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. 2012;7(8):e42740.
doi: 10.1371/journal.pone.0042740. Epub 2012 Aug 10.

Reviving the African wolf Canis lupus lupaster in North and West Africa: a mitochondrial lineage ranging more than 6,000 km wide

Philippe Gaubert  1 , Cécile Bloch, Slim Benyacoub, Adnan Abdelhamid, Paolo Pagani, Chabi Adéyèmi Marc Sylvestre Djagoun, Arnaud Couloux, Sylvain Dufour
Affiliations

Reviving the African wolf Canis lupus lupaster in North and West Africa: a mitochondrial lineage ranging more than 6,000 km wide

Philippe Gaubert et al. PLoS One. 2012.

Abstract

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.

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Conflict of interest statement

Competing Interests: The authors have declared that no competing interests exist.

Figures

Figure 1
Figure 1
A- Geographic distribution of the African Canis included in this study. Numbers indicate samples collected for this study (see Table 1), and letters refer to the nucleotide sequences available in the literature or Genbank for cytochrome b. Square, circles and stars represent mtDNA-typed Canis lupus/familiaris, C. l. lupaster, and Canis spp. other than C. lupus, respectively: 1- near Sharm el Shikh city, south Sinai, Egypt; 2- coastal region between Skikda and El-Kala, Algeria; 3- Terarabat, Adrar des Iforas, Mali; 4- near Kheune, east of P.N. du Djoudj, Senegal; 5- Koubia, Dalaba and Tougue districts, Guinea; 6- Béterou, Benin; a- northern Egypt ; b- Menz region, Ethiopia ; c- southern Kenya and northern Tanzania . B- The typical ‘wolf-like’ (left) and ‘jackal-like’ (right) phenotypes observed near Kheune, Senegal. (photographs: C. Bloch).
Figure 2
Figure 2. Bayesian phylogenetic analysis of the wolf-like clade based on cytochrome b and control region.
The model HKY + I + Γ was applied to the coherent fragment “cytochrome b + control region”, assuming a constant size coalescent model. Values at nodes correspond to posterior probabilities ≥0.90. Clades were collapsed for better readability of the tree. Scale bar represents 2% sequence divergence.
Figure 3
Figure 3. Median-joining networks of Canis lupus and jackal haplotypes based on cytochrome b.
Circle size and branches are proportional to haplotype frequency and number of mutation steps among haplotypes, respectively. Numbers refer to mutation steps separating the main haplogroups (circled). To improve overall visualization of the network, branch proportions were not respected within the Canis lupus/familiaris haplogroup.
Figure 4
Figure 4. Habitats of the African wolf.
A- Agricultural landscape with wildland (Asphodelus microcarpus), coastal region between Skikda and El-Kala, Algeria (photograph: S. Benyacoub); B- Sandy Sahelian savannah with scarce shrub cover (Tamarix senegalensis), near Kheune, Senegal (photograph: C. Bloch).
Figure 5
Figure 5. Phenotypic and behavioral traits between Canis species near Kheune, Senegal.
A- Aggressive posture of the African wolf towards golden jackals; B- Aggressive posture of a golden jackal towards congeners; C- Typical ‘wolf-like’ phenotype, sampled in this study (T1361); D- Typical ‘jackal-like’ phenotypes, sampled in this study (T1360); E- Food guarding of golden jackals on a dead carcass of cow; E- African wolf (left) fighting with golden jackal (right) to access the dead carcass; F- Simulated mating between two male golden jackals; G- Phenotype of the feral dogs living in sympatry with the African wolf and the golden jackal. (photographs: C. Bloch).
Figure 6
Figure 6. Phenotypic variation in the golden jackal and the African wolf near Kheune, Senegal.
A- Typical ‘jackal-like’ phenotype; B- ‘Jackal-like’ phenotype tending towards C; C- ‘Intermediate’ phenotype between golden jackal and African wolf; D- ‘wolf-like’ phenotype tending towards C; E- Typical ‘wolf-like’ phenotype. (photographs: C. Bloch).
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