Taxonomic and neutral genetic variation within the complex of West-Eurasian beeches

1. Taxonomic and neutral genetic variation within the complex of West-Eurasian beeches Dušan Gömöry & Ladislav Paule Technical University in Zvolen, Zvolen, Slovakia

2. Complex Fagus sylvatica L. (Flora Europaea, Greuter & Burdet 1981): Generally recognized taxa Fagus sylvatica ssp. sylvatica Europa Fagus sylvatica ssp. orientalis (Fagus orientalis Lipsky) Thrakia, Ponthic Mts., Amanus Mts., Grand Caucasus, Small Caucasus, Transcaucasia, Alborz Mts. (F. hohenackeriana Palibin?) Questionable taxa ? Fagus moesiaca (Maly) Czeczott Serbia, Macedonia, Bosnia?, Bulgaria? Greece? Albania? ? Fagus taurica Popl. Crimea

3. Fagus orientalis Distribution taurica F. orientalis-stands in SE Europe – legacy of the Osman Empire? moesiaca orientalis

4. A MT 1 EU. North-Spain B MT 3 EU. Austria C MT 3 EU. Austria D MT 3 B. Bulgaria E MT 1 G. West Georgia F MT 1 I. North Iran G MT 1 I. North Iran Various morphotypes of beeches in West-Eurasia Denk et al. (2002) Plant Syst Evol 232: 213–236

5. Differentiation sylvatica vs. orientalis Bayesian analysis (Pritchard et al. 2000) 12 isozyme loci, 279 populations Morphometry, 9 leaf and fruit traits Chloroplast PCR-RFLP Markers highly divergent haplotypes In Crimea Two groups most probable, but possible substructure Gömöry et al. (2007) Bot J Linn Soc 154: 165–174. Demesure et al. 1996 Evolution 50: 2515–2520 Denket al. (2002)Plant Syst Evol 232: 213–236

6. Thr WAMi EAMi Aman WCa ECa Transc Alborz Crimea F. orientalis No recognizable species- or geographic structure secondary peak – possible substructure Phylogeny of the Fagus sylvatica complex sylF. sylvatica sylvatica syl* SW Balkan ori F. sylvatica orientalis ori* Krim-Halbinsel Denket al. (2002) Plant Syst Evol 232: 213–236 nrDNA ITS 3 groups most probable Gömöry et al. (2007) Bot J Linn Soc 154: 165–174 12 allozyme loci, 279 populations

7. Geographic patterns of the genetic variation Variation in both species, differentiation only in F. orientalis Variation and much differentiation only in F. orientalis Variation in only in F. sylvatica, but few differentiation Hotspots of allelic richness in F. orientalis Gömöry et al. (2007) Bot J Linn Soc 154: 165–174 + unpubl. Data 12 allozyme loci, 279 populations

8. Mdh-C s s s s s s m m m m o s Fagus moesiaca – is it possible to delineate the “species”? 12 allozyme loci, genetic distances, PCoA Clinal pattern No clear boundary Isolation by distance Gömöry et al. (1999)J Evol Biol 12: 746-754

9. Fagus sylvatica s. str. – genetic variation 7 allozyme loci, 608 populations, SAMOVA cpSSR cpDNA PCR-RFLP Vendramin in Magri et al. (2006)New Phytologist 171: 199–221 Vettori et al. (2004) Theor Appl Genet 109: 1–9 Comps/Gömöry in Magri et al. (2006)New Phytologist 171: 199–221 Geburek in Magri et al. (2006)New Phytologist 171: 199–221

10. Postglacial colonisation (FOSSILVA) Projection of SAMOVA groups onto paleobotanicsites ▲makrofossils ●C14 dated pollen > 2% Magri et al. (2006)New Phytologist 171: 199–221

11. Genetic footprints of postglacial migration Opposite trends of allelic richness and gene diversity Excess of gene diversity against expectations of mutation-drift equilibrium at the periphery – a consequence of recurrent founder events Comps et al. (2001)Genetics 157: 389–397

12. Caveats for provenance research – • differences in adaptive traits need • not necessarily be due to adaptation • taxonomic differences within the innerhalb des • distribution range (Balkan provenances) • origin from different glacial refugia • (Central Europe vs. Italy or S France) • lower initial allelic richness in the northern • (western?) marginal populations

13. survival height growth Just for illustration: international beech provenance trial, series 1998, plot Tale/Slovakia marginal population different refugium (?) different taxon

14. Thank you for your attention