Reviving the African Wolf Canis lupus lupaster in North and West Africa: A Mitochondrial Lineage Ranging More than 6,000 km Wide

The recent discovery of a lineage of gray wolf in North-East Africa suggests the presence of a cryptic canid on the continent, the African wolf Canis lupus lupaster. We analyzed the mtDNA diversity (cytochrome b and control region) of a series of African Canis including wolf-like animals from North and West Africa. Our objectives were to assess the actual range of C. l. lupaster, to further estimate the genetic characteristics and demographic history of its lineage, and to question its taxonomic delineation from the golden jackal C. aureus, with which it has been considered synonymous. We confirmed the existence of four distinct lineages within the gray wolf, including C. lupus/familiaris (Holarctic wolves and dogs), C. l. pallipes, C. l. chanco and C. l. lupaster. Taxonomic assignment procedures identified wolf-like individuals from Algeria, Mali and Senegal, as belonging to C. l. lupaster, expanding its known distribution c. 6,000 km to the west. We estimated that the African wolf lineage (i) had the highest level of genetic diversity within C. lupus, (ii) coalesced during the Late Pleistocene, contemporaneously with Holarctic wolves and dogs, and (iii) had an effective population size of c. 80,000 females. Our results suggest that the African wolf is a relatively ancient gray wolf lineage with a fairly large, past effective population size, as also suggested by the Pleistocene fossil record. Unique field observations in Senegal allowed us to provide a morphological and behavioral diagnosis of the African wolf that clearly distinguished it from the sympatric golden jackal. However, the detection of C. l. lupaster mtDNA haplotypes in C. aureus from Senegal brings the delineation between the African wolf and the golden jackal into question. In terms of conservation, it appears urgent to further characterize the status of the African wolf with regard to the African golden jackal.     

Phylogeography of the Golden Jackal (Canis aureus) in India

The golden jackal (Canis aureus) is one of the most common and widely distributed carnivores in India but phylogeographic studies on the species have been limited across its range. Recent studies have observed absence of mitochondrial (mt) DNA diversity in European populations while some North African populations of golden jackal were found to carry gray wolf (Canis lupus lupaster) mtDNA lineages. In the present study, we sequenced 440 basepairs (bp) of control region (CR) and 412 bp of cytochrome b (cyt b) gene of mtDNA from 62 golden jackals sampled from India (n = 55), Israel (n = 2) and Bulgaria (n = 5), to obtain a total of eighteen haplotypes, comprising sixteen from India and one each from Israel and Bulgaria. Except for three previously described haplotypes represented by one cyt b and one CR haplotype both from India, and one CR haplotype from Bulgaria, all haplotypes identified in this study are new. Genetic diversity was high in golden jackals compared to that reported for other canids in India. Unlike the paraphyletic status of African conspecifics with the gray wolf, the Indian (and other Eurasian) golden jackal clustered in a distinct but shallow monophyletic clade, displaying no evidence of admixture with sympatric and related gray wolf and domestic dog clades in the region. Phylogeographic analyses indicated no clear pattern of genetic structuring of the golden jackal haplotypes and the median joining network revealed a star-shaped polytomy indicative of recent expansion of the species from India. Indian haplotypes were observed to be interior and thus ancestral compared to haplotypes from Europe and Israel, which were peripheral and hence more derived. Molecular tests for demographic expansion confirmed a recent event of expansion of golden jackals in the Indian subcontinent, which can be traced back ~ 37,000 years ago during the late Pleistocene. Our results suggest that golden jackals have had a potentially longer evolutionary history in India than in other parts of the world, although further sampling from Africa, the Middle East and south-east Asia is needed to test this hypothesis.     

A cryptic mitochondrial DNA link between North European and West African dogs

Domestic dogs have an ancient origin and a long history in Africa. Nevertheless, the timing and sources of their introduction into Africa remain enigmatic. Herein, we analyse variation in mitochondrial DNA(mt DNA) D-loop sequences from 345 Nigerian and 37 Kenyan village dogs plus 1530 published sequences of dogs from other parts of Africa, Europe and West Asia. All Kenyan dogs can be assigned to one of three haplogroups(matrilines; clades): A, B, and C, while Nigerian dogs can be assigned to one of four haplogroups A, B, C, and D. None of the African dogs exhibits a matrilineal contribution from the African wolf(Canis lupus lupaster). The genetic signal of a recent demographic expansion is detected in Nigerian dogs from West Africa. The analyses of mitochondrial genomes reveal a maternal genetic link between modern West African and North European dogs indicated by sub-haplogroup D1(but not the entire haplogroup D) coalescing around 12,000 years ago. Incorporating molecular anthropological evidence,we propose that sub-haplogroup D1 in West African dogs could be traced back to the late-glacial dispersals, potentially associated with human hunter-gatherer migration from southwestern Europe.     

Sympatric wolf and coyote populations of the western Great Lakes region are reproductively isolated

Interpretation of the genetic composition and taxonomic history of wolves in the western Great Lakes region (WGLR) of the United States has long been debated and has become more important to their conservation given the recent changes in their status under the Endangered Species Act. Currently, the two competing hypotheses on WGLR wolves are that they resulted from hybridization between (i) grey wolves (Canis lupus) and western coyotes (C. latrans) or (ii) between grey wolves and eastern wolves (C. lycaon). We performed a genetic analysis of sympatric wolves and coyotes from the region to assess the degree of reproductive isolation between them and to clarify the taxonomic status of WGLR wolves. Based on data from maternal, paternal and bi‐parental genetic markers, we demonstrate a clear genetic distinction between sympatric wolves and coyotes and conclude that they are reproductively isolated and that wolf–coyote hybridization in the WGLR is uncommon. The data reject the hypothesis that wolves in the WGLR derive from hybridization between grey wolves and western coyotes, and we conclude that the extant WGLR wolf population is derived from hybridization between grey wolves and eastern wolves. Grey‐eastern wolf hybrids (C. lupus × lycaon) comprise a substantial population that extends across Michigan, Wisconsin, Minnesota and western Ontario. These findings have important implications for the conservation and management of wolves in North America, specifically concerning the overestimation of grey wolf numbers in the United States and the need to address policies for hybrids.     

Deep divergence among mitochondrial lineages in African jackals

Recently, molecular analyses revealed that African and Eurasian golden jackals are distinct species. This finding suggests re‐investigation of the phylogenetic relationships and taxonomy of other African members of the Canidae. Here, we provide a study on the phylogenetic relationship between populations of African jackals Lupulella mesomelas and L. adusta inferred from 962 bp of the mitochondrial cytochrome b (cytb) gene. As expected from its disjunct distribution, with one population in eastern Africa and the other one in southern Africa, we found two mitochondrial lineages within L. mesomelas, which diverged about 2.5 million years ago (Ma). In contrast, in L. adusta with its more continuous distribution in sub‐Saharan Africa, we found only a shallower genetic diversification, with the exception of the West African population, which diverged around 1.4 Ma from the Central and East African populations. Both divergence ages are older than, for example the 1.1–0.9 million years between the grey wolf Canis lupus and the African golden wolf C. lupaster. One taxonomic implication of our findings might be that the two L. mesomelas populations warrant species status. However, genome‐wide data with adequate geographical sampling are needed to substantiate our results.     

Interspecific killing between wolves and golden jackals in Iran

Interspecific killing is a widespread phenomenon among mammalian carnivores; being common, for instance, within the canid guild. The canid guild in Hamadan province, west of Iran, comprises three species: the gray wolf Canis lupus pallipes, the golden jackal Canis aureus, and the red fox Vulpes vulpes. In this area, habitat transformation and agricultural land uses have significantly reduced wild prey populations. In search of food, members of the canid guild are often driven to human settlements. Due to inefficient waste management, the major source of food for canids here is anthropogenic food resources, such as livestock and pet carcasses, and domestic waste illegally dumped near poultry farms and rural areas. Here, we described one of the first reports in the literature of a case of interspecific killing between a wolf and a golden jackal. On June 13, 2016, a collared wolf (adult male) killed an adult jackal (adult male). Further studies are required to understand intraguild competition among red fox, golden jackals, and wolves in this area.     

Native Great Lakes wolves were not restored

Wolves from the Great Lakes area were historically decimated due to habitat loss and predator control programmes. Under the protection of the US Endangered Species Act, the population has rebounded to approximately 3000 individuals. We show that the pre-recovery population was dominated by mitochondrial DNA haplotypes from an endemic American wolf referred to here as the Great Lakes wolf. In contrast, the recent population is admixed, and probably derives also from the grey wolf (Canis lupus) of Old World origin and the coyote (Canis latrans). Consequently, the pre-recovery population has not been restored, casting doubt on delisting actions.     

Canine carnassials: character displacement in the wolves, jackals and foxes of Israel

Previous research implies that competitive character displacement in felids and mustelids of Israel is expressed by canine size. Anatomy and observed killing behaviour of canids suggest that canines in this group are less adapted for the stylized role they play in felids and mustelids. Thus we hypothesized that character displacement, if it exists in canids, should not be manifested more clearly by canine size than by other traits. Five sympatric and at least partially syntopic canids occupy Israel, while in North Africa the largest (wolf) and smallest (Blanford's fox) are absent. Sexual size dimorphism in Israeli canids is generally less than in felids and mustelids (in which we analysed each sex as a separate ‘morphospecies’), so we used mixed-sex samples to represent each species. The three largest species (wolf, golden jackal and red fox) are also represented by Middle Palaeolithic samples in Israel, and all three had larger carnassial lengths then. Carnassial lengths, canine diameters and skull lengths are all remarkable evenly spaced among the five recent species in Israel. In Egypt, no trait manifests significant equality. Despite regional fluctuations in size, the carnassial length ratios of the three smaller species (foxes) are strikingly constant (1.18–1.21) throughout the region, while the ratios for the three larger species (wolf, jackal and red fox), sympatric only in Israel, are larger (1.33–1.34). Finally, mean carnassial length of jackals is constant across North Africa, while skull length and canine diameter both increase from Algeria through Egypt. All three traits are larger in Egypt than in Israel. We tentatively ascribe the equal ratios in Israel to competitive character displacement, though this hypothesis is speculative because of numerous lacunae in knowledge of diet, killing behaviour, available resources and extent of food limitation. Furthermore, humans have greatly affected range, density and ecology of wolves and jackals in the last century. Larger sizes in the Palaeolithic may well be manifestations of Bergmann's rule. The constancy of carnassial length in North African jackals, notwithstanding a longitudinal cline in CBLs of these populations, and the constant ratio between jackal and red fox carnassial length are both consistent with a hypothesis of character release in the absence of the wolf.     

African wolf diet,predation on livestock and conflict in the Guassa mountains of Ethiopia

The African wolf (Canis lupus lupaster) was first identified in 2011 in the Ethiopian highlands, with its status as a new species confirmed in 2015. We studied the diet of a confirmed African wolf population in the Menz‐Guassa Community Conservation Area of central Ethiopia from scat samples collected by den sites from August to November 2010. Rodents were found to be the principal food items occurring in 88.1% of scats (n = 101), followed by plant material (34.7%) and insects (21.8%). Information on reported livestock predation and ensuing conflict with the agro‐pastoral community was obtained through a questionnaire survey. Interview respondents listed the African wolf as the most serious predator of livestock, accounting for 74.6% of the reported kills (n = 492) and 78.9% of the economic loss. Over 70% of reported livestock predation occurred during the dry season (January–April). Better livestock management during this period may significantly reduce conflict. As sympatric Ethiopian wolves primarily feed on rodents, further studies on the foraging ecology, niche overlap and interspecific interactions between the two species should be studied to determine the extent of competition between the two species.     

Ancient origin and evolution of the Indian wolf: evidence from mitochondrial DNA typing of wolves from Trans-Himalayan region and Pennisular India

The two wolf types found in India are represented by two isolated populations and believed to be two sub-species of Canis lupus. One of these wolf, locally called Himalayan wolf (HW) or Tibetan wolf, is found only in the upper Trans-Himalayan region from Himachal Pradesh to Leh in Kasmir and is considered to be C. lupus chanco. The other relatively larger population is of Indian Gray wolf (GW) that is found in the peninsular India and considered to be C. lupus pallipes. Both these wolves are accorded endangered species status under the Indian Wildlife Protection Act. In 1998 for the first time in India, we initiated molecular characterization studies to understand their genetic structure and taxonomic status. Since then, we have analyzed the genetic variability in 18 of the total of 21 HW samples available in Zoological parks along with representative samples of GW, wild dogs and jackals. Our study of mitochondrial DNA diversity across three different taxonomically informative domains i.e., cytochrome-B gene, 16S rDNA and hypervariable d-loop control region revealed HW to be genetically distinct from the GW as well as from all other wolves of the world, including C. lupus chanco from China. Most importantly, d-loop haplotypic diversity revealed both HW and GW from India to be significantly diverse from other wolf populations globally and showed that these represent the most ancient lineages among them. Phylogenetic analysis revealed the Indian wolves as two independent lineages in a clade distinct and basal to the clade of all wolves from outside of India. Conservative estimate of evolutionary time-span suggests more than one million years of separation and independent evolution of HW and GW. We hypothesize that Indian wolves represent a post-jackal pre-wolf ancestral radiation that migrated to India about 1-2 mya and underwent independent evolution without contamination from other wolf like canids. The study thus, suggests that Indian subcontinent had been one major center of origin and diversification of the wolf and related canids. Further, the significant degree of genetic differentiation of HW from GW and of these two from other wolves, suggest the interesting possibility of them to be new wolf species/subspecies in evolution that may need to be described possibly as C. himalayaensis and C. indica (or as C. lupus himalyaensis and C. lupus indica), respectively. Thus for the first time, the study reveals new ancient wolf lineages in India and also highlights the need to revisit the origin, evolution and dispersion of wolf populations in Asia and elsewhere. Simultaneously, it increases the conservation importance of Indian wolves warranting urgent measures for their effective protection and management, especially of the small HW population that at present is not even recognized in the canid Red List.     

Serum concentrations of vitamin D metabolites, vitamins A and E, and carotenoids in six canid and four ursid species at four zoos

Nutritional status for six captive canid species (n=34) and four captive ursid species (n=18) were analyzed. The species analyzed included: African wild dog (Lycaon pictus), arctic fox (Alopex lagopus), gray wolf (Canis lupus), maned wolf (Chrysocyon brachyurus), Mexican wolf (Canis lupus baleiyi), red wolf (Canis rufus), brown bear (Ursus arctos), polar bear (Ursus maritimus), spectacled bear (Tremarctos ornatus), and sun bear (Ursus malayanus). Diet information was collected for these animals from each participating zoo (Brookfield Zoo, Fort Worth Zoo, Lincoln Park Zoological Gardens, and North Carolina Zoological Park). The nutritional composition of the diet for each species at each institution met probable dietary requirements. Blood samples were collected from each animal and analyzed for vitamin D metabolites 25(OH)D and 1,25(OH)(2)D, vitamin A (retinol, retinyl stearate, retinyl palmitate), vitamin E (alpha-tocopherol and gamma-tocopherol) and selected carotenoids. Family differences were found for 25(OH)D, retinol, retinyl stearate, retinyl palmitate and gamma-tocopherol. Species differences were found for all detectable measurements. Carotenoids were not detected in any species. The large number of animals contributing to these data, provides a substantial base for comparing the nutritional status of healthy animals and the differences among them.     

Chromosomal evolution of the Canidae. I. Species with high diploid numbers

The Giemsa banding patterns of seven canid species, including the grey wolf (Canis lupus), the maned wolf (Chrysocyon brachyurus), the bush dog (Speothos venaticus), the crab-eating fox (Cerdocyon thous), the grey fox (Urocyon cinereoargenteus), the bat-eared fox (Otocyon megalotis), and the fennec (Fennecus zerda), are presented and compared. Relative to other members of Canidae, these species have high diploid complements (2n greater than 64) consisting of largely acrocentric chromosomes. They show a considerable degree of chromosome homoeology, but relative to the grey wolf, each species is either missing chromosomes or has unique chromosomal additions and rearrangements. Differences in chromosome morphology among the seven species were used to reconstruct their phylogenetic history. The results suggest that the South American canids are closely related to each other and are derived from a wolf-like progenitor. The fennec and the bat-eared fox seem to be recent derivatives of a lineage that branched early from the wolf-like canids and which also includes the grey fox.     

Mitochondrial DNA coding region sequences support the phylogenetic distinction of two Indian wolf species

Two small endangered populations of Indian wolves were recently shown to be distant from other wolf and dog mtDNA lineages characterized so far. None of the inner branches in the tree of canid species based on partial hypervariable D-loop sequences were, however, statistically supported by the data raising the question whether the two Indian wolf lineages represent two new species, occupying an intermediate position between Canis latrans and C. lupus or have diverged from the sub-species of C. lupus due to isolation and drift. Here we report complete D-loop, cytochrome b, and 16S rRNA sequences data for 23 additional wolves from India analysed in the context of other canid species. Extended analyses of D-loop data and partial sequences of 16S rRNA showed highly reticulated pattern and were unable to resolve unambiguously the phylogenetic relationship of Indian wolves among other canid species. The phylogenetic reconstructions of cytochrome b sequences, however gave significant statistical support for the inner branches supporting genetic distinction of the two Indian wolf lineages within themselves as well as from all other wolves of the world, including individuals belonging to subspecies C. lupus chanco and C. lupus pallipes to which the two Indian wolf populations have been traditionally assigned. Their genetic differentiation relative to worldwide variation of wolves supports the suggestion to treat them as separate wolf species, C. himalayensis and C. indica .     

Food habits of the world's grey wolves

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Substantial functional diversity accompanies limited major histocompatibility complex class II variability in golden jackal (Canis aureus): A comparison between two wild Canis species in Croatia

The golden jackal (Canis aureus) is one of the less studied carnivores and research on its major histocompatibility complex (MHC) variability is just at its early stages. MHC genes encode cell-surface receptors that serve to bind and present antigens to T cells, which is essential to initiating specific immunological responses in vertebrates. In this paper we present for the first time patterns of genetic diversity and natural selection on MHC class II DLA-DRB1, DQA1 and DQB1 loci in the golden jackal using samples from two geographically distinct regions in Croatia and further compare them to the values found in its congener grey wolf (Canis lupus). Diversity of golden jackals at all three loci was markedly lower than that of grey wolves (allelic richness values were 4, 2 and 3 in jackal versus 11.9, 6.6 and 10.2 in wolves for DRB1, DQA1 and DQB1, respectively) and can be attributed to a genetic drift rather than to the lack of historical positive selection. The finding of high evolutionary distances (16.3% for DRB1 and 8.5% for DQB1) and a substantial number of codons predicted to be under the influence of positive selection (11 for DRB1 and 9 for DQB1) suggests that the investigated golden jackal population still contains considerable functional diversity necessary for the presentation of varied foreign peptides. In contrast to neutral genetic variation, our results suggest that the Dalmatian population has a higher MHC diversity than the Slavonian population, casting doubt on its supposed isolation and calling for a more extensive investigation of the MHC variability of southern Balkan jackal populations.     

Genetic analysis of historic western Great Lakes region wolf samples reveals early Canis lupus/lycaon hybridization

The genetic status of wolves in the western Great Lakes region has received increased attention following the decision to remove them from protection under the US Endangered Species Act. A recent study of mitochondrial DNA has suggested that the recovered wolf population is not genetically representative of the historic population. We present microsatellite genotype data on three historic samples and compare them with extant populations, and interpret published genetic data to show that the pre-recovery population was admixed over a century ago by eastern wolf (Canis lycaon) and grey wolf (Canis lupus) hybridization. The DNA profiles of the historic samples are similar to those of extant animals in the region, suggesting that the current Great Lakes wolves are representative of the historic population.     

Analysis of <Emphasis Type="Italic">Canis</Emphasis> mitochondrial DNA demonstrates high concordance between the control region and ATPase genes

Background  

Phylogenetic studies of wild Canis species have relied heavily on the mitochondrial DNA control region (mtDNA CR) to infer species relationships and evolutionary lineages. Previous analyses of the CR provided evidence for a North American evolved eastern wolf (C. lycaon), that is more closely related to red wolves (C. rufus) and coyotes (C. latrans) than grey wolves (C. lupus). Eastern wolf origins, however, continue to be questioned. Therefore, we analyzed mtDNA from 89 wolves and coyotes across North America and Eurasia at 347 base pairs (bp) of the CR and 1067 bp that included the ATPase6 and ATPase8 genes. Phylogenies and divergence estimates were used to clarify the evolutionary history of eastern wolves, and regional comparisons of nonsynonomous to synonomous substitutions (dN/dS) at the ATPase6 and ATPase8 genes were used to elucidate the potential role of selection in shaping mtDNA geographic distribution.     

Domestic dog origin of canine distemper virus in free-ranging wolves in Portugal as revealed by hemagglutinin gene characterization

Serologic evidence for canine distemper virus (CDV) has been described in grey wolves but, to our knowledge, virus strains circulating in wolves have not been characterized genetically. The emergence of CDV in several non-dog hosts has been associated with amino acid substitutions at sites 530 and 549 of the hemagglutinin (H) protein. We sequenced the H gene of wild-type canine distemper virus obtained from two free-ranging Iberian wolves (Canis lupus signatus) and from one domestic dog (Canis familiaris). More differences were found between the two wolf sequences than between one of the wolves (wolf 75) and the dog. The latter two had a very high nucleotide similarity resulting in identical H gene amino acid sequences. Possible explanations include geographic and especially temporal proximity of the CDV obtained from wolf 75 and the domestic dog, taken in 2007-2008, as opposed to that from wolf 3 taken more distantly in 1998. Analysis of the deduced amino acids of the viral hemagglutinin revealed a glycine (G) and a tyrosine (Y) at amino acid positions 530 and 549, respectively, of the partial signaling lymphocytic activation molecule (SLAM)-receptor binding region which is typically found in viral strains obtained from domestic dogs. This suggests that the CDV found in these wolves resulted from transmission events from local domestic dogs rather than from wildlife species.     

Chromosomal evolution of the Canidae. II. Divergence from the primitive carnivore karyotype

The Giemsa-banding patterns of chromosomes from the arctic fox (Alopex lagopus), the red fox (Vulpes vulpes), the kit fox (Vulpes macrotis), and the raccoon dog (Nyctereutes procyonoides) are compared. Despite their traditional placement in different genera, the arctic fox and the kit fox have an identical chromosome morphology and G-banding pattern. The red fox has extensive chromosome arm homoeology with these two species, but has only two entire chromosomes in common. All three species share some chromosomes with the raccoon dog, as does the high diploid-numbered grey wolf (Canis lupus, 2n = 78). Moreover, some chromosomes of the raccoon dog show partial or complete homoeology with metacentric feline chromosomes which suggests that these are primitive canid chromosomes. We present the history of chromosomal rearrangements within the Canidae family based on the assumption that a metacentric-dominated karyotype is primitive for the group.     

Stable isotopes, ecological integration and environmental change: wolves record atmospheric carbon isotope trend better than tree rings

Large-scale patterns of isotope ratios are detectable in the tissues of organisms, but the variability in these patterns often obscures detection of environmental trends. We show that plants and animals at lower trophic levels are relatively poor indicators of the temporal trend in atmospheric carbon isotope ratios (delta13C) when compared with animals at higher trophic levels. First, we tested how differences in atmospheric delta13C values were transferred across three trophic levels. Second, we compared contemporary delta13C trends (1961-2004) in atmospheric CO2 to delta13C patterns in a tree species (jack pine, Pinus banksiana), large herbivore (moose, Alces alces) and large carnivore (grey wolf, Canis lupus) from North America. Third, we compared palaeontological (approx. 30000 to 12000 14C years before present) atmospheric CO2 trends to delta13C patterns in a tree species (Pinus flexilis, Juniperus sp.), a megaherbivore (bison, Bison antiquus) and a large carnivore (dire wolf, Canis dirus) from the La Brea tar pits (southern California, USA) and Great Basin (western USA). Contrary to previous expectations, we found that the environmental isotope pattern is better represented with increasing trophic level. Our results indicate that museum specimens of large carnivores would best reflect large-scale spatial and temporal patterns of carbon isotopes in the palaeontological record because top predators can act as ecological integrators of environmental change.