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
Fagus orientalis
Distribution
taurica
moesiaca
F. orientalis-stands
in SE Europe –
legacy of the
Osman Empire?
orientalis
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
Differentiation sylvatica vs. orientalis
60.00
1
subsp. sylvatica
subsp. sylvatica Calabria
subsp. sylvatica Balkans
0,5
subsp. orientalis Crimea
subsp. orientalis
regression
0
40
longitude [degrees]
60.00
60
<0.20
<0.30
<0.40
<0.50
>0.50
>0.60
>0.70
>0.80
>0.90
Latitude
50.00
40.00
30.00
0.00
10.00
20.00
30.00
40.00
Longitude
50.00
60.00
1
<0.20
<0.30
<0.40
subsp. sylvatica
<0.50
subsp.
sylvatica Calabria
>0.50
subsp.
sylvatica Balkans
>0.60
>0.70
subsp.
orientalis Crimea
highly
divergent
haplotypes
>0.80
subsp.
orientalis
In Crimea
>0.90
regression
50.00
0,5
40.00
-165000
30.00
0.00
0
0
10.00
20
20.00
30.00
40
Longitude
longitude [degrees]
ln(Prob (X |K ))
20
Latitude
0
proportion of the ' orientalis '
gene pool
proportion of the 'orientalis'
gene pool
Bayesian analysis (Pritchard
Morphometry,
et al. 2000)
9 leaf
and fruit traits
Chloroplast
PCR-RFLP
Markers
12 isozyme loci, 279 populations
-170000
40.00
-175000
-180000
-185000
-190000
50.00
60.00
60
Two groups most
probable,
but possible
substructure
-195000
0
2 Syst
4 Evol
6
8
10213–236
12
14
Gömöry et al. (2007)
Demesure
Bot J et
Linn
al. Soc
1996154:
Evolution
50:Plant
2515–2520
Denk
et al.165–174.
(2002)
232:
number of clusters (K )
16
ln(Prob (X |K ))
-165000
-170000
-175000
proportion of
gene pool
-180000
No recognizable
species- -185000
or
-190000
geographic
-195000
structure
0
1,0
F. orientalis
secondary peak –
possible substructure
2
4
6
8
10
12
14
16
Phylogeny
of
the
Fagus
sylvatica
complex
number of clusters (K )
P3o
0,5
P2o
P1o
0,0
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
Thr
WAMi EAMi Aman WCa ECa
Transc
Alborz
Crimea
Denk et al. (2002)
Plant Syst Evol
232: 213–236
Gömöry et al. (2007) Bot J Linn Soc 154: 165–174
12ITS
allozyme loci, 279 populations
nrDNA
ln(Prob (X |K ))
-30000
-31000
syl
syl*
-33000
ori
-34000
ori*
-32000
-35000
F. sylvatica sylvatica
SW Balkan
F. sylvatica orientalis
Krim-Halbinsel
3 groups
most probable
-36000
0
2
4
6
8
number of groups (K )
10
Geographic patterns of the genetic variation
A[100]
A[4000]
ne
FST
F. sylvatica
F. orientalis
2.21
3.40
1.458
0.0397
2.35
4.27
1.455
0.1424
Variation
in
and
both
only
much
in
species,
F.differentiation
sylvatica,
differentiation
but
only
few
in
differentiation
only
F. orientalis
in F. orientalis
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
Fagus moesiaca – is it possible to delineate the “species”?
12 allozyme loci, genetic distances, PCoA
Mdh-C
s
s
s
s
s
s
m
m
Clinal pattern
No clear
boundary
m
m
s
Isolation by
distance
Gömöry et al. (1999) J Evol Biol 12: 746-754
o
Fagus sylvatica s. str. – genetic variation
7 allozyme
cpDNA
cpSSR
PCR-RFLP
loci, 608 populations, SAMOVA
Vendramin in Magri et al. (2006) New Phytologist 171: 199–221
Comps/Gömöry
Geburek
inetMagri
al. (2006)
etAppl
al. New
(2006)
Phytologist
New
171: 199–221
Vettori et in
al.Magri
(2004)
Theor
Genet
109: Phytologist
1–9 171: 199–221
Postglacial colonisation (FOSSILVA)
Projection of SAMOVA groups onto
paleobotanic sites
makrofossils
● C14 dated pollen > 2%
▲
Magri et al. (2006) New Phytologist 171: 199–221
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
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
Just for illustration: international beech provenance
trial, series 1998, plot Tale/Slovakia
height
growth
survival
marginal population
different refugium (?)
different taxon
Thank you for your attention