barriers to dispersal. We hypothesize that dispersing grey wolves select habitats similar to the one in which they were reared, and that this differentiation is maintained largely through behavioural mechanisms. Main conclusions The identification of strong genetic structure on a scale within the dispersing capabilities of an individual suggests that ecological factors are driving wolf differentiation in British Columbia. Coastal wolves are highly distinct and representative of a unique ecosystem, whereas inland British Columbia grey wolves are more similar to adjacent populations of wolves located in Alaska, Alberta and Northwest Territories. Given their unique ecological, morphological, behavioural and genetic characteristics, grey wolves of coastal British Columbia should be considered an Evolutionary Significant Unit (ESU) and, consequently, warrant special conservation status. If ecology can drive differentiation in a highly mobile generalist such as the grey wolf, ecology probably drives differentiation in many other species as well.     

Megafaunal extinctions and the disappearance of a specialized wolf ecomorph

The gray wolf (Canis lupus) is one of the few large predators to survive the Late Pleistocene megafaunal extinctions [1]. Nevertheless, wolves disappeared from northern North America in the Late Pleistocene, suggesting they were affected by factors that eliminated other species. Using skeletal material collected from Pleistocene permafrost deposits of eastern Beringia, we present a comprehensive analysis of an extinct vertebrate by exploring genetic (mtDNA), morphologic, and isotopic (delta(13)C, delta(15)N) data to reveal the evolutionary relationships, as well as diet and feeding behavior, of ancient wolves. Remarkably, the Late Pleistocene wolves are genetically unique and morphologically distinct. None of the 16 mtDNA haplotypes recovered from a sample of 20 Pleistocene eastern-Beringian wolves was shared with any modern wolf, and instead they appear most closely related to Late Pleistocene wolves of Eurasia. Moreover, skull shape, tooth wear, and isotopic data suggest that eastern-Beringian wolves were specialized hunters and scavengers of extinct megafauna. Thus, a previously unrecognized, uniquely adapted, and genetically distinct wolf ecomorph suffered extinction in the Late Pleistocene, along with other megafauna. Consequently, the survival of the species in North America depended on the presence of more generalized forms elsewhere.     

中国狼不同地理种群遗传多样性及系统进化分析

为了解中国狼不同地理种群遗传多样性及系统发育情况,从中国境内狼的主要分布区青海、新疆、内蒙古和吉林4个地区采集样品,用分子生物学技术手段成功地获得44个个体线粒体DNA控制区第一高变区(HVRⅠ)序列和40个线粒体Cyt b部分序列。线粒体控制区HVRⅠ共检测到51个变异位点,位点变异率为8.76%;线粒体Cyt b部分序列发现31个变异位点,位点变异率为5.33%,未见插入及缺失现象,变异类型全部为碱基置换。共定义了16个线粒体HVRⅠ单倍型,其中吉林与内蒙种群存在共享单倍型,估计这两地间种群亲缘关系较近。4个地理种群中新疆种群拥有较高的遗传多样性(0.94)。中国狼种群总体平均核苷酸多态性为2.27%,与世界其他国家地区相比,中国狼种群拥有相对较高的遗传多样性。通过线粒体HVRⅠ单倍型构建的系统进化树可以看出,中国狼在进化上分为2大支,其中位于青藏高原的青海种群独立为一支,推测其可能长期作为独立种群进化。基于青海种群与新疆,内蒙种群的线粒体Cyt b遗传距离,推测中国狼2个世系可能在更新世冰川时期青藏高原受地质作用急速隆起后出现分歧,分歧时间大约在1.1 MY前。     

Biogeographic variation in skull morphology across the Kra Isthmus in dusky leaf monkeys

Despite the growing literature on the underlying factors of geographical phenotypic variation, little is known about how and to what extent biogeographic barriers in South‐East Asia have shaped morphological variation in primates. We aimed to investigate the geographical variations in skull morphology in dusky leaf monkeys by decomposing them into clinal (latitudinal), non‐clinal spatial (discrete difference between regions north and south of the Isthmus of Kra), and environment‐related components. We applied geometric morphometrics to measure 53 adult male specimens from 36 localities, covering the regions both north and south of the Isthmus of Kra. A linear model was used to test the effects of region (north vs. south of the Isthmus of Kra), latitude, and environmental factors (temperature and rainfall) on the size and shape of skulls. A part of variation in skull shape differed moderately between the regions in the north and south of the Isthmus of Kra, and this difference cannot be explained by latitudinal and environmental factors. However, for size and the majority of variations in shape, we detected limited contributions of region and the two environmental factors. Shape differentiation that was unexplained by latitudinal and environmental factors suggests that dusky leaf monkeys may have experienced a population division due to habitat constriction around the Isthmus of Kra. However, this divergence probably has been obscured by subsequent gene flow between populations after habitat recovery.     

Cross‐continental comparison of parasite communities in a wide‐ranging carnivore suggests associations with prey diversity and host density

1. Parasites are integral to ecosystem functioning yet often overlooked. Improved understanding of host–parasite associations is important, particularly for wide‐ranging species for which host range shifts and climate change could alter host–parasite interactions and their effects on ecosystem function.
2. Among the most widely distributed mammals with diverse diets, gray wolves (Canis lupus) host parasites that are transmitted among canids and via prey species. Wolf–parasite associations may therefore influence the population dynamics and ecological functions of both wolves and their prey. Our goal was to identify large‐scale processes that shape host–parasite interactions across populations, with the wolf as a model organism.
3. By compiling data from various studies, we examined the fecal prevalence of gastrointestinal parasites in six wolf populations from two continents in relation to wolf density, diet diversity, and other ecological conditions.
4. As expected, we found that the fecal prevalence of parasites transmitted directly to wolves via contact with other canids or their excreta was positively associated with wolf density. Contrary to our expectations, the fecal prevalence of parasites transmitted via prey was negatively associated with prey diversity. We also found that parasite communities reflected landscape characteristics and specific prey items available to wolves.
5. Several parasite taxa identified in this study, including hookworms and coccidian protozoans, can cause morbidity and mortality in canids, especially in pups, or in combination with other stressors. The density–prevalence relationship for parasites with simple life cycles may reflect a regulatory role of gastrointestinal parasites on wolf populations. Our result that fecal prevalence of parasites was lower in wolves with more diverse diets could provide insight into the mechanisms by which biodiversity may regulate disease. A diverse suite of predator–prey interactions could regulate the effects of parasitism on prey populations and mitigate the transmission of infectious agents, including zoonoses, spread via trophic interactions.

     

Evidence of genetic distinction and long-term population decline in wolves (Canis lupus) in the Italian Apennines

Historical information suggests the occurrence of an extensive human-caused contraction in the distribution range of wolves (Canis lupus) during the last few centuries in Europe. Wolves disappeared from the Alps in the 1920s, and thereafter continued to decline in peninsular Italy until the 1970s, when approximately 100 individuals survived, isolated in the central Apennines. In this study we performed a coalescent analysis of multilocus DNA markers to infer patterns and timing of historical population changes in wolves surviving in the Apennines. This population showed a unique mitochondrial DNA control-region haplotype, the absence of private alleles and lower heterozygosity at microsatellite loci, as compared to other wolf populations. Multivariate, clustering and Bayesian assignment procedures consistently assigned all the wolf genotypes sampled in Italy to a single group, supporting their genetic distinction. Bottleneck tests showed evidences of population decline in the Italian wolves, but not in other populations. Results of a Bayesian coalescent model indicate that wolves in Italy underwent a 100- to 1000-fold population contraction over the past 2000-10,000 years. The population decline was stronger and longer in peninsular Italy than elsewhere in Europe, suggesting that wolves have apparently been genetically isolated for thousands of generations south of the Alps. Ice caps covering the Alps at the Last Glacial Maximum (c. 18,000 years before present), and the wide expansion of the Po River, which cut the alluvial plains throughout the Holocene, might have provided effective geographical barriers to wolf dispersal. More recently, the admixture of Alpine and Apennine wolf populations could have been prevented by deforestation, which was already widespread in the fifteenth century in northern Italy. This study suggests that, despite the high potential rates of dispersal and gene flow, local wolf populations may not have mixed for long periods of time.     

中国古人类进化连续性新辩

本文通过对一些形态特征的分析说明研究人类起源和进化应该注意的一些思维方法的问题,并且为多地区进化说增添了新的证据:1)中国更新世头骨共同具有的眼眶及其外侧下缘形状等特征在近代头骨中已有所改变,在考虑其意义时不应该忽略时间和基因交流起过的作用;2)中国更新世头骨的共同特征并不因为它们也可见于其他地区而丧失其意义,因为证明连续性的共同特征不需要是地区独有的;3)铲形门齿的结构在近代人中变弱,不削弱它作为连续进化证据的价值,这种弱化的现象是人类进化过程中形态纤巧化的一种表现;4)在辩论时不要对有关形态作不正确的观察和曲解对方的观点和论据;5)头骨正中矢状突隆、下颌圆枕、夹紧状鼻梁和第三臼齿先天性缺失等四项特征在近代非洲人中阙如,却存在于东亚近代人,本文的分析认为这种现象增强多地区进化说的说服力。     

高原鼢鼠(Eospalax baileyi)头骨形态的地理分化

运用单因素方差分析、主成分分析、判别分析和聚类分析,对高原鼢鼠(Eospalax baileyi)4个地区(临潭、天祝、玛曲和碌曲)8个地理群体的头骨形态特征进行综合分析,比较13个特征变量的地理分异,挖掘形态特征的主要变量,并探讨其形态变异与环境的关系。结果发现,群体间13个特征变量均存在显著差异,主成分分析表明13个头骨特征变量可用3个主成分来表述,其累计贡献率达96.981%。颅全长、上齿列长及齿隙长是高原鼢鼠头骨发生分化的主要变量。天祝种群与碌曲种群,及玛曲种群与临潭种群都有重叠,其中碌曲的两个种群相似性最高,玛曲种群和临潭种群与其他种群几乎无重叠,显示完全分化,聚类分析结果与判别分析结果一致。头骨形态与地理因子的相关性分析发现头骨形态大小与海拔成显著的正相关关系。综合分析认为地理隔离、栖息地海拔因子影响下的生态条件等是高原鼢鼠头骨分化的主要因素。     

A new mitochondrial haplotype confirms the distinctiveness of the Italian wolf (Canis lupus) population

In the past century the Italian wolf has been repeatedly indicated as a distinct subspecies, Canis lupus italicus, due to its unique morphology and its distinctive mtDNA control region (CR) monomorphism. However, recent studies on wolf x dog hybridization in Italy documented the presence of a second mtDNA CR haplotype (W16), previously found only in wolves from Eastern Europe, casting doubts on the genetic uniqueness of the Italian wolves. To test whether this second haplotype belongs to the Italian wolf population, we genotyped 92 wolf DNA samples from Italy, Slovenia, Greece and Bulgaria at four mtDNA regions (control-region, ATP6, COIII and ND4 genes) and at 39 autosomal microsatellites. Results confirm the presence of two mtDNA multi-fragment haplotypes (WH14 and WH19) in the Italian wolves, distinct from all the other European wolves. Network analyses of the multi-fragment mtDNA haplotypes identified two strongly differentiated clades, with the Italian wolf WH14 and WH19 multi-fragment haplotypes rooted together. Finally, Bayesian clustering clearly assigned all the wolves sampled in Italy to the Italian population, regardless of the two different multi-fragment haplotypes. These results demonstrate that the W16 CR haplotype is part of the genetic pool of the Italian wolf population, reconfirming its distinctiveness from other European wolves. Overall, considering the presence of unique mtDNA and Y-linked haplotypes, the sharply different frequencies of genome-wide autosomal alleles and the distinct morphological features of Italian wolves, we believe that this population should be considered a distinct subspecies.     

Population Viability, Nature Reserves, and the Outlook for Gray Wolf Conservation in North America

Theoretical work on population viability and extinction probabilities, empirical data from Canis lupus (gray wolf) populations, and expert opinion provide only general and conflicting conclusions about the number of wolves and the size of areas needed for conservation of wolf populations. There is no threshold population size or proven reserve design that guarantees long-term (century or more) survival for a gray wolf population. Most theoretical analyses of population viability have assumed a single, isolated population and lack of management intervention, neither of which is likely for wolves. Data on survival of actual wolf populations suggest greater resiliency than is indicated by theory. In our view, the previous theoretical treatments of population viability have not been appropriate to wolves, have contributed little to their conservation, and have created unnecessary dilemmas for wolf recovery programs by overstating the required population size. Nonetheless, viability as commonly understood may be problematic for small populations at the fringe of or outside the contiguous species range, unless they are part of a metapopulation. The capability of existing nature reserves to support viable wolf populations appears related to a variety of in situ circumstances, including size, shape and topography of the reserve; productivity, numbers, dispersion, and seasonal movement of prey; extent of poaching inside; degree of persecution outside; exposure to enzootica; attitudes of local people; and proximity to other wolf populations. We estimate that a population of 100 or more wolves and a reserve of several thousand square kilometers may be necessary to maintain a viable population in complete isolation, although 3000 km² or even 500–1000 km² may be adequate under favorable circumstances. In most cases, management intervention is probably necessary to assure the viability of relatively small, isolated populations. Because most reserves may be inadequate by themselves to ensure the long-term survival of wolf populations, favorable human attitudes toward the species and its management must be recognized as paramount, and cooperation of neighboring management jurisdictions will be increasingly important.     

Evidence of wolf dispersal in anthropogenic habitats of the Polish Carpathian Mountains

In the course of their maturation, most young wolves leave their natal pack and disperse in search for mating partners, improved food availability and new territories. We investigated whether this dispersal is affected by anthropogenic infrastructure in a 5,000 km2 area of the eastern region of the Polish Carpathian Mountains occupied by wolves. A radio-collared male wolf covered 230 km while dispersing through forested hills and densely populated valleys. To test if such dispersal is common in the population we analysed by microsatellite genotyping 39 samples taken from live-trapped wolves or wolves found dead in the study area. Although the obtained genotypes were assigned to different clusters in Bayesian tests, we could not ascribe this structure to landscape features, but rather to shared ancestry of wolf individuals found in distant locations. Moreover, we could not detect a spatial genetic structure in the wolf population, indicating a random occurrence of genotypes within the study area. Observation of the dispersing wolf and the absence of spatial genetic structure imply that wolves are still able to roam the entire area despite high densities of roads and a dense human population. Thus, we concluded that the existing anthropogenic infrastructure does not restrict wolf dispersal in the area and the studied wolves represent a coherent part of the Polish Carpathian wolf population.     

Geographic Variation in the Skull Morphometry of Four Populations of Batrachuperus karlschmidti(Urodela: Hynobiidae)

Geographic variation of morphology is an important topic of evolutionary biology, and research on geographic variation can provide insights on the formation, evolution, and adaptation of species and subspecies. The vertebrate skull is a developmentally and functionally complex morphological structure with multiple functions, that is susceptible to vary according to selection pressure. In this study, geographic variations in skull morphology of Batrachuperus karlschmidti from four different geographic populations(Shade, Gexi,Shangluokema, and Xinduqiao) were examined via geometric morphometrics. No significant differences were found among these populations with regard to skull size; however, significant variation was found in skull shape. The most notable shape changes are the relative sizes and positions of the frontal, maxilla,pterygoid, and vomer. Skull shape changes were not related to allometry. However, due to limitation of sample populations and size, the results of this study need to be further verified by more sample populations and individuals in the future. The results of this study contribute to our knowledge about these aspects of morphological variability in this species as well as in hynobiid salamanders.     

Extinct Beringian wolf morphotype found in the continental U.S. has implications for wolf migration and evolution

Pleistocene diversity was much higher than today, for example there were three distinct wolf morphotypes (dire, gray, Beringian) in North America versus one today (gray). Previous fossil evidence suggested that these three groups overlapped ecologically, but split the landscape geographically. The Natural Trap Cave (NTC) fossil site in Wyoming, USA is an ideally placed late Pleistocene site to study the geographical movement of species from northern to middle North America before, during, and after the last glacial maximum. Until now, it has been unclear what type of wolf was present at NTC. We analyzed morphometrics of three wolf groups (dire, extant North American gray, Alaskan Beringian) to determine which wolves were present at NTC and what this indicates about wolf diversity and migration in Pleistocene North America. Results show NTC wolves group with Alaskan Beringian wolves. This provides the first morphological evidence for Beringian wolves in mid‐continental North America. Their location at NTC and their radiocarbon ages suggest that they followed a temporary channel through the glaciers. Results suggest high levels of competition and diversity in Pleistocene North American wolves. The presence of mid‐continental Beringian morphotypes adds important data for untangling the history of immigration and evolution of Canis in North America.     

Spatial Heterogeneity in Human Activities Favors the Persistence of Wolves in Agroecosystems

As human populations expand, there is increasing demand and pressure for land. Under this scenario, behavioural flexibility and adaptation become important processes leading to the persistence of large carnivores in human-dominated landscapes such as agroecosystems. A growing interest has recently emerged on the outcome of the coexistence between wolves and humans in these systems. It has been suggested that spatial heterogeneity in human activities would be a major environmental factor modulating vulnerability and persistence of this contentious species in agroecosystems. Here, we combined information from 35 den sites detected between 2011 and 2012 in agroecosystems of western Iran (Hamedan province), a set of environmental variables measured at landscape and fine spatial scales, and generalized linear models to identify patterns of den site selection by wolves in a highly-modified agroecosystem. On a landscape level, wolves selected a mixture of rangelands with scattered dry-farms on hillsides (showing a low human use) to locate their dens, avoiding areas with high densities of settlements and primary roads. On a fine spatial scale, wolves primarily excavated dens into the sides of elevated steep-slope hills with availability of water bodies in the vicinity of den sites, and wolves were relegated to dig in places with coarse-soil particles. Our results suggest that vulnerability of wolves in human-dominated landscapes could be compensated by the existence of spatial heterogeneity in human activities. Such heterogeneity would favor wolf persistence in agroecosystems favoring a land sharing model of coexistence between wolves and people.     

Population structure and gene flow in a newly harvested gray wolf (<Emphasis Type="Italic">Canis lupus</Emphasis>) population

The genetic effects of harvest may be especially important in species that form social groups, such as gray wolves (Canis lupus). Though much research exists on the ecology and population dynamics of gray wolves, little research has focused on how anthropogenic harvest relates to the genetics of wolf populations. To analyze the short-term genetic consequences of the first two years of public wolf harvest in Minnesota following delisting under the Endangered Species Act, we genotyped harvested individuals at 18 microsatellite loci and quantified changes in population genetic structure and diversity in the first post-harvest year. If the harvest rate was high enough to create detectable genetic changes, population structure and differentiation between clusters could both increase because of decreased natal dispersal and increased disperser mortality, or they could decrease because of increased immigration from outside the population. In the Minnesota population, heterozygosity and allelic richness were not significantly different between years. However, population genetic structure increased and effective migration decreased among the sampled wolves. While the role of anthropogenic harvest in these changes cannot be distinguished from other confounding factors, this analysis suggests that harvest has a non-negligible effect and indicates the need for continued study to determine whether harvest-induced changes in genetic structure affect the evolutionary trajectory of harvested populations.