The close geological and faunal connections of the Carpathians suggest the existence of highly dynamic contacts with the mountains of the Balkan Peninsula during the Upper Pleistocene. These connections show a contrasting picture compared to the refugia of the Iberian and the Appenine peninsula which have been much more sheltered by the glaciated mountains of the Pyrenees and the Alps, respectively. At least two major arboreal refugia can be traced here: the Illyrian refugium related to the Dinarids and its foothills and the Carpatho-Dacian refugium related to the Carpathians and its foothills. Some areas attached to these refuges served as periodic habitats over climatically favourable periods. These are regarded as fluctuation zones (Varga 1995; Sumegi et al. 1998; Deli and Sumegi 1999).
Since the Carpathian Basin occupied a transitional position between the Balkanic refugia and the cold-continental tundro-steppe zone during the glacial periods, the post-glacial re-population of the Carpathian Basin proceeded (1) by long-distance dispersal from the more remote (atlanto- and ponto-) Mediterranean and Southern Continental refugia, and (2) also from some adjacent local survival areas, e.g. from North-Western Balkanic ("Illyrian") versus South Transylvanian ("Dacian") arboreal refugia. In such cases, the arrows of the Northwards dispersal of the SouthWestern and South-Eastern populations surround the arid central part of the basin. These components of the flora and fauna extend Northwards through the foothills of the Eastern Alps and Southwest-Pannonian hilly regions on the one hand, and through the hilly regions of the Banat area and the Western foothills of the Transylvanian "Island" mountains (Apuseni Mts.), on the other. In some cases, the populations of the South-Western and South-Eastern "strains" do not display any significant taxonomical differentiation, e.g. the silver lime (Tilia tomentosa) or
Table 3 Endemic subspecies of Erebia (Nymphalidae, Satyrinae) and Glacies (Geometridae, Ennominae) in the Carpathians (maps of Erebia spp.: Varga 1975; Varga and Schmitt 2008)
Erebia manto traianus (Hormuzaki, 1895) Erebia epiphron transsylvanica
(Rebel, 1908) Erebia pharte belaensis v. Goltz, 1937 (= E. pharte carpatina A. Popescu-Gorj & A. Szabó, 1986) Erebia sudetica radnaensis
(Rebel, 1915) Erebia gorge rudkowskii
(Bang-Haas, 1933) Erebia gorge fridericikoenigi
(Varga 1998) Erebia cassioides neleus
(Freyer, 1833) Erebia melas melas (Herbst, 1796)
Erebia melas carpathicola (Alexinschi and Popescu-Gorj, 1959) Erebia melas runcensis
(König, 1965) Erebia pronoe regalis (Hormuzaki, 1937) Erebia pandrose roberti
(Peschke, 1920). Erebia pandrose cibiniaca
(Dannehl, 1927) Glacies canaliculata schwingenschussi (Wehrli, 1921)
Glacies noricana carpathica (Schwingenschuss, 1915)
Glacies bentelii retyezatensis (Bartha, 1933) -subspecific status dubious! Glacies coracina dioszeghyi (Schmidt, 1930)
Typical locality Rareu (NE Carpathians)
Bucegi (SE Carpathians)
(= Belanske Tatry) Rodna Mts. (NE Carpathians)
Rodna Mts. (NE Carpathians)
(SE Carpathians) "Banater Gebirge"
"Transylvania" (Mti Cernei)
Haghma^ul Mare (E Carpathians)
Cheile Runcului (Mti Apuseni)
Bucegi (SE Carpathians)
"Belaer Kalkalpen" (= Belanske Tatry), Mt. Bujacie Retezat Mts. (Lac. Zenoaga)
Retezat Mts. (Lac. Zenoaga)
NE-SE-S Carpathians the whole Carpathian arc
N + E Carpathians, S Carpathians
NE Carpathians, S Carpathians Tatra Mts., NE Carpathians
Mti Cernei (Domogled)
Tatra Mts, NE Carpathians
SE-S Carpathians higher mountains of the whole Carpathian arc
Belanske Tatry, Bucegi, Retezat
some butterflies and moths (Pyronia tithonus, Aplasta ononaria, Idaea nitidata, Zanclognatha tenuialis). Much more evidence is provided by the re-population of the Carpathian Basin from different directions in the cases of vicarious pairs of closely related species or in subspecies of polycentric species. Such cases can mostly be mentioned in land gastropods, e.g. Pomatias elegans - P. rivulare,
Chilostoma illyricum - Ch. banaticum, or in flightless insects, e.g. short-winged Orthoptera: Odontopodisma schmidti - O. rubripes, Isophya modestior - I. stysi (Orci et al. 2005).
The Western Balkanic ("Illyrean") influences are most significant in the Southern and South-Western parts of Transdanubia. These areas are characterised by a humid sub-Mediterranean climate and do not have a significant rainfall deficit in the summer period. They belong to the belt of mesophilous zonal forests of Fagion illyricum and Querco-Carpinion illyricum and the Illyrean-Pannonian hardwood gallery forests (Fraxino pannonicae-Ulmetum) characterised by a richness in tertiary/inter-glacial relict, often geophytic plant species (Horvat et al. 1974).
The Transylvanian ("Dacian") influences are connected with the forested areas of the Eastern Carpathians and often transmitted by the Western Transylvanian mountains (Mahunka 1993, 2007; Varga 1989, 1995, 2003a,b). The occurrence of Dacian elements is typical of the Eastern part of the Hungarian Middle Range, especially in the higher parts of the volcanic Eperjes-Tokaj range and in the Karst areas of N Hungary and S Slovakia. Eastern Balkanic influences reach also the Hungarian Middle Range by relict-like occurrences of some Balkanic and Balkanic-Anatolian elements (e.g. Noctuidae: A. syriacus and D. schmidtii), especially in the warm foothill zone where the sub-Mediterranean influences are also significant. Relict occurrences of Dacian elements (bush-crickets: Isophya stysi, Leptophyes discoidalis, Pholidoptera transsylvanica; ground-beetles: Carabus hampei ormayi) have been recently discovered on the small, island-like volcanic hills of the Bereg lowland.
The influences of the Northern Carpathians are also significant in the NE part of the Hungarian Middle Range. There is a characteristic difference between the Eperjes-Tokaj volcanic chain on the one hand, and the limestone plateau of the Bukk Mts. and the N Hungarian karst on the other. The biotic contact of the Eperjes-Tokaj range with the Carpathians is young, obviously post-glacial, and can be characterised mostly by the presence of species, which are either typical of the montane forest belts of the Carpathians (e.g. numerous land snails: Bielzia coerulans, Vestia gulo and ground-beetles: Carabus obsoletus, C. zawadszkyi, Abax schuep-peli) or widely dispersed in the Northern part of Central Europe, often having a Euro-Siberian distribution. The Bukk Mts., however, display an insular character. Its Carpathian and de-Alpine elements (e.g. land snails: Spelaeodiscus triaria, Phenacolimax annularis, the Geometrid moth Entephria cyanata gerennae) are isolated relicts. In the Aggtelek Karst area, the immediate contact with the higher limestone plateaus of Slovakia is combined with the occurrence of Carpathian (land snails: B. coerulans, Cochlodina cerata, Trichia unidentata; ground-beetles: C. obsoletus, C. zawadszkyi, A. schueppeli, Trichotichnus laevicollis carpathicus), boreal and xeromontane species at surprisingly low altitudes, influenced by the conspicuous meso-climatic and geomorphological features of this area. Some influences of the Northern and the Eastern Carpathians are to be observed at the NE marginal areas of the Pannonian lowland, i.e. along the upper course of the river Tisza and its tributaries (e.g. occurrence of land gastropods Vitrea diaphana, B. coerulans, Balea stabilis, Perforatella dibothrion, P. vicina).
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