Phylogeography - site de Sylvain Ursenbacher
Phylogeography - site de Sylvain Ursenbacher
Phylogeography - site de Sylvain Ursenbacher
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<strong>Phylogeography</strong><br />
of European reptiles<br />
<strong>Sylvain</strong> <strong>Ursenbacher</strong><br />
Department of Environmental Sciences<br />
Section of Conservation Biology<br />
University of Basel
<strong>Phylogeography</strong> and DNA<br />
• <strong>Phylogeography</strong>: historical processes that may be responsible for<br />
the contemporary geographic distributions of individuals<br />
• previously based on morphology (but coevolution)<br />
• with PCR: sequencing and genetic relationships<br />
• using mtDNA<br />
only female genetic structure<br />
• also introns or genetic diversity of nuclear genes<br />
• increase of phylogeography since about 1990
<strong>Phylogeography</strong> in Europe: introduction<br />
• research of general pattern of genetic structure between species<br />
(animals and plants)<br />
• P Taberlet / GM Hewitt<br />
Taberlet et al (1998)
<strong>Phylogeography</strong> in Europe: introduction<br />
Chorthippus parallelus Erinaceus spp Ursus arctos<br />
• Hewitt (1999, 2000)<br />
Alnus glutinosa Quercus spp Sorex araneus<br />
Fagus sylvatica Abies alba Arvicola terrestris<br />
Triturus cristatus Mus musculus Crocidura suaveolens
<strong>Phylogeography</strong> in Europe: introduction<br />
• southern refugia for temperate species
<strong>Phylogeography</strong> in Europe: introduction<br />
• Differentiation<br />
Hewitt, 1996
<strong>Phylogeography</strong> in Europe: introduction<br />
• timing: based on molecular clock /<br />
complex bayesian methods<br />
estimated e.g. with fossil on islands,<br />
historical split between species, ...<br />
• for most mammals: only Pleistocene<br />
(max. 1.5 mya)
eptiles characteristics<br />
• possibility to hibernate (not <strong>de</strong>pending on winter conditions)<br />
• directly <strong>de</strong>pendant on the temperature (ectothermic animals)<br />
• low migration rate (most species)<br />
• slow morphological differentiation<br />
• ...<br />
• impact of previous Pleistocene temperature fluctuations
<strong>Phylogeography</strong> of several European species<br />
• “warm” species<br />
• Vipera ammodytes (Balkan peninsula)<br />
• Podarcis species (Balkan peninsula, Greece and Aegean Islands)<br />
• “medium temperate” species<br />
• Vipera aspis (Italy, Spain, France, Switzerland)<br />
• Emys orbicularis (Spain to Caspian sea)<br />
• “cold-tolerant” species<br />
• Vipera berus (France to Sakhalin Island)<br />
• Zootoca vivipara (Spain to Sakhalin Island)
“warm” species: Vipera ammodytes<br />
• (GB) nose-horned or sand viper<br />
• (D) Sandotter, hornotter
“warm” species: Vipera ammodytes<br />
-<br />
-<br />
-<br />
-<br />
-<br />
-<br />
81<br />
66<br />
80<br />
88<br />
66<br />
54<br />
86<br />
77<br />
85<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
94<br />
91<br />
92<br />
54 -<br />
52<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
SW1<br />
SW2<br />
SW3<br />
SW4<br />
NE6<br />
NE2<br />
NE1<br />
NE4<br />
NE5<br />
NE3<br />
MO5<br />
MO2<br />
MO3<br />
MO1<br />
MO4<br />
<strong>Ursenbacher</strong> et al. 2008<br />
99<br />
98<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
NW14, NW16<br />
NW13<br />
NW17<br />
NW15<br />
NW18<br />
NW1<br />
NW2<br />
NW3, NW4, NW6<br />
NW5<br />
NW7, NW8<br />
NW11<br />
NW9<br />
NW12<br />
NW10<br />
AM3<br />
AM2<br />
AM1<br />
T1<br />
PE1<br />
PE3<br />
PE2<br />
CY2<br />
CY1<br />
CY3<br />
CY4<br />
V. aspis<br />
S6<br />
S7<br />
S1<br />
S2<br />
S3<br />
S8<br />
S5<br />
S9<br />
S4<br />
S10<br />
E5<br />
E1<br />
E3<br />
E4<br />
E2<br />
South-eastern cla<strong>de</strong><br />
Asia Minor subcla<strong>de</strong><br />
Southern subcla<strong>de</strong><br />
Eastern subcla<strong>de</strong><br />
Turkish subcla<strong>de</strong><br />
Peloponnese<br />
cla<strong>de</strong><br />
Cycla<strong>de</strong>s<br />
cla<strong>de</strong><br />
South-western<br />
cla<strong>de</strong><br />
North-western<br />
cla<strong>de</strong><br />
North-eastern<br />
cla<strong>de</strong><br />
Montenegrin<br />
cla<strong>de</strong>
“warm” species: Vipera ammodytes<br />
timing estimated with BEAST (Drummond and Rambaut, 2002)
“warm” species: Vipera ammodytes
“warm” species: Vipera ammodytes<br />
Figure 3. Parsimony network of cytochrome b haplotypes. Numbered circles: recor<strong>de</strong>d haplotypes. Small circles stand<br />
for missing haplotypes. A: The haplotype assumed as ancient for postglacial spread to the north
“warm” species: Podarcis species in Balkan peninsula<br />
• (GB) Balkan wall lizard group<br />
• (D) Mauerei<strong>de</strong>chse Groupe
“warm” species: Podarcis species in Balkan peninsula<br />
129 samples; cytochrome b and 16S, 927 bp.<br />
Poulakakis et al. 2005
“warm” species: Podarcis species in Balkan peninsula
“medium temperate” species: Vipera aspis<br />
• (GB) Asp viper<br />
• (D) Aspisviper / Juraviper
“medium temperate” species: Vipera aspis<br />
53 samples; Control region, 671 bp.<br />
H1<br />
H3<br />
?<br />
zinnikeri<br />
aspis<br />
H1<br />
H6<br />
H11<br />
H9<br />
H1<br />
H1<br />
H6<br />
H8<br />
H1<br />
H6<br />
H13<br />
H7<br />
?<br />
H12<br />
H10<br />
H1<br />
1A<br />
1B<br />
H1<br />
H1<br />
?<br />
atra<br />
H1<br />
H2<br />
H1<br />
H1<br />
H1<br />
H1<br />
H16<br />
H15<br />
H1<br />
H1<br />
H16<br />
H4<br />
H17<br />
H5<br />
H1<br />
H1 H1<br />
H1 H1<br />
H16<br />
H1<br />
H16<br />
H1<br />
H18<br />
H16<br />
francisciredi<br />
H16<br />
H14<br />
2B<br />
H15<br />
hugyi<br />
2A<br />
H20<br />
H19<br />
H20<br />
<strong>Ursenbacher</strong> et al. 2006
“medium temperate” species: Vipera aspis<br />
• first: 2 refuges<br />
! Spain<br />
! Italy<br />
• Alps are also a contact zone for several species<br />
R1<br />
i.e.: Sorex araneus, Arvicola terrestris, Triturus sp., Salmo trutta,<br />
Apis mellifera, Chorthippus parallelus, ...<br />
R2
“medium temperate” species: Vipera aspis<br />
• first: 2 refuges<br />
! Spain<br />
! Italy<br />
• first: 4 refuges<br />
! Spain<br />
! France<br />
! Italy<br />
! Italy<br />
R1<br />
R1’<br />
R2’<br />
R2
“medium temperate” species: Emys orbicularis<br />
• (GB) European pond turtle / European pond tortoise<br />
• (D) Europäische Sumpfschildkröte
“medium temperate” species: Emys orbicularis<br />
>1100 samples; cytochrome b, 1031 bp.<br />
Fritz et al. 2007
“cold-tolerant” species: Zootoca (Lacerta) vivipara<br />
• (GB) European common lizard<br />
• (D) Wal<strong>de</strong>i<strong>de</strong>chse / Bergei<strong>de</strong>chse / Moorei<strong>de</strong>chse
30<br />
31 32 33 34<br />
“cold-tolerant” species: Zootoca (Lacerta) vivipara<br />
522 samples; cytochrome b and 16S, 1660 bp<br />
101<br />
45°<br />
102<br />
130<br />
124<br />
64<br />
61<br />
63<br />
67<br />
50°<br />
65<br />
55°<br />
88<br />
89<br />
92<br />
35<br />
36<br />
90<br />
93<br />
42<br />
91<br />
37-41<br />
43<br />
94<br />
44<br />
99<br />
95,104-106<br />
50°<br />
100<br />
109<br />
101<br />
117,121<br />
107,108<br />
98<br />
96<br />
112<br />
110<br />
111<br />
1-3 6<br />
4,5<br />
8<br />
7<br />
9-13<br />
60°<br />
70°<br />
80°<br />
123<br />
122<br />
113 29<br />
114,116 139<br />
47,48<br />
14-22<br />
102<br />
45,46<br />
23 24,25<br />
26-28<br />
130<br />
49<br />
140-141 142<br />
124<br />
126,127<br />
50-52<br />
137<br />
55<br />
53,54<br />
64<br />
125<br />
128 129<br />
136<br />
138<br />
59<br />
56<br />
65<br />
67<br />
60<br />
58<br />
62<br />
131<br />
135<br />
133<br />
134<br />
132<br />
5° 0° 5° 10° 15° 20° 25°<br />
103<br />
74 75<br />
68-70<br />
71,72<br />
73<br />
77<br />
76<br />
66<br />
78 79<br />
81<br />
80<br />
50° 60° 70° 80° 90° 100° 110° 120° 130°<br />
82<br />
83<br />
84<br />
85<br />
86<br />
87<br />
A<br />
B<br />
61<br />
100<br />
100<br />
100<br />
90<br />
82<br />
57<br />
99<br />
97<br />
94<br />
83<br />
75<br />
62<br />
94<br />
73<br />
100<br />
99<br />
73<br />
70<br />
99<br />
97<br />
85<br />
69<br />
91<br />
91<br />
98<br />
94<br />
88<br />
71<br />
99<br />
98<br />
87<br />
85<br />
100<br />
100<br />
82<br />
66<br />
51<br />
66<br />
100<br />
99<br />
64<br />
61<br />
86<br />
80<br />
96<br />
84<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
98<br />
84 94<br />
81 74<br />
67<br />
62<br />
63<br />
83<br />
83<br />
100<br />
100<br />
72<br />
72<br />
61<br />
63<br />
96<br />
94<br />
79<br />
78<br />
Surget-Groba et al (2006)<br />
VB6<br />
VB16<br />
VB5<br />
VB1<br />
VB2<br />
VB3<br />
VB4<br />
VB7<br />
VB17<br />
VB18<br />
VB8<br />
VB11<br />
VB12<br />
VB13<br />
VB14<br />
VB15<br />
VB9<br />
VB22<br />
VB21<br />
VB10<br />
VU1<br />
VU3<br />
VU2<br />
VU4<br />
VU5<br />
VU6<br />
VU7<br />
VU8<br />
VU9<br />
OC1<br />
OC5<br />
OC2<br />
OC4<br />
OF2<br />
OF3<br />
OF1<br />
OF4<br />
PA1<br />
PA2<br />
VH1<br />
VH2<br />
OS4<br />
OS5<br />
OS3<br />
OS8<br />
OS9<br />
OS7<br />
OS10<br />
Podarcis muralis<br />
Lacerta bilineata<br />
Cla<strong>de</strong> E: Western Viviparous<br />
(Western and Northern Europe: 88-138)<br />
Cla<strong>de</strong> D: Eastern Viviparous<br />
(Eastern Europe and Asia: 50-87)<br />
Cla<strong>de</strong> B: Western Oviparous<br />
(Southern France and Northern Spain:<br />
30-44)<br />
Cla<strong>de</strong> C: Central Viviparous I<br />
(Austria and Hungary: 45-49)<br />
Cla<strong>de</strong> F: Central Viviparous II<br />
(Austria and Hungary: 139-142)<br />
Cla<strong>de</strong> A: Eastern Oviparous<br />
(Italy, Austria and Slovenia: 1-29)
100<br />
100<br />
100<br />
90<br />
82<br />
57<br />
99<br />
97<br />
94<br />
83<br />
“cold-tolerant” species: Zootoca (Lacerta) vivipara<br />
75<br />
62<br />
94<br />
73<br />
100<br />
99<br />
73<br />
70<br />
99<br />
97<br />
85<br />
69<br />
91<br />
91<br />
98<br />
94<br />
88<br />
71<br />
99<br />
98<br />
87<br />
85<br />
100<br />
100<br />
82<br />
66<br />
51<br />
66<br />
100<br />
99<br />
64<br />
61<br />
86<br />
80<br />
96<br />
84<br />
100<br />
100<br />
100<br />
100<br />
100<br />
100<br />
98<br />
84 94<br />
81 74<br />
67<br />
62<br />
63<br />
83<br />
83<br />
100<br />
100<br />
72<br />
72<br />
61<br />
63<br />
96<br />
94<br />
79<br />
78<br />
VB6<br />
VB16<br />
VB5<br />
VB1<br />
VB2<br />
VB3<br />
VB4<br />
VB7<br />
VB17<br />
VB18<br />
VB8<br />
VB11<br />
VB12<br />
VB13<br />
VB14<br />
VB15<br />
VB9<br />
VB22<br />
VB21<br />
VB10<br />
VU1<br />
VU3<br />
VU2<br />
VU4<br />
VU5<br />
VU6<br />
VU7<br />
VU8<br />
VU9<br />
OC1<br />
OC5<br />
OC2<br />
OC4<br />
OF2<br />
OF3<br />
OF1<br />
OF4<br />
PA1<br />
PA2<br />
VH1<br />
VH2<br />
OS4<br />
OS5<br />
OS3<br />
OS8<br />
OS9<br />
OS7<br />
OS10<br />
Podarcis muralis<br />
Lacerta bilineata<br />
Cla<strong>de</strong> E: Western Viviparous<br />
(Western and Northern Europe: 88-138)<br />
Cla<strong>de</strong> D: Eastern Viviparous<br />
(Eastern Europe and Asia: 50-87)<br />
Cla<strong>de</strong> B: Western Oviparous<br />
(Southern France and Northern Spain:<br />
30-44)<br />
Cla<strong>de</strong> C: Central Viviparous I<br />
(Austria and Hungary: 45-49)<br />
Cla<strong>de</strong> F: Central Viviparous II<br />
(Austria and Hungary: 139-142)<br />
Cla<strong>de</strong> A: Eastern Oviparous<br />
(Italy, Austria and Slovenia: 1-29)<br />
A One<br />
reversal<br />
Cla<strong>de</strong> E<br />
Cla<strong>de</strong> D<br />
Cla<strong>de</strong> F<br />
Cla<strong>de</strong> C<br />
Cla<strong>de</strong> B<br />
Cla<strong>de</strong> A<br />
Surget-Groba et al (2006)<br />
B No<br />
reversal<br />
Viviparous cla<strong>de</strong>s<br />
Oviparous cla<strong>de</strong>s<br />
Evolutionary transition
“cold-tolerant” species: Vipera berus<br />
• (GB) ad<strong>de</strong>r / common ad<strong>de</strong>r<br />
• (D) Kreuzotter
“cold-tolerant” species: Vipera berus<br />
Material & Methods<br />
80 samples, 60 diff. populations<br />
Blood, sloughed skin, scales, tip tail,<br />
tissues from different museums<br />
PCR, Sequencing (cyt b + CR)<br />
cyt b + CR = 1981 bp.<br />
45 haplotypes of the CR<br />
50 samples sequenced (CR + cyt b)<br />
<strong>Ursenbacher</strong> et al. 2006
Ch1<br />
Au1<br />
Ch3<br />
Fr1<br />
Fr3<br />
Ch2<br />
Ge<br />
Ne<br />
Sw2<br />
De<br />
Cz1<br />
Cz2<br />
Br2<br />
Sw1<br />
Sw3<br />
Br1<br />
No2<br />
Es<br />
Fi1<br />
Fi2<br />
Fi3<br />
Ru2<br />
Ru3<br />
Ru4<br />
Ru1<br />
Ru6<br />
Ru5<br />
Sk1<br />
Po<br />
Sk2<br />
Ro2<br />
Ro3<br />
Ro4<br />
Mo<br />
Bo3<br />
Se1<br />
Se2<br />
Bu4<br />
Bu2<br />
Bo1<br />
Bo2<br />
Au3<br />
It3<br />
Sv1<br />
Au2<br />
Sv2<br />
It2<br />
Ch5<br />
It1<br />
Ch5<br />
64<br />
73<br />
65<br />
91<br />
98<br />
94<br />
97<br />
95<br />
93<br />
-<br />
59<br />
-<br />
94<br />
90<br />
93<br />
54<br />
-<br />
64<br />
92<br />
91<br />
93<br />
92<br />
91<br />
93<br />
100<br />
100<br />
100<br />
87<br />
88<br />
88<br />
69<br />
90<br />
72<br />
88<br />
97<br />
92<br />
90<br />
94<br />
89<br />
94<br />
83<br />
94<br />
Ch1<br />
Au1<br />
Ch3<br />
Fr1<br />
Fr3<br />
Ch2<br />
Ge<br />
Ne<br />
Sw2<br />
De<br />
Cz1<br />
Cz2<br />
Br2<br />
Sw1<br />
Sw3<br />
Br1<br />
No2<br />
Es<br />
Fi1<br />
Fi2<br />
Fi3<br />
Ru2<br />
Ru3<br />
Ru4<br />
Ru1<br />
Ru6<br />
Ru5<br />
Sk1<br />
Po<br />
Sk2<br />
Ro2<br />
Ro3<br />
Ro4<br />
Mo<br />
Bo3<br />
Se1<br />
Se2<br />
Bu4<br />
Bu2<br />
Bo1<br />
Bo2<br />
Au3<br />
It3<br />
Sv1<br />
Au2<br />
Sv2<br />
It2<br />
Ch5<br />
It1<br />
Ch5<br />
64<br />
73<br />
65<br />
91<br />
98<br />
94<br />
97<br />
95<br />
93<br />
-<br />
59<br />
-<br />
94<br />
90<br />
93<br />
54<br />
-<br />
64<br />
92<br />
91<br />
93<br />
92<br />
91<br />
93<br />
100<br />
100<br />
100<br />
87<br />
88<br />
88<br />
69<br />
90<br />
72<br />
88<br />
97<br />
92<br />
90<br />
94<br />
89<br />
94<br />
83<br />
94<br />
ML tree; bootstrap value for 10'000 replicates: NJ, MP, ML<br />
“cold-tolerant” species: Vipera berus
“cold-tolerant” species: Vipera berus<br />
3 Refugial areas<br />
Italy<br />
Balkan peninsula<br />
Caucasus Mountains<br />
V. seoanei
Ch1<br />
Au1<br />
Ch3<br />
Fr1<br />
Fr3<br />
Ch2<br />
Ge<br />
Ne<br />
Sw2<br />
De<br />
Cz1<br />
Cz2<br />
Br2<br />
Sw1<br />
Sw3<br />
Br1<br />
No2<br />
Es<br />
Fi1<br />
Fi2<br />
Fi3<br />
Ru2<br />
Ru3<br />
Ru4<br />
Ru1<br />
Ru6<br />
Ru5<br />
Sk1<br />
Po<br />
Sk2<br />
Ro2<br />
Ro3<br />
Ro4<br />
Mo<br />
Bo3<br />
Se1<br />
Se2<br />
Bu4<br />
Bu2<br />
Bo1<br />
Bo2<br />
Au3<br />
It3<br />
Sv1<br />
Au2<br />
Sv2<br />
It2<br />
Ch5<br />
It1<br />
Ch5<br />
64<br />
73<br />
65<br />
91<br />
98<br />
94<br />
97<br />
95<br />
93<br />
-<br />
59<br />
-<br />
94<br />
90<br />
93<br />
54<br />
-<br />
64<br />
92<br />
91<br />
93<br />
92<br />
91<br />
93<br />
100<br />
100<br />
100<br />
87<br />
88<br />
88<br />
69<br />
90<br />
72<br />
88<br />
97<br />
92<br />
90<br />
94<br />
89<br />
94<br />
83<br />
94<br />
Ch1<br />
Au1<br />
Ch3<br />
Fr1<br />
Fr3<br />
Ch2<br />
Ge<br />
Ne<br />
Sw2<br />
De<br />
Cz1<br />
Cz2<br />
Br2<br />
Sw1<br />
Sw3<br />
Br1<br />
No2<br />
Es<br />
Fi1<br />
Fi2<br />
Fi3<br />
Ru2<br />
Ru3<br />
Ru4<br />
Ru1<br />
Ru6<br />
Ru5<br />
Sk1<br />
Po<br />
Sk2<br />
Ro2<br />
Ro3<br />
Ro4<br />
Mo<br />
Bo3<br />
Se1<br />
Se2<br />
Bu4<br />
Bu2<br />
Bo1<br />
Bo2<br />
Au3<br />
It3<br />
Sv1<br />
Au2<br />
Sv2<br />
It2<br />
Ch5<br />
It1<br />
Ch5<br />
64<br />
73<br />
65<br />
91<br />
98<br />
94<br />
97<br />
95<br />
93<br />
-<br />
59<br />
-<br />
94<br />
90<br />
93<br />
54<br />
-<br />
64<br />
92<br />
91<br />
93<br />
92<br />
91<br />
93<br />
100<br />
100<br />
100<br />
87<br />
88<br />
88<br />
69<br />
90<br />
72<br />
88<br />
97<br />
92<br />
90<br />
94<br />
89<br />
94<br />
83<br />
94<br />
ML tree; bootstrap value for 10'000 replicates: NJ, MP, ML<br />
“cold-tolerant” species: Vipera berus
“cold-tolerant” species: Vipera berus<br />
3 Refugial areas<br />
Italy<br />
Balkan peninsula<br />
Caucasus Mountains<br />
Sub-structure in the Northern Cla<strong>de</strong><br />
France<br />
Central Europe<br />
Romania<br />
Russia<br />
V. seoanei
“cold-tolerant” species: Vipera berus<br />
• V. berus in the Jura Mountains: isolated<br />
• Samples<br />
• 10 populations in the Jura Mountains<br />
• 3 in the Alps, 2 Massif Central,<br />
• 1 Atlantic Coast<br />
• Microsatellites<br />
• 13 microsatellites <strong>de</strong>veloped:<br />
• 7 without null alleles, follow<br />
• Hardy-Weinberg, not linked, …<br />
•<br />
<strong>Ursenbacher</strong> et al. 2008<br />
lac Léman<br />
Lausanne<br />
Neuchâtel
“cold-tolerant” species: Vipera berus<br />
• Allelic richness<br />
•<br />
Jura Mountains + Alps:<br />
A R ! 2.8 (2.11-3.14)<br />
Massif Central + Atlantic Coast:<br />
A R ! 3.9 (3.80-4.04)<br />
Possible reasons:<br />
• recent culling? (last centuries)<br />
• bottleneck? not <strong>de</strong>tected by<br />
BOTTLENECK (200-1500 years ago)<br />
• Ne: populations not smaller in the Alps<br />
• historical reasons: recolonisation after<br />
the last glaciation<br />
Jura Mountains Prealps<br />
Massif Central<br />
Atlantic Coast<br />
3<br />
2.5<br />
2<br />
1.5<br />
1<br />
0.5<br />
0<br />
4.5<br />
4<br />
3.5
“cold-tolerant” species: Vipera berus<br />
Jura Mountains + Alps:<br />
A R ! 2.8 (2.11-3.14)<br />
Massif Central + Atlantic Coast:<br />
A R ! 3.9 (3.80-4.04)<br />
Probable historical reasons:<br />
recolonisation after last glaciation<br />
(confirmed by the phylogeographic<br />
analysis)
Trends observed in reptiles<br />
• “warm” species<br />
• split during the Pliocene (3-5 Myr)<br />
• subsequent split (Pleistocene/Eocene)<br />
• reduction of genetic diversity during Pleistocene cold periods<br />
• “medium temperate” species<br />
• use of complementary refugia (not only Italy, Spain and Balkan Peninsula)<br />
• colonisation around drainage basins (species related to ponds - Emys)<br />
• “cold-tolerant” species:<br />
• split during Pleistocene
References<br />
• Fritz U, Guicking D, Kami H, Arakelyan M, Auer M, Ayaz D, Fernán<strong>de</strong>z CA, Bakiev AG, Celani A, D!uki G, Fahd S, Hava" P, Joger U, Khabibullin<br />
VF, Mazanaeva LF, #irok$ P, Tripepi S, Vélez AV, Velo Antón G, Win M (2007) Mitochondrial phylogeography of European pond turtles (Emys<br />
orbicularis, Emys trinacris) – an update Amphibia-Reptilia 28: 418-426<br />
• Hewitt GM (1996) Some genetic consequences of ice ages, and their role in divergence and speciation Biological Journal of the Linnean<br />
Society 58, 247-276.<br />
• Hewitt GM (1999) Post-glacial re-colonization of European biota Biological Journal of Linnean Society 68, 87-112.<br />
• Hewitt GM (2000) The genetic legacy of the Quaternary ice ages Nature 405, 907-913.<br />
• Poulakakis N, Lymberakis P, Valakos E, Zouros E, Mylonas M (2005) Phylogenetic relationships and biogeography of Podarcis species from the<br />
Balkan Peninsula, by Bayesian and maximum likelihood analyses of mitochondrial DNA sequences Molecular Phylogenetics and Evolution 37,<br />
845-857.<br />
• Surget-Groba Y, Heulin B, Guillaume CP, Puky M, Semenov D, Orlova V, Kupriyanova L, Ghira I, Smajda B (2006) Multiple origins of viviparity,<br />
or reversal from viviparity to oviparity? The European common lizard (Zootoca vivipara, Lacertidae) and the evolution of parity Biological<br />
Journal of the Linnean Society 87, 1-11.<br />
• Taberlet P, Fumagalli L, Wust-Saucy AG, Cosson JF (1998) Comparative phylogeography and postglacial colonization routes in Europe<br />
Molecular Ecology 7, 453-464.<br />
• <strong>Ursenbacher</strong> S, Carlsson M, Helfer V, Tegelström H, Fumagalli L (2006a) <strong>Phylogeography</strong> and Pleistocene refugia of the Ad<strong>de</strong>r (Vipera berus)<br />
as inferred from mitochondrial DNA sequence data Molecular Ecology 15, 3425-3437.<br />
• <strong>Ursenbacher</strong> S, Conelli A, Golay P, Monney JC, Zuffi MAL, Thiery G, Durand T, Fumagalli L (2006b) <strong>Phylogeography</strong> of the asp viper (Vipera<br />
aspis) inferred from mitochondrial DNA sequence data: evi<strong>de</strong>nce for multiple Mediterranean refugial areas Molecular Phylogenetics and<br />
Evolution 38, 546-552.<br />
• <strong>Ursenbacher</strong> S, Monney J-C, Fumagalli L (2008a) Population genetic structure in the remnant populations of ad<strong>de</strong>rs (Vipera berus) in the<br />
Swiss and French Jura Mountains: implications for conservation Conservation Genetics.<br />
• <strong>Ursenbacher</strong> S, Schweiger S, Tomovic LJ, Crnobrnja-Isailovi% J, Fumagalli L, Mayer W (2008b) <strong>Phylogeography</strong> of the nose-horned viper<br />
(Vipera ammodytes, (Linnaeus, 1758)): evi<strong>de</strong>nce of multiple refugia in the Balkan penisula Molecular Phylogenetics and Evolution 46, 1116-1128.
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