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