Endemic Taxa and Autochtonous Evolution in the Carpathian Basin

The level of endemism generally correlates with the geological age of the refugia where relict-like taxa have been evolved and/or could survive. The Carpathian Basin belongs to the geologically young areas of Europe. Its relief developed under the influence of the Alpine orogenesis and by retreat of the Paratethys and the Pannonian inland sea. Moreover, the phylogeography of some freshwater invertebrates (e.g. Neritidae snails, see: Bunje 2007; Feher et al. 2007) is clearly connected with the evolution of the Ponto-Pannonian water basin and of the Danube catchment area. In addition, there are several taxonomical groups with considerable proportion of endemic species, e.g. the land gastropods (Soos 1943) the earthworms (Lumbricidae: Csuzdi and Pop 2007) or some soil arthropods (e.g. Opiliones, Diplopoda: Korsos 1994; Collembola: Danyi and Traser 2007). Their core areas clearly coincide with the younger tertiary land masses within and near the Carpathian Basin.

Most endemic species are narrow specialists, inhabiting extreme habitats, such as thermal springs, karstic caves and karstic springs (Table 1). Several endemic troglobionta have been described in gastropods, pseudo-scorpions, harvestmen, spiders and springtails, often occurring within a single or a few caves of karstic mountains. Several species of earthworms, millipedes, centipedes and assels can be considered as holo-endemic species of the Western Transylvanian (Apuseni) mountains (Csuzdi and Pop 2007; Varga and Rakosy 2008).

Endemic terrestrial insects of the Carpathians are, as a rule, short-winged, flightless species such as the bush-crickets Isophya, Poecilimon spp.; some stenotopic relict grasshoppers (Capraiuscola ebneri, Podismopsis transsylvanica, Uvarovitettix transsylvanica, Zubovskia banatica; Kis 1965, 1980); numerous species of the

Table 1 Examples of stenotopic species of extreme habitats endemic to the Carpathians and the Carpathian basin

Taxonomic group

Species

Habitat type

Occurrence

Gastropoda

Melanopsis parreysi

Thermal springs

Baile Felix (Oradea)

Gastropoda

Theodoxus prevostianus

Thermal springs

Hungarian Middle Range

Gastropoda:

Paladilhia hungarica

Karstic water in caves

Mecsek Mts. (Abaliget)

Gastropoda

Paladilhiopsis

Karstic water in

Mti. Apuseni

transsylvanica

caves

Gastropoda:

Paladilhiopsis leruthi

Karstic water in caves

Mti. Apuseni

Amphipoda:

Niphargus tatrensis

Karstic water in

Calcareous mts. in the

caves

Northern Carpathians

Gastropoda

Bythinella (Sadleriana)

Karstic springs

Calcareous mts. in the

pannonica

Northern Carpathians

Palpigradi:

Eukoenenia vagvoelgyii

Karstic caves

Aggtelek karst,

Slovakian karst

Araneae

Centromerus chappuisi

Karstic caves

Mti. Apuseni

Araneae

Troglohypantes

Karstic caves

Mti. Apuseni

racovitzai

Araneae

Nesticus biroi

Karstic caves

Mti. Apuseni

Diplopoda

Haasea hungarica

Karstic caves

Mecsek Mts. (Abaliget)

Diplopoda

Hungarosoma bokori

Karstic caves

Mecsek Mts. (Abaliget)

Isopoda

Mesoniscus graniger

Karstic caves

Aggtelek karst,

Slovakian karst

Collembola

Pumilinura dudichi

Karstic caves

Aggtelek karst

Collembola

Protaphorura kadici

Karstic caves

Aggtelek karst

Collembola

Arrhopalites dudichi

Karstic caves

Aggtelek karst

Collembola

Arrhopalites

Karstic caves

Aggtelek karst

hungaricus

Collembola

Arrhopalites buekkensis

Karstic caves

Bükk Mts. (N. Hungary)

Coleoptera,

Duvalius bokori

Karstic caves

Slovakian karst

Carabidae

Coleoptera,

Duvalius gebhardti

Karstic caves

Bükk Mts.

Carabidae

Coleoptera,

Duvalius

Karstic caves

Calcareous mts. in the

Carabidae

microphthalmus

Northern Carpathians

Coleoptera,

Duvalius hungaricus

Karstic caves

Aggtelek karst,

Carabidae

Slovakian karst

Coleoptera,

Duvalius redtenbacheri

Karstic caves

Mti. Apuseni

Carabidae

Coleoptera,

Duvalius biroi

Karstic caves

Mti. Apuseni

Carabidae

Table 2 Examples of endemic species of forest soils in Transylvania

Lumbricida Lumbricida

Allolobophora dacica Octodrilus spp. (12 spp.!) Neobisium brevipes Ischyropsalis dacica Trachysphaera biharica Typhloiulus serbani Monotarsobius paucioculatus Monotarsobius spelaeus Monotarsobius burzenlandicus Lithobius dacicus Cylisticus major Trachelipus spinulatus Biharoniscus racovitzai

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Forest soil + litter Mti. Apuseni

Pseudoscorpiones

Opiliones

Diplopoda

Diplopoda

Myriopoda

Myriopoda

Myriopoda

Myriopoda

Isopoda

Isopoda Isopoda ground beetles (Duvalius, Trechus, Patrobus, Morphocarabus spp.) and weevils (e.g. Otiorrhynchus spp.). A bulk of these endemic taxa is confined to the Eastern and Southern Carpathians, to the Apuseni Mts.and to the mountains of Banat, which could preserve relict species (e.g. the tertiary relict gastropods Chilostoma banati-cum, Pomatias rivulare) or some narrow endemic species of Isopoda and Diplopoda (Table 2) in refugia without permafrost phenomena during the last glaciations (Bennett et al. 1991; Krolopp and Sumegi 1995; Willis et al. 1995).

In the more mobile insect groups, the proportion of endemism lies rather low (e.g. in Odonata no endemic taxa occur in the Carpathian Basin). Most endemic Lepidoptera of the Carpathian Basin belong to Microlepidoptera, which have flightless females and are strictly specialised to some food plants living on halo-phyta in the saline grasslands of the Ferto-Neusiedlersee area (Kasy 1965) and those of the Great Hungarian plain (Kiskunsag and Hortobagy). Endemic subspecies of Geometridae and Noctuidae evolved as peripheric isolates of turano-eremic species from the late-glacial, kryoxerotic periods, e.g. Narraga tessularia kasyi, Saragossa porosa kenderesensis (on food plants: Artemisia santonicum, A. pontica) and Hadula dianthi hungarica (on Gypsophila muralis). Some endemic taxa in the sandy areas of the Pannonian lowland are specialised predators or parasitoids, e.g. the spider Dictyna szaboi and the pompilid wasp Cryptocheilus szabopatayi. Further species described as endemics later proved widely-dispersed steppicolous species. In their majority, the endemics of the lower, hilly parts of the Carpathian Basin, however, represent thermophilous post-(inter?-) glacial relicts with connections to the Balkan Peninsula, Asia Minor or Southern Russia (e.g. Apamea sicula tallosi in warm-humid alluvial areas, Dioszeghyana schmidtii schmidtii and Asteroscopus syriacus decipulae in Pannonian xerothermic oak forests, Polymixis rufocincta isolata in the Villanyi Mts.; Chersotis fimbriola fimbriola, Euxoa vitta vitta, E. hastifera pomazensis and Cucullia mixta lorica in the dolomitic areas of the Transdanubian Middle Range, Chersotis fimbriola baloghi in the Aggtelek Karst). Balkanic connections have also been observed in butterfly species, which are restricted to special, Pontic-Pannonian steppic food plants, e.g. Plebeius sephi-rus (feeding on Astragalus exscapus, A. dasyanthus), Melitaea telona kovacsi (on Cirsium pannonicum).

Several endemic Macrolepidoptera subspecies of the Carpathians belong to the genera Erebia and Glacies (Table 3). A few endemic taxa are only widespread in the Carpathians and in the neighbouring mountainous areas, e.g. Aricia artaxerxes isse-kutzi, Photedes captiuncula delattini, while others are confined to the Southern Carpathians, often with Balkanic connections: Erebia cassioides neleus, Coenonympha rhodopensis schmidtii (Varga 1975, 2003a). The subspecies of Erebia melas inhabit island-like, calcareous mountain stocks, E. melas runcensis in the Apuseni Mts., E. melas melas in the Cernei Mts. and E. melas carpathicola in the Eastern Carpathians). All these data clearly demonstrate that the Carpathians, especially the Eastern and Southern parts, together with the mountains of Western Transylvania (Apuseni Mts. and Banat) can be considered as core areas for the survival and autochtonous evolution in many invertebrate groups of limited mobility.

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