European Platform on Biodiversity Research Strategy Meeting "World Biodiversity and European . Strategies in Taxonomy: Research in a Changing World" Pr ůhonice, Czech Republic, May 2009

Genetic diversity of in Europe and conservation

Jan Zima

Institute of Vertebrate Biology , Academy of Sciences of the Czech Republic Kvetna 8, CZ-60365 Brno, Czech Republic [email protected] Talk Outline

 Fauna of European mammals  Diversity and taxonomy of the European mammalian biota  Sources and origins of the diversity  Phylogeographic studies  Northern and cryptic refugia  Revealing traces of introduction attempts  Hybridization and hybrid zones  Conservation implications Mammalian species richness in Europe (native species only)

taxon number of species Rodentia 91 Lagomorpha 8 Eulipotyphla 36 Chiroptera 42 Artiodactyla 16 Cetacea 33 Carnivora 26 (Fissipedia)

European Assessment Carnivora 8 2007 (Pinnipedia) TOTAL 260 Mammalian species richness in Europe (introduced species included, cetaceans excluded)

taxon number of species Diprotodontia 1 Lagomorpha 10 Rodentia 106 Primates 1 Eulipotyphla 39 Chiroptera 42 Artiodactyla 26 Carnivora 25 TOTAL 250

Aulagnier et al.: Guide des mammifères d’Europe, 2008 Mammalian distribution mapping in Europe Species diversity of European mammals

Mitchell-Jones et al. 1999, European Mammal Assessment Academic Press 2007 Mammals Species richness of the mammals and vertebrates in the Czech Republic (no. of species)

All vertebrates

►

Krojerová-Prokešová et al. 2008: Folia Zool. Biodiversity hotspots and endemism in Europe

Williams 1999 European Mammal Assessment 2007 Mammalian species richness in Europe

• 219 autochthonous species • 31 introduced species • 33 strictly marine species (cetaceans)

• 45 species differentiated recently Karyotypic races in the lesser horseshoe bat (Rhinolophus hipposideros )

54

Zima et al. 1992; Puerma et al. 2008, Genetica Karyotypic races in the European pine , subterraneus

● 52-chromosome race

X Y ○ 54-chromosome race

X Y Macholán et al. 2001: J. Zool. Karyotypic races in the garden dormouse (Eliomys quercinus )

50- chromosome race from central Europe

Zima et al. 1997, Folia Zool. Type localities of the karyotypic races of the common shrew (Sorex araneus )

The karyotype of the Drnholec race

Zima et al. 1996, Hereditas; Wójcik et al. 2003, Mammalia Distribution of cryptic species from the group

Microtus arvalis

M. levis

M. arvalis M. obscurus Traditional view of glacial refugia situated in the Mediterranean area

Bilton et al. 1998: Proc. R. Soc B Alternative views of glacial refugia and postglacial colonization Mediterranean Europe as an area of endemism for small mammals rather than source for nortwards postglacial colonization

Bilton et al. 1998: Proc. R. Soc B Originally identified phylogeographic clades in bank

Bilton et al . 1998, Proc . R. Soc. B; Deffontaine et al. 2005 , Mol. Ecol . New data: Maximum likelihood tree of Central and South-eastern European populations of bank voles

W Balkan / Central European

Ca rpa thi an Carpathian s

ps Alp B a al ka an Eastern s

E Balkan

Kotlík et al. 2006, PNAS Estimates of the divergence time Isolation with migration model, Bayesian coalescent method (MCMC)

0,015 2.1 (1.5 -2.9) ky

0,010 Probability Probability

0,005 Marginal Posterior Marginal Posterior 0,000 0 1 2 3 4 5 6 7 8 9 t = t * u

Kotlík et al. 2006, PNAS Estimates of the divergence time Range of molecular clocks: 3.6% to 13% sequence divergence / MY

90% HPD ML estimate

Riss glacial Würm glacial LGM LGM

Riss / W ürm Holocene interglacial The height of the last glaciation in Europe

Kotlík et al. 2006, PNAS , Microtus tatricus: the only mammal endemic to the Carpathian Mountains

Martínková et al. 2003, 2004, Folia Zool. Distribution of chromosome diploid number among European pine voles (subgen. ) Phylogenetic relationships among voles of the genus Microtus, the position of the Terricola clade, and phylogenetic relationships of M. tatricus and M. bavaricus

Jaarola et al. 2004, Mol. Phylogenet. Evol., Martínková et al. 2007, Folia Zool. Distribution of the pine vole species endemic to the Alps

M. bavaricus

↓ ↓

M. multiplex -liechtensteini M. liechtensteini

↑

↑↑↑ ↑↑

Spitzenberger 2001 Phylogenetic position of the Bavarian pine vole and its karyotype (2n=46)

Martínková et al. 2007 Folia Zool. Genetic structure and evidence for recent population decline in Eurasian otter

Hájková et al. 2006, J. Zool. The hybrid zone between Mus musculus and M. domesticus in Europe

-musculus

Macholán et al. 2007 Evolution Clines in the hybrid zone and frequency discontinuities for various genetic markers Phylogeographic studies of mammals in tropical regions Phylogenetic relationships within populations of a rodent genus Praomys in western Africa

D

C2

C1

B

A

Nicolas et al. 2008, Mol. Ecol. Geographic distribution of the clades

Clade D

Clade A Clade B

Clade C1 Clade C2

Nicolas et al. 2008, Mol. Ecol. Outgroup Possible phylogeographic scenario BeninClade A C319 Clade B východníClade klád C1/C2

západní klád B Clade D (separated to two not supported západnísubclades klád by A Senegal river)

Nicolas et al. 2008, Mol. Ecol. Median Joining Network of haplotypes found in Praomys rostratus in western Africa

h1 à h25

h26

13

h27 à h44

4

7 H59 à h172 h45 10

h46 4 h47 5 4 h48 à h58

h59 à h80

The lenght of branches is proportional to the number of mutations The size of circeles is proportional to the number of individuals with respective haplotype

Nicolas et al. 2008, Mol. Ecol. yérendé Khoneya

Distribution of haplotype lineages of Preaomys rostratus in western Africa

Nicolas et al. 2008, Mol. Ecol. Possible taxonomic changes resulting from combined molecular analyses of mtDNA and microsatellites: possible discovery of a new species and invalidation of P. derooi weak congruence between both the sets of data

3 „P. sp.n.“ mtDNA P. derooi 2

1 mtDNA P. sp. n.

0

-1 Factorial Factorial axis 2 -2 „P. derooi “

-3

-4 -1.5 -1 -0.5 0 0.5 1 1.5 2 Factorial axis 1

mtDNA = P. sp. n., microsatellites (+ phenotype, habitat etc.) = P. derooi

Nicolas et al. 2008, Mol. Ecol. Conclusions and Recommendations

• Even in the well-studied groups as mammals, the taxonomy is still in progress and many changes are steadily introduced. • The current concepts of the species are not always sufficient to recognize reliably the species status. Well-supported phylogenetic lineages should be handled as valid species. Forget the subspecies? • Support should be given to conservation genetic research. • Conservation of sites of high scientific interest such as cryptic refugia or hybrid zones should be considered seriously. • Experience gained in European research should be applied in other regions and training capacities should be provided to local students. • Conservation legislation should not generate administrative obstacles preventing to perform the research work effectively. Acknowledgements

Josef Bryja Jeremy B. Searle Petra Hájková Natalia Martínková Petr Kotlík Maarit Jaarola Jaroslav Piálek Friederike Spitzenberger Miloš Macholán Adam Kone čný David Bilton Boris Kryštufek Anthony Mitchell-Jones … and many others