Phylogenetic evidence for the ancient Himalayan wolf: towards a clarification of its taxonomic status based on genetic sampling from western Nepal

Abstract

Wolves in the Himalayan region form a monophyletic lineage distinct from the present-day Holarctic grey wolf Canis lupus spp. (Linnaeus 1758) found across Eurasia and North America. Here, we analyse phylogenetic relationships and the geographic distribution of mitochondrial DNA haplotypes of the contemporary Himalayan wolf (proposed in previous studies as Canis himalayensis) found in Central Asia. We combine genetic data from a living Himalayan wolf population collected in northwestern Nepal in this study with already published genetic data, and confirm the Himalayan wolf lineage based on mitochondrial genomic data (508 bp cytochrome b and 242 bp D-loop), and X- and Y-linked zinc-finger protein gene (ZFX and ZFY) sequences. We then compare the genetic profile of the Himalayan wolf lineage found in northwestern Nepal with canid reference sequences from around the globe with maximum likelihood and Bayesian phylogeny building methods to demonstrate that the Himalayan wolf forms a distinct monophyletic clade supported by posterior probabilities/bootstrap for D-loop of greater than 0.92/85 and cytochrome b greater than 0.99/93. The Himalayan wolf shows a unique Y-chromosome (ZFY) haplotype, and shares an X-chromosome haplotype (ZFX) with the newly postulated African wolf. Our results imply that the Himalayan wolf distribution range extends from the Himalayan range north across the Tibetan Plateau up to the Qinghai Lakes region in Qinghai Province in the People's Republic of China. Based on its phylogenetic distinction and its older age of divergence relative to the Holarctic grey wolf, the Himalayan wolf merits formal classification as a distinct taxon of special conservation concern.

Phylogeny of the Caninae (Carnivora): Combining morphology, behaviour, genes and fossils

“…The recently identified “African wolf” Canis anthus (syn. C. lupaster ; see Koepfli et al., ; Viranta, Atickem, Werdelin, & Stenseth, ) was scored under Canis aureus traditionally, as it is here also; the problematic “ C. himalayensis ” and “ C. indicus ” were not included (see Werhahn et al., ). The Falkland Islands wolf ( Dusicyon australis ), exterminated as recently as 1876, was treated as a Recent species.…”

Himalayan wolf distribution and admixture based on multiple genetic markers

“…The Himalayan wolf (also called the Tibetan wolf) has been shown to comprise a genetically distinctive lineage from the Holarctic grey wolves (Aggarwal, Kivisild, Ramadevi, & Singh, 2007;Sharma, Maldonado, Jhala, & Fleischer, 2004;Werhahn et al, 2018;Werhahn, Senn, et al, 2017). In mitochondrial DNA (mtDNA) analysis, Himalayan wolves constitute a monophyletic clade that is sister to the Holarctic grey wolf subspecies (Aggarwal et al, 2007;Koepfli et al, 2015;Sharma et al, 2004;Werhahn et al, 2018), whereas nuclear genome analysis (Fan et al, 2016) indicates that they form a clade within the Holarctic grey wolf complex.…”

“…Himalayan wolves exhibit zinc finger (ZF) protein gene haplotypes on both sex chromosomes that differ from those found in Holarctic grey wolves. Specifically, the ZF protein gene haplotype on the Himalayan X chromosome is shared with the African wolf, recently posited as Canis lupaster (Álvares et al, 2019;Viranta, Atickem, Werdelin, & Stenseth, 2017), while the ZF haplotype on the Himalayan Y chromosome is unique among wolf lineages (Werhahn et al, 2018;Werhahn, Senn, et al, 2017). The Himalayan lineage exhibits mutations on four non-synonymous SNPs in three functional nuclear genes related to the hypoxia pathway (vonHoldt, Fan, Vecchyo, & Wayne, 2017;Werhahn et al, 2018;Zhang et al, 2014), which are likely to affect high-altitude adaptation, that is, the evolution of physiological mechanisms to cope with lifelong hypoxic conditions in high-altitude habitats (Storz, Scott, & Cheviron, 2010).…”

Perspectives of traditional Himalayan communities on fostering coexistence with Himalayan wolf and snow leopard

“…The Himalayan wolf (currently recommended as Canis lupus chanco by Álvares et al, ; also see Werhahn et al, , Werhahn et al, ) and snow leopard are top carnivores coexisting in the Himalayas and the Qinghai‐Tibetan Plateau (QTP) of Asia. As top carnivores, they regulate ecosystem health and processes through trophic interactions with mesopredators, herbivores and the vegetation, facilitating biodiversity and ecosystem resilience (Ripple et al, ), and may contribute to nutrient cycling by supporting scavenger diversity (Wilmers, Crabtree, Smith, Murphy, & Getz, ).…”