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Phylogenetic Evidence for the Ancient Himalayan Wolf: Towards a Clarification of Its Taxonomic Status Based on Genetic Sampling From Western Nepal

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Phylogenetic Evidence for the Ancient Himalayan Wolf: Towards a Clarification of Its Taxonomic Status Based on Genetic Sampling From Western Nepal

Geraldine Werhahn et al. R Soc Open Sci.

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.

Keywords: Canis himalayensis; Canis lupus chanco; Himalaya; Himalayan wolf; Nepal; phylogeny.

Conflict of interest statement

We declare we have no competing interests.

Figures

Figure 1.
Figure 1.
Overview of the current genetic evidence of the Himalayan wolf distribution. The data shown originates from the current and previous studies (Himalayan wolf field collected samples, dark green; Himalayan wolf museum specimens, light green). For overview, samples originating from Holarctic grey wolf lineages found in the region are also shown, i.e. Mongolian grey wolf Canis lupus chanco (dark blue) and Indian grey wolf Canis lupus pallipes (light blue). This study generated 72 Himalayan wolf sequences from 104 field collected samples in Humla, Nepal. The additional data shown derive from other studies [6,7,13].
Figure 2.
Figure 2.
Bayesian phylogeny based on (a) 242 bp D-loop mtDNA sequences and (b) 508 bp cytochrome b mtDNA sequences with Bayesian posterior probability values at nodes. Asterisks at nodes indicate greater than or equal to 80% bootstrap support based on maximum-likelihood analyses and greater than or equal to 0.90 posterior probability from Bayesian inference. The Himalayan wolf (green) is a distinct monophyletic clade relative to the Holarctic grey wolf clade (blue) which also includes domestic dogs. Haplotypes without accession numbers are sequences generated in this study (electronic supplementary material, table S1).
Figure 2.
Figure 2.
Bayesian phylogeny based on (a) 242 bp D-loop mtDNA sequences and (b) 508 bp cytochrome b mtDNA sequences with Bayesian posterior probability values at nodes. Asterisks at nodes indicate greater than or equal to 80% bootstrap support based on maximum-likelihood analyses and greater than or equal to 0.90 posterior probability from Bayesian inference. The Himalayan wolf (green) is a distinct monophyletic clade relative to the Holarctic grey wolf clade (blue) which also includes domestic dogs. Haplotypes without accession numbers are sequences generated in this study (electronic supplementary material, table S1).
Figure 3.
Figure 3.
Median-joining networks based on (a) 242 bp D-loop haplotypes of Canis species and (b) 508 bp cytochrome b haplotypes of Canis species. Himalayan wolves (HW, green), African wolves (AW, red) form monophyletic clades. The grey wolf complex clusters with grey wolves from around the globe (GW, blue) including Indian grey wolves (IW, light blue) and domestic dogs (DD, yellow).
Figure 4.
Figure 4.
Haplotype network showing the ZFY and ZFX final intron sequences of Himalayan wolf (green), African wolf (red), Holarctic grey wolf (blue) and golden jackal (red). The black dots on the internode represent indels and substitutions between the haplotypes.
Figure 5.
Figure 5.
Himalayan wolf adults in Humla, Nepal. Photograph (a) shows a pale-coloured wolf individual, and (b) a black-coloured wolf individual (© Geraldine Werhahn).

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References

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