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Genetic Differentiation of an Endangered Capercaillie (Tetrao Urogallus) Population at the Southern Edge of the Species Range

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Genetic Differentiation of an Endangered Capercaillie (Tetrao Urogallus) Population at the Southern Edge of the Species Range Conserv Genet (2007) 8:659–670 DOI 10.1007/s10592-006-9212-z ORIGINAL PAPER Genetic differentiation of an endangered capercaillie (Tetrao urogallus) population at the Southern edge of the species range R. Rodrı´guez-Mun˜ oz Æ P. M. Mirol Æ G. Segelbacher Æ A. Ferna´ndez Æ T. Tregenza Received: 27 February 2006 / Accepted: 31 August 2006 / Published online: 9 November 2006 Ó Springer Science+Business Media B.V. 2006 Abstract The low-latitude limits of species ranges limits of a number of boreal species. The capercaillie are thought to be particularly important as long-term is one such species with a range extending northwards stores of genetic diversity and hot spots for specia- to cover most of Europe from Iberia to Scandinavia tion. The Iberian Peninsula, one of the main glacial and East to Siberia. The Cantabrian Range, in North refugia in Europe, houses the southern distribution Spain, constitutes the contemporary south-western distribution limit of the species. In contrast to all other populations, which live in pure or mixed coniferous forests, the Cantabrian population is un- R. Rodrı´guez-Mun˜ oz Á T. Tregenza (&) ique in inhabiting pure deciduous forests. We have Centre for Ecology and Conservation, School of assessed the existence of genetic differentiation be- Biosciences, University of Exeter, Cornwall Campus, tween this and other European populations using Penryn TR10 9EZ, UK e-mail: [email protected] microsatellite and mitochondrial DNA (mtDNA) extracted from capercaillie feathers. Samples were R. Rodrı´guez-Mun˜ oz e-mail: [email protected] collected between 2001 and 2004 across most of the current distribution of the Cantabrian population. P. M. Mirol Mitochondrial DNA analysis showed that the Can- CIGEBA, Facultad de Ciencias Veterinarias, Universidad tabrian birds form a distinct clade with respect to all Nacional de La Plata, B1900AVW La Plata, Argentina e-mail: [email protected] the other European populations analysed, including the Alps, Black Forest, Scandinavia and Russia, G. Segelbacher which are all members of a discrete clade. Microsat- Max Planck Institute of Ornithology, Vogelwarte ellite DNA from Cantabrian birds reveals the lowest Radolfzell, Schlossallee 2, D-78315 Radolfzell, Germany e-mail: [email protected] genetic variation within the species in Europe. The existence of birds from both mtDNA clades in the G. Segelbacher Pyrenees and evidence from microsatellite frequen- Department of Wildlife Ecology and Management, cies for two different groups, points to the existence University Freiburg, Tennenbacher Str. 4, D-79106 Freiburg, Germany of a Pyrenean contact zone between European and Cantabrian type birds. The ecological and genetic A. Ferna´ndez differences of the Cantabrian capercaillies qualify Estacio´ n Biolo´ gica de Don˜ ana, Consejo Superior de them as an Evolutionarily Significant Unit and sup- Investigaciones Cientı´ficas. , Pabello´ n del Peru´ , Avda. Maria Luisa s/n, 41 013 Sevilla, Spain port the idea of the importance of the rear edge for e-mail: [email protected] speciation. Implications for capercaillie taxonomy and conservation are discussed. Present Address: P. M. Mirol Á Á Á Á Museo Argentino de Ciencias Naturales, Angel Gallardo Keywords ESU Glaciation Refugia Taxonomy 470, C1405DJR Buenos Aires, Argentina Hybrid zone Á Contact zone Á Grouse 123 660 Conserv Genet (2007) 8:659–670 Introduction species previously described in Finland from morphol- ogy, display song and allozyme loci (Liukkonen-Anttila Besides its intrinsic interest in relation to the temporal et al. 2004). Although no accurate data are available on processes driving speciation and species distributions historical distribution, recent isolation of the Canta- (Barraclough and Nee 2001), the identification of brian population from its nearest neighbour in the Py- species subunits from genetic analyses has recently renees is thought to date from at least the 18th century become a matter of interest from a conservation per- (Castroviejo et al. 1974). In contrast with the coniferous spective. Historically, nature conservation has focused forests inhabited by all other populations, Cantabrian on the protection of ecosystems and species as a whole. forests are deciduous, with a dominance of oak New insights support the importance of preserving (Quercus petraea and Quercus pyrenaica) and beech genetic variation within species, defining Evolution- (Fagus sylvatica). Evidence from palynological studies, arily Significant Units (ESUs) as distinct population indicates that conifers were replaced by deciduous segments (Moritz 2002). Differences among popula- forests during the Holocene, with only a small relict tions of the same species can arise either from pro- Scots pine (Pinus sylvestris) forest having survived cesses of divergent selection or from neutral processes (Garcı´a Anto´ netal. 1997). The absence of conifers is a particularly where there are colonisation events or major habitat difference between capercaillies living in long-term historical isolation (Moritz 2002; Tregenza the Cantabrian range and those living elsewhere 2002). The importance of quaternary glaciations in (Quevedo et al. 2006). Pine needles are one of the main moulding the current phylogeographical patterns is food sources for capercaillies elsewhere in the world, well established (Avise and Walker 1998; Hewitt 2000). whereas those inhabiting the Cantabrian Range have a Temperature cycles linked to glacial periods have distinct diet, with beech buds and holly tree leaves produced alternate expansions and contractions to- substituting the pine needles as the main source of food gether with North and Southward shifts of species during the winter (Castroviejo 1975; Rodrı´guez and ranges (Hewitt 2000). During peaks of ice extension, Obeso 2000). Based on this ecological divergence and most species have been relegated to a few milder cli- some significant differences in morphology between the mate areas located at low latitude (Taberlet et al. Cantabrian and the Pyrenean capercaillies, the former 1998). The importance of the low latitude range edges population was described as belonging to a different as long-term stores of genetic diversity and hot spots subspecies (Castroviejo 1967), the Cantabrian caper- for speciation has been recently stressed (Martin and caillie (T. u. cantabricus). Because of its geographic McKay 2004; Hampe and Petit 2005). location, at the edge of the species distribution and in The Iberian Peninsula was one of the main glacial one of the main glacial refugia across Europe (Hewitt refugia in Europe (Taberlet et al. 1998), with a number 2000), the study of this population is particularly of well-studied contemporary contact zones between interesting. The Cantabrian capercaillie presumably historically Iberian populations and the descendants of survived through the Ice Ages somewhere in Iberia, and more Easterly refugia (Butlin et al. 1992; Guillaume its contemporary distribution together with its rela- et al. 2000). Close to the Pyrenees, to the West, the tionships with other capercaillie populations are pres- Cantabrian Range runs parallel to the North Iberian ent day evidence of its evolutionary history. coast, acting as a boundary between two very distinct It has been persuasively argued that the capercaillie biogeographical regions, the Atlantic to the North and is a good indicator or ‘umbrella’ species, since it is the Mediterranean to the South (Go´ mez and Lunt confined to undisturbed environments with high spe- 2006). This particular position has allowed this range to cies diversity (Suter et al. 2002; Pakkala et al. 2003). maintain stable populations of many Boreal and Additionally, within the Cantabrian Range the caper- Central European species during warm periods as well caillie is, together with the brown bear, the major as acting as a refugia during the glaciations (Go´ mez flagship species in relation to the conservation of pro- and Lunt 2006). tected areas. Thus, in practical terms, preservation of The capercaillie (Tetrao urogallus) is a bird species this population is closely linked to the preservation of inhabiting coniferous and mixed forests from Western the whole forest ecosystem. Over the last two decades Europe to Eastern Siberia (Storch 2000). Phenotypic both the population size and the distribution area of characters have led to the description of twelve differ- the Cantabrian capercaillie have decreased dramati- ent subspecies (de Juana 1994), but there is still a cally (Obeso 2003). Understanding the phylogeogra- shortage of genetic analyses supporting current classi- phy and the genetic status of this population are two fication. Moreover, a recent study based on mtDNA key issues for the rationalization of any recovery action found no clear evidence for the presence of the sub- to be taken. 123 Conserv Genet (2007) 8:659–670 661 By combining microsatellite and mtDNA analysis, for this species (Segelbacher and Storch 2002). Feath- we aim to assess (1) the degree of genetic differentia- ers were processed by cutting the tips into small pieces tion of the Cantabrian capercaillie population, (2) its to be used together with the remains of skin attached present genetic diversity, and (3) the position of the around them. Pyrenees in the current phylogeographic scenario of the capercaillie in Europe. Mitochondrial sequencing The amplification of a fragment of the control region Materials and methods (CR) of the mtDNA was carried out with primers PHDL (5¢-AGG ACT ACG GCT TGA AAA
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