Sand Lizards Lacerta aglis Complex

Lacerta agilis Linnaeus, 1758, is a widespread species in Europe and Northwestern Asia. Nine subspecies have been described (Bischoff 1988). A phylogeographic analysis based on cytochrome b sequences (Kalyabina et al. 2001, Kalyabina-Hauf et al., 2004a) confirmed most of the subspecies, and revalidated a 10th one, L.a.tauridica Suchov, 1926, from Crimea. If the branching events (Fig. 4) are linked with the geographical distribution of the subspecies, the Caucasian/Black Sea region

L. trilineata

L. bilineata

L. viridis

' L bilineata 11

' L bilineata 155 L bilineata 12

■ L bilineata 13 L bilineata 14

■ L bilineata 17 L bilineata 18

■ L bilineata 19 L bilineata 27 L bilineata 28

■ L bilineata 43 L bilineata 53 L bilineata 15 L bilineata 41

L bilineata 51

L viridis 49 L viridis 4 L viridis 50 L viridis 5 L viridis 158 L viridis 52 L viridis 6 L viridis 7

Rhineland

Italy/

France/

Spain

Cres/Croatia Vosges

L viridis 22 Lp L viridis 35 L bilineata 48

Maribor/Slovenia

Kaiserstuhl/D

Greece

Northern Turkey

Fig. 3 Maximum parsimony tree for Green lizards, derived from mitochondrial cytochrome b sequences. Note that several individuals were initially determined as L. bilineata (from Trieste, Italy, and Kaiserstuhl, Germany), but have haplotypes of L. viridis, whereas lizards from the Croatian island of Cres are unambiguously L. bilineata emerges as the group's most probable area of origin (Joger et al. 2007). The most ancestral surviving clade is L. a. boemica Suchov, 1929, from the Eastern Caucasus. It may deserve species status.

The Western clade (distributed from Western Russia to the Pyrenees), in which the earliest internal branches lead to the easternmost subspecies - L. a. chersonensis and the next branching to an unnamed Carpathian group - must have moved gradually westwards.

The Eastern clade (from the Caucasus West to Crimea and East to Lake Baikal), and a Balkan subspecies (L. a. bosnica) are the other two major clades. As L. a. bosnica is sister of the Eastern clade, while the Western clade branched off from the common stem earlier, the most probable phylogeographic hypothesis must take into account several waves of colonization from East to West. A molecular

L. media

100,

LLLaaal2l2449

LLLaaa2ll2689

La lo5

(Greece)

Balkans (bosnica)

grusinica brevicaudata

SE. Europe Caucasus > Central Asia exigua group

La l92 rLta3249

1001

rto iLal5

Carpathian group

E. Europe (chersonensis)

boemica strigata L. viridis

Fig. 4 Maximum Parsimony phylogram (consensus of 300 equally parsimonious trees) of the Lacerta agilis complex, derived from mitochondrial cytochrome b sequences. L. media, L. strigata, and L. viridis are used as outgroups. Bootstrap values indicated. The L. exigua group unites subspecies from Caucasus (L. a. grusinica, L. a. brevicaudata) as well as Southeastern European exigua s.str. and various central Asian agilis

La l52 La l6o clock places these waves into different periods, from upper Pliocene to middle Pleistocene. The colonization of Central Asia appears to be more recent (maybe even Holocene).

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