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

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

Dušan Gömöry & Ladislav Paule

Technical University in Zvolen, Zvolen, Slovakia

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

moesiaca

taurica

orientalis

F. orientalis-standsin SE Europe – legacy of theOsman Empire?

DistributionFagus orientalis

Various morphotypes of beeches in West-Eurasia

Denk et al. (2002) Plant Syst Evol 232: 213–236

A MT 1 EU. North-SpainB MT 3 EU. AustriaC MT 3 EU. AustriaD MT 3 B. BulgariaE MT 1 G. West GeorgiaF MT 1 I. North Iran G MT 1 I. North Iran

30.00

40.00

50.00

60.00

0.00 10.00 20.00 30.00 40.00 50.00 60.00

Longitude

Lat

itu

de

<0.20<0.30<0.40<0.50>0.50>0.60>0.70>0.80>0.90

Gömöry et al. (2007) Bot J Linn Soc 154: 165–174.

Differentiation sylvatica vs. orientalis

Bayesian analysis (Pritchard et al. 2000)12 isozyme loci, 279 populations

0

0,5

1

0 20 40 60

longitude [degrees]

pro

po

rtio

n o

f th

e 'o

rie

nta

lis

' g

en

e p

oo

l

subsp. sylvatica

subsp. sylvatica Calabria

subsp. sylvatica Balkans

subsp. orientalis Crimea

subsp. orientalis

regression

0

0,5

1

0 20 40 60

longitude [degrees]

pro

po

rtio

n o

f th

e '

ori

en

tali

s'

ge

ne

po

ol

subsp. sylvatica

subsp. sylvatica Calabria

subsp. sylvatica Balkans

subsp. orientalis Crimea

subsp. orientalis

regression

30.00

40.00

50.00

60.00

0.00 10.00 20.00 30.00 40.00 50.00 60.00

Longitude

Lat

itu

de

<0.20

<0.30

<0.40

<0.50

>0.50

>0.60

>0.70

>0.80

>0.90

Morphometry, 9 leaf and fruit traits

Denk et al. (2002) Plant Syst Evol 232: 213–236

Chloroplast PCR-RFLP Markers

Demesure et al. 1996 Evolution 50: 2515–2520-195000

-190000

-185000

-180000

-175000

-170000

-165000

0 2 4 6 8 10 12 14 16

number of clusters (K )

ln(P

rob

(X|K

))

highly divergent haplotypesIn Crimea

Two groups most probable,but possiblesubstructure

0,0

0,5

1,0

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59

prop

ortio

n of

gen

e po

ol

P3o

P2o

P1o

Thr WAMi EAMi Aman WCa ECa Transc Alborz Crimea

Gömöry et al. (2007) Bot J Linn Soc 154: 165–17412 allozyme loci, 279 populations

Denk et al. (2002) Plant Syst Evol 232: 213–236

nrDNA ITS

Phylogeny of the Fagus sylvatica complex

syl F. sylvatica sylvaticasyl* SW Balkanori F. sylvatica orientalisori* Krim-Halbinsel

-195000

-190000

-185000

-180000

-175000

-170000

-165000

0 2 4 6 8 10 12 14 16

number of clusters (K )

ln(P

rob

(X|K

))

-36000

-35000

-34000

-33000

-32000

-31000

-30000

0 2 4 6 8 10

number of groups (K )

ln(P

rob

(X|K

))

No recognizablespecies- orgeographicstructure

F. orientalissecondary peak –possible substructure

3 groups most probable

Geographic patterns of the genetic variation

Gömöry et al. (2007) Bot J Linn Soc 154: 165–174 + unpubl. Data12 allozyme loci, 279 populations

F. sylvatica F. orientalis

A[100]

A[4000]

ne

FST

2.21

3.40

1.458

0.0397

2.35

4.27

1.455

0.1424

Hotspots of allelic richness in F. orientalisVariation in both species, differentiation only in F. orientalisVariation in only in F. sylvatica, but few differentiationVariation and much differentiation only in F. orientalis

Fagus moesiaca – is it possible to delineate the “species”?12 allozyme loci, genetic distances, PCoA

Gömöry et al. (1999) J Evol Biol 12: 746-754

Mdh-C

s

s

s

s

s

s

m

mm

mo

s

Clinal patternNo clear boundary

Isolation bydistance

Comps/Gömöry in Magri et al. (2006) New Phytologist 171: 199–221

Fagus sylvatica s. str. – genetic variation

7 allozyme loci, 608 populations, SAMOVAcpSSRcpDNA PCR-RFLP

Vendramin in Magri et al. (2006) New Phytologist 171: 199–221Vettori et al. (2004) Theor Appl Genet 109: 1–9Geburek in Magri et al. (2006) New Phytologist 171: 199–221

Magri et al. (2006) New Phytologist 171: 199–221

Postglacial colonisation (FOSSILVA)

Projection of SAMOVA groups ontopaleobotanic sites

▲ makrofossils● C14 dated pollen > 2%

Comps et al. (2001) Genetics 157: 389–397

Genetic footprints of postglacial migration

Opposite trends of allelic richness and genediversity

Excess of gene diversity against expectations of mutation-driftequilibrium at the periphery – a consequence of recurrent founder events

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

survival

Just for illustration: international beech provenancetrial, series 1998, plot Tale/Slovakia

height growth

different taxon

different refugium (?)

marginal population

Thank you for your attention