are rare, but have been shown, e.g. for Rhododendron ferrugineum with five
(Charrier et al. 2014) and Cardamine alpina with a number of lineages that is
difficult to quantify (Lihová et al. 2008).
Similarly to the Pyrenees, two genetic lineages were frequently observed in the
Carpathians, with one of these in the mountains’ northern part and the second in the
eastern and southern regions (e.g. Pauls et al. 2006; Mráz et al. 2007; Ronikier et al.
2008a; Ujvárosi et al. 2010; Theissinger et al. 2012). An additional endemic genetic
lineage in the Apuseni mountains (island mountains in the Carpathian Basin) was
obtained for the stonefly Arcynopteryx dichroa (Theissinger et al. 2012). In
Soldanella species, two major lineages were distinguished, but in contrast to the
above pattern, the northern lineage also included the eastern Carpathians and thus
was geographically more extended. Furthermore, some geographically restricted
endemic species are known in this group (Zhang et al. 2001), hence underlining that
the biogeographic history of the high mountain elements of the Carpathians can be
more complex than the two-refugia theory.
The Balkan Peninsula presents a complex pattern of numerous small blocks of
high mountain systems in relatively close geographic proximity to each other.
Unfortunately, this area is still poorly studied phylogeographically. Morphological
studies in butterflies, however, strongly support an east-west split for several high
mountain species (Varga 1975), speaking for long lasting separation of the
mountain areas east and west of the Central Balkan Depression. This split was
supported by a genetic study of the stonefly Arcynopteryx dichroa (Theissinger
et al. 2012), but the butterfly species Erebia ottomana and Coenonympha
rhodopensis showed relatively uniform genetic constitutions throughout the area,
thus supporting the hypothesis that in these cases gene flow connected populations
throughout the Balkan mountains during glacial periods (Louy et al. 2013, 2014a).
3.3 Genetic Links Between High Mountain Systems
As Europe’s most important high mountain system, the Alps have multiple biogeographic links to all neighbouring mountains (Fig. 3.3; Schmitt 2009). In a
number of cases, in both animals and plants, identical genetic lineages exist in the
south-western Alps and in the Pyrenees (e.g. Kropf et al. 2002; Martin et al. 2002;
Schönswetter et al. 2002, 2004b; Gaudeul 2006; Schmitt et al. 2006; Reisch 2008).
This often repeated pattern is in most cases thought to be the result of glacial
distributions in the hilly areas of southern France and postglacial retreat into the
adjoining mountain ranges, leading to a rather young vicariance event, which is still
not mirrored in the genetic make-up of the now disconnected population groups.
Quite similar phenomena, with similar biogeographic explanations, are known
between the north-eastern Alps and the Tatra mountains (e.g. Kropf et al. 2003;
Muster and Berendonk 2006; Schönswetter et al. 2006; Suda et al. 2007; Paun et al.
2008; Triponez et al. 2011; Schmitt et al. 2014) as well as the south-eastern Alps
and the north-western Balkan mountains (Triponez et al. 2011; examples for
3 Molecular Biogeography of the High Mountain Systems …
67
(Charrier et al. 2014) and Cardamine alpina with a number of lineages that is
difficult to quantify (Lihová et al. 2008).
Similarly to the Pyrenees, two genetic lineages were frequently observed in the
Carpathians, with one of these in the mountains’ northern part and the second in the
eastern and southern regions (e.g. Pauls et al. 2006; Mráz et al. 2007; Ronikier et al.
2008a; Ujvárosi et al. 2010; Theissinger et al. 2012). An additional endemic genetic
lineage in the Apuseni mountains (island mountains in the Carpathian Basin) was
obtained for the stonefly Arcynopteryx dichroa (Theissinger et al. 2012). In
Soldanella species, two major lineages were distinguished, but in contrast to the
above pattern, the northern lineage also included the eastern Carpathians and thus
was geographically more extended. Furthermore, some geographically restricted
endemic species are known in this group (Zhang et al. 2001), hence underlining that
the biogeographic history of the high mountain elements of the Carpathians can be
more complex than the two-refugia theory.
The Balkan Peninsula presents a complex pattern of numerous small blocks of
high mountain systems in relatively close geographic proximity to each other.
Unfortunately, this area is still poorly studied phylogeographically. Morphological
studies in butterflies, however, strongly support an east-west split for several high
mountain species (Varga 1975), speaking for long lasting separation of the
mountain areas east and west of the Central Balkan Depression. This split was
supported by a genetic study of the stonefly Arcynopteryx dichroa (Theissinger
et al. 2012), but the butterfly species Erebia ottomana and Coenonympha
rhodopensis showed relatively uniform genetic constitutions throughout the area,
thus supporting the hypothesis that in these cases gene flow connected populations
throughout the Balkan mountains during glacial periods (Louy et al. 2013, 2014a).
3.3 Genetic Links Between High Mountain Systems
As Europe’s most important high mountain system, the Alps have multiple biogeographic links to all neighbouring mountains (Fig. 3.3; Schmitt 2009). In a
number of cases, in both animals and plants, identical genetic lineages exist in the
south-western Alps and in the Pyrenees (e.g. Kropf et al. 2002; Martin et al. 2002;
Schönswetter et al. 2002, 2004b; Gaudeul 2006; Schmitt et al. 2006; Reisch 2008).
This often repeated pattern is in most cases thought to be the result of glacial
distributions in the hilly areas of southern France and postglacial retreat into the
adjoining mountain ranges, leading to a rather young vicariance event, which is still
not mirrored in the genetic make-up of the now disconnected population groups.
Quite similar phenomena, with similar biogeographic explanations, are known
between the north-eastern Alps and the Tatra mountains (e.g. Kropf et al. 2003;
Muster and Berendonk 2006; Schönswetter et al. 2006; Suda et al. 2007; Paun et al.
2008; Triponez et al. 2011; Schmitt et al. 2014) as well as the south-eastern Alps
and the north-western Balkan mountains (Triponez et al. 2011; examples for
3 Molecular Biogeography of the High Mountain Systems …
67