Post-Glacial Dispersal Strategies of Orthoptera and Lepidoptera in Europe and in the Carpathian Basin

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Post-glacial dispersal strategies of Orthoptera and Lepidoptera in Europe and in the Carpathian basin

Zoltan Varga

Abstract Ecologically transitional regions are characterised by high species diversity due to the overlap of species with different geographical origins caused by dispersal processes along gradients, e.g. the overlap of species belonging to different zonobiomes. The overlap of azonal communities along short- distance gradients results in the organisation of community-complexes and landscape mosaics connected by habitat ecotones characterised by overlap of several faunal types, like Mediterranean, Balkanic, Siberian, Ponto-Caspian, Ponto-Pannonian, Turano-Eremic and Xeromontane elements in the Pannonian forest-steppe complexes. For some invertebrate groups, such as the land gastropods or some soil arthropods, the Carpathian basin contains a considerable proportion of endemic species. The eastern and southern Carpathians, together with the mountains of western Transsylvania, can be considered as core areas of survival and autochtonous evolution in some invertebrate groups with limited mobility. In mobile insect groups only peripherically isolated sibling (sub)species have evolved, which display manifold biogeographic connections, for instance to the Balkan peninsula, Asia minor or southern Russia. The distribution of Mediterranean-Manchurian bicentric faunal elements, connected to the Ponto-Caspian waterway- system, shows long-distance disjunctions from the eastern Asiatic taxa. Gallery forests of the Illyrian and Pannonian types and wetlands accompanying the large rivers of the Pannonian lowland have served as corridors of the northward expansion of these species. The post-glacial repopulation of the Carpathian basin from different directions has been supported by Illyrian versus Dacian vicarious pairs of sister (sub)species. Cases of vicarious ‘Siberian’- Mediterranean sibling taxa have also been established. Recent results confirm that the montane part of the range of several ‘Siberian’ boreo-montane species does not have its origin in post-glacial ‘Siberian’ invasion. Populations of some butterfly species have been studied in different geographical scales. Patterns of metapopulation structure and of genetic differentiation have been established.

Key words: biogeography, Lepidoptera, Orthoptera, dispersal, Carpathian basin.

Introduction: biodiversity and 1985, Eldredge & Salthe 1984). Genealogical phylogeography diversity is usually connected with some core The continuity of life is maintained by distinct areas which can be outlined by areographical biological individuals of limited life-span. methods and tested by tools of the phylogenetic Similarly, the continuity of changing genetic analysis (Cracraft 1983, Cracraft & Prum 1988). information, called evolution, is carried by spe- The other aspect of the biological diversity is the cies of limited life-span. Species diversity is compositional one. Due to dispersal, overlap and generated by branching processes called specia- survival of species belonging to different bio- tion: by partition of the genetic variation into geographical types some regions can be distinct breeding systems with specific mate characterised by the accumulated presence of recognition signals (SMRS) and separated by floristic and faunistic elements of diverse age reproductive isolation. The subdivision of the and different geographical origin. formerly at least potentially coherent gene-pool The study of the structure and dynamics of results in the genealogical diversity (Eldredge ranges by genetic markers, called ‘phylogeo-

Proc. 13th Int. Coll. EIS, September 2001: pp. 93-105 (2003) 93 Proceedings 13th international colloquium European Invertebrate Survey, Leiden, 2-5 September 2001 graphy’ (Avise 1989, 1994, 1999, Avise & Hamrick 1996, Roderick 1996) has become to one of the most rapidly evolving fields of population biology. Phylogeny means the topology of the process of evolution at different hierarchical levels of organisation (Avise 1994): the level of genetic identity versus non-identity, the level of kinship in sexually reproducing organisms, the level of differentiation into ecotypes, geographical races and species (the level of emerging reproductive barriers) and the level of phylogenetic lineages separated by reproductive isolation (monophyletic supraspecific units). The study of genetic differentiation of genealogical lineages and the analysis of the processes govern- ing their geographical distribution (phylogeogra- Figure 1 phy) has demonstrated that the cladograms of The graph and sectors of the ‘centrifugally’ decreasing groups of related species (supraspecific taxa) species number of a faunal type under ideal conditions. should match the historical ‘area-cladograms’ of A. centrifugal decrease of the species number, the geographical ranges occupied by them (Eng- B. cumulative curve of the normal type of distribution, hoff 1993, 1995, Oosterbroek & Arntzen 1992). X. arithmetic mean of the normal distribution (Gauss The processes which generate spatial patterns of curve), I. point of inflexion, 1. section of the zonal dis- diversity, exhibit some basic features: tribution of the faunal type, 2. section of the dis- The genetic differentiation of populations aggregation (fluctuation) of the faunal type (with results in ‘elementary’ branching processes peripheric subspeciation), 3. section of the biogeo- which form a reticulating network of popula- graphical extinction of the faunal type. tions connected by more or less regular, or restricted gene-flow, as a consequence of full or limited exchangeability of individuals. zonobiomes and azonal communities in There are ‘hot spots’ (core areas of diversity), forest-steppe areas of the Carpathian basin characterised by the occurrence of numerous (Varga 1995). strictly endemic species. These areas appear In this paper, the biogeographical terminology is to be repeated more or less regularly in the adopted from Natura 2000 (Council of Europe different taxonomical groups. The ‘hot spots’ 1998). are considered to have a long-term ecological stability (Fjeldså & Lovett 1997, Fjeldså et Overlaps by dispersal along al. 1997), which cause the accumulation of gradients the genetic information (‘gene-centres’). The In the Palaearctic region, the most important pos- genealogical connections of endemic taxa, sibility of long-distance extension is the east- belonging to well-defined core areas, can be west dispersal along geographical latitudes, analysed by phylogenetic methods (‘areas of north to the chains of the Eurasiatic mountain endemism’, Harold & Mooi 1994). system. A large number of ‘trans-Palaearctic’, There are ecologically transitional regions Euro-Siberian areas has been established by this with high numbers of species, but without mechanism, which resulted in the formation of high proportions of endemism (Williams et several peripherically isolated and often taxon- al. 1999, 2000). They are characterised by omically differentiated populations at the zone of the overlap of ranges of species of different ‘disaggregation’ of the distribution at the moun- geographical origins caused by dispersal pro- tain regions of Central and South Europe (Varga cesses along gradients. An example is the 1977). Supposed the normal distribution of area overlap of species belonging to different sizes belonging to the same core area (to the

94 Varga - Post-glacial dispersal of Orthoptera and Lepidoptera in Europe and Carpathian basin same faunal type), the graph of the centrifugally especially in the boundary regions of lowland decreasing species number represents the inverse and hilly areas, with a variety of relatively stable of the cumulative graph of the normal distribu- versus mobile ecotones. tion, which can be subdivided into three sectors The orographically varied landscapes of Central by tangents of maximal convexity and concavity Europe and the Carpathian basin display several of the graph (fig. 1): the sector of 1. zonal distri- different cases of short-distance overlaps. bution, 2. disaggregation and 3. the extinction of Southern parts of Central Europe, North of the the faunal type in question. The normal distribu- Alps and Balkans (like southern France, southern tion of the area sizes will be performed only Germany, hilly parts of the Carpathian basin, under ideal conditions. Under real circumstances Williams et al. 1999, 2000) belong to the regions it is strongly influenced by climatic, orographical of Europe having the highest number of species and other conditions. For instance, the Pyrenees in vascular plants, in some groups of insects and and the Carpathians have served as effective bar- vertebrates, without a significant fraction of riers of dispersal in numerous atlanto-mediter- point-endemics. This clearly demonstrates the ranean (including Lusitanian) and ponto-medi- importance of the overlap of different floral and terranean (including Balkanic) species, respec- faunal elements populated this region post-glaci- tively. Thus, in these faunal types the number of ally from different core areas. species with restricted areas is relatively high, as The mosaic-like structure of the traditional illustrated by the numerous Iberian and Balkanic Central European cultural landscape with a endemics. variety of managed open habitats and their eco- In the Palaearctic region there are two main con- tones (hedges, ridges, lines of trees, forest fringes sequences of dispersal along gradients: etc.) in a forested matrix proved to be rather suit- 1. The overlap of zonobiomes along long-distan- able to preserve the connectivity of habitats and ce gradients: formation of transitions with over- also a high level of compositional diversity. A laps of ‘zonal’ sectors of several faunal types, large number of species has been successfully e.g. the overlap of Mediterranean (s.l.), Ponto- adapted to moderate perturbations (extensive Caspian and Siberian (West-Siberian, Mandzhu- grazing, mowing by hand, coppicing), typical for rian, East-Siberian) faunal types in the East traditional land use (fig. 2). European forest-steppe, with gradual decrease of the Mediterranean component and increase of the Endemism and autochtonous Siberian component along a west-eastern gra- evolution in the Carpathian basin dient (De Lattin 1957, 1967). This appears to be Core areas are characterised by numerous ende- typical for the orographically less varied eastern mic taxa. The level of endemism is generally cor- European (Boreal, Continental and Steppic) related with the geological age of the refugia regions. where relict-like taxa have been evolved and/or 2. The overlap of azonal communities along could survive (Fjeldså & Lovett 1997). From a short-distance gradients resulting in the organisa- geological point of view, the Carpathian basin tion of community-complexes (‘sigmeta’ of belongs to the youngest areas of Europe. Its relief German phytosociological tradition) and lands- has developed under the influence of the Alpine cape mosaics connected by habitat ecotones. orogenesis and by retreat of the Pannonian inland These community-complexes are characterised sea. There are, however, some taxonomical by the overlap of several faunal types, both in groups which show considerable proportion of zonal sectors and in sectors of disaggregation, endemic species, such as the land gastropods e.g. the overlap of Mediterranean, Balkanic, (Soós 1943), or some soil arthropods, like Siberian, Ponto-Caspian, Ponto-Pannonian, Opiliones and Diplopoda (Korsós 1994). Turano-Eremic and Xeromontane faunal ele- Most endemics are narrow specialists, inhabiting ments in the Pannonian forest-steppe complexes extreme habitats: thermal springs (Gastropoda: (Varga 1989a, 1992, 1995, 1996, 1997). They Melanopsis parreysi Philippi, Theodoxus pre- have been proved to be rather dynamic overlaps, vostianus C. Pfeiffer), karstic caves (Gastropoda: often with short-term fluctuation dynamics, Paladilhia hungarica Soós, Paladilhiopsis spe-

95 Proceedings 13th international colloquium European Invertebrate Survey, Leiden, 2-5 September 2001

been isolated in ‘massifs de refuge’ (Orthoptera: Isophya, Odontopodisma, Poecilimon species; the relict-like species: Capraiuscola ebneri Galvagni, Podismopsis transsylvanica Ramme, Uvarovitettix transsylvanica Bazyluk & Kis, Zubovskia banatica Kis; Kis 1962a,b, 1965, 1980; Coleoptera, Carabidae: Duvalius, Morpho- carabus, Patrobus, Trechus species; Curculio- nidae: Otiorrhynchus species) (some of these species have been suggested for inclusion into the Annex II-IV of the Natura 2000). The bulk of these endemic taxa is confined to the Eastern and Southern Carpathians, to the Apuseni mountains and to the mountains of Banat, which could preserve relict species (such as the Tertiary relict gastropods Chilostoma banaticum Rossmaessler, Pomatias rivulare Eichwald) or point-endemics (Orthoptera: Capraiuscola ebneri Galvagni, Podismopsis transsylvanica Ramme, Uvarovi- Figure 2 tettix transsylvanica Bazyluk & Kis, Zubovskia Habitat types and species diversity in the European cul- banatica Kis; Isopoda: Hyloniscus transsylvani- tural landscape. 1. virgin forest, 2. nature-like mixed cus Verhoeff, Protracheoniscus politus C.L. forest after logging and abandoning, 3. young forest Koch; Diplopoda: Entomobielzia gaetica after coppicing (‘Niederwald’), 4. grazed steppe forest Verhoeff, Karpathophilon dacicus Verhoeff) in (‘Hudewald’), 5. homogenous managed forest (‘Hoch- refugia of deciduous forests, in areas with not wald’), 6. dense medium-aged forest after repeated cop- frozen soil during the younger glaciations picing, 7. coniferous plantation, 8. deciduous, inten- (Bennett et al. 1991, Krolopp & Sümegi 1995, sively used plantation, 9. wooded pasture, 10. extensi- Willis et al. 1995). vely managed greenland (hayfield), 11. abandoned, In the more mobile insect groups, the proportion semi-natural grassland with tall-forbs (‘Versaumung’), of endemism lies rather low. For instance, in 12. abandoned, semi-natural grassland with tall-forbs Odonata no endemic taxa occur in the Carpathian and scrubs, 13. intensive greenland, 14. traditional agri- basin. Most endemic Lepidoptera of the Car- cultural land, 15. intensively used, monoculture-like pathian basin belong to Microlepidoptera, which agricultural land. Area surrounded with broken line: are strictly specialised to some food-plants and habitat types with high species diversity. the females are flightless, for instance species of Coleophoridae: Coleophora hungariae Gozmány (Camphorosma annua), C. peisoniella Kasy cies; Palpigradi: Eukoenenia vagvoelgyii Szalay; (Artemisia santonicum), Gelechiidae: Ilseopsis Amphipoda: Niphargus species; Isopoda: Meso- hungariae Staudinger, I. salinella salicorniae niscus graniger I. Frivaldszky; Diplopoda: E. Hering (Salicornia prostrata), living on halo- Brachydesmus troglobius Daday, Haasea hunga- phytic plants in the saline grasslands of the rica Verhoeff, Hungarosoma bokori Verhoeff; Ferto-Neusiedlersee´´ area and of the Great Hun- Carabidae: Anophthalmus, Duvalius, Typhlo- garian plain (Kasy 1965, 1981). Endemic sub- trechus species) and karst springs (Gastropoda: species of Geometridae and Noctuidae have Sadleriana pannonica Frauenfeld; Trichoptera: evolved in the same habitats, as peripheric iso- Drusus, Rhyacophila species, mostly in the lates of turano-eremic species from the late- Apus¸ eni mountains and the eastern and southern glacial, kryoxerotic periods, like Narraga tessu- Carpathians). laria kasyi Moucha & Povolny, Saragossa poro- The endemic insects of the Carpathians are, as a sa kenderesensis Kovács (Artemisia santonicum, rule, short-winged, flightless forms which have A. pontica) and Discestra dianthi hungarica

96 Varga - Post-glacial dispersal of Orthoptera and Lepidoptera in Europe and Carpathian basin

Wagner (Gypsophila muralis). Some endemic ly demonstrate that the Carpathians, especially taxa of the sandy areas of the Pannonian lowland the eastern and southern parts, together with the are specialised predators or parasitoids, such as mountains of western Transsylvania (Apuºeni the spider Dictyna szaboi Chyzer and the pompi- and Banat mountains) can be considered as core lid wasp Cryptocheilus szabopatayi Móczár. areas of survival and autochtonous evolution in Further species described as endemics later many invertebrate groups of limited mobility. On proved to be widespread steppicolous species. the other hand, in the mobile insect groups, only In majority the endemics of the lower, hilly parts peripherically isolated subspecies have been of the Carpathian basin, however, represent evolved, which display manifold biogeographic thermophilous post-(inter?-)glacial relicts with connections. connections to the Balkan peninsula, Asia minor or southern Russia (for instance, in warm-humid Relict species with long-distance alluvial areas: Apamea sicula tallosi Kovács & disjunctions in the Carpathian Varga, in Pannonian xerothermic oak forests: basin Dioszeghyana s. schmidtii Diószeghy, Astero- Several Mediterranean-Manchurian bicentric scopus syriacus decipulae Kovács, in the Villány faunal elements with disjunct ranges occur in the mountains: Polymixis rufocincta isolata Ronkay Carpathian basin. The distribution of this species & Uherkovich; in the dolomitic areas of the group is connected to the Ponto-Caspian water- Transdanubian Middle Range: Chersotis f. fim- way-system, and displays long-distance disjunc- briola Esper, Euxoa v. vitta Esper, E. hastifera tions from the eastern Asiatic vicariant taxa, pomazensis Kovács, Cucullia mixta lorica which are often only subspecifically differentia- Ronkay & Ronkay; in the Aggtelek Karst: ted (Lepidoptera: Apatura m. metis Freyer – Chersotis fimbriola baloghi Hacker & Varga). Apatura metis substituta Butler, Chariaspilates Similar connections can be observed in butterfly formosarius hungaricus Vojnits – Chariaspilates species, restricted to special, Pontic-Pannonian f. formosarius Eversmann, Rhyparioides m. food plants, like Plebeius sephirus I. Frivaldszky metelkanus Lederer – Rhyparioides metelkanus (Astragalus dasyanthus, A. exscapus), Melitaea flavidus Bremer, Arytrura musculus subspecies – telona kovacsi Varga (Cirsium pannonicum). Arytrura m. musculus Ménétries). Some species Some endemic Macrolepidoptera subspecies of of this group occur at the lower course of the the Carpathians belong to the genera Erebia Danube and Drava and also in swampy-boggy (E. epiphron transsylvanica Rebel, E. pharte areas of the lowlands in Transdanubia, in the belaensis Goltz, E. manto traianus Hormuzachi, Banat and eastern Hungary (Polypogon grypha- E. gorge fridericikoenigi Varga, E. pandrose lis Herrich-Schaeffer, Herminia tenuialis Rebel, roberti Peschke, E. pronoe regalis Hormuzachi) Diachrysia nadeja Oberthur and also the species and Glacies (G. coracina dioszeghyi Schmidt, mentioned above). The refugia of these faunal G. noricana carpathica Schwingenschuss, elements had probably been at the lower course G. canaliculata schwingenschussi Wehrli). Some of the Danube and its tributaries. Gallery forests endemic taxa are widespread in the Carpathians of the Illyrian and Pannonian type and alluvial and in the neighbouring mountainous areas, like wetlands accompanying the large rivers of the Aricia artaxerxes issekutzi I. Balogh and Phote- Pannonian lowland have served as corridors of des captiuncula delattini Varga, others are con- the northward expansion of these species. fined to the southern Carpathians, often with bal- Different types of long-distance disjunctions kanic connections: Erebia cassioides neleus have been observed in the relict-like steppe and Freyer, Coenonympha rhodopensis schmidtii semi-desert species. The polytypic butterfly Diószeghy. The subspecies of Erebia melas in- Melanargia russiae Esper, widespread in West habit island-like, calcareous mountain stocks, and Central Asia, South Siberia and in mountains Erebia m. melas Herbst in Cernei mountain, of Italy and Balkan peninsula, had locally occur- E. melas carpathicola P. Gorj & Alexinschi in red – as M. russiae clotho Herbst – on tall-grass the eastern Carpathians and E. melas runcensis clearings of birch gallery forests of the sandy König in Apus¸ eni mountain). All these data clear- lowland in Kiskunság. Its extinction was partly

97 Proceedings 13th international colloquium European Invertebrate Survey, Leiden, 2-5 September 2001 the consequence of overcollecting, but mostly of hypotheses. For instance, numerous genetic ana- destroying of the habitats (re-forestation with lyses could verify the existence of different gla- black locust Robinia pseudoacacia). The habitats cial refugia within the Mediterranean region of Chondrosoma fiduciarium Anker (Kasy 1965, (Assmann et al. 1994; Cooper et al. 1995, 1981) are also tall-grass lowland and hilly step- Demesure et al. 1996, Dumoulin-Lapègue et al. pes, often mixed with slightly saline patches. 1997, Hewitt 1996, 2000, Lagercrantz & Ryman Other species are confined to open dolomitic 1990, Schmitt & Seitz 2001a, Schmitt et al. rocky swards (Phyllometra culminaria Evers- 2001, Taberlet & Bouvet 1994, Taberlet et al. mann, Lignyoptera fumidaria Hübner, Cucullia 1994, 1998, Wallis & Arntzen 1989) and demon- mixta lorica Ronkay & Ronkay) or to open sandy strated, that several species have repopulated and rupicolous grasslands (Oxytripia orbiculosa Central Europe in the post-glacial period from Esper, also vanishing). The rapidly growing gar- different refugia in more waves. The ‘secondary den suburbs of the Buda hills and the metastasis- subdivision’ (De Lattin 1957, 1967, Reinig 1950) like expansion of ‘green-field’ investments of the Mediterranean refugium has also been around the capital are the most recent threatening repeatedly demonstrated, for instance in Chor- factors, often with considerable political and thippus parallelus Zetterstedt, in the European economical interests in the ‘background’. hedgehog (Santucci et al. 1998) or by the com- There are numerous halophytic food-plant speci- parative area analysis of several plant and mam- alists of the alkaline habitats, mentioned above, malian species (Cooper et al. 1995, Hewitt 1996, which are specifically or subspecifically diffe- Taberlet et al. 1998). The importance of the rentiated. They represent marginal isolates of Siberian refugium (s.l.) has been pointed out turano-eremic species, widespread in Trans- (Nève 1996, Taberlet & Bouvet 1994, 1998) and caspia and Central Asia. The common character- evidence has been found supporting some kinds istic feature of the area structure of these species of the Mediterranean-Siberian polycentry in is the large-scale disjunction between the several cases (Nève 1996, Schmitt 1999, Schmitt Carpathian basin and the Eastern Ucranian-South et al. 2000, Schmitt & Seitz 2001b). In contrast Russian steppes. with the opinion of De Lattin (1957, 1967), these results confirm the statement of the author Polycentric postglacial repopulation (Varga 1975, 1977, 1989b) that the montane part patterns in the Carpathian basin: of the ranges of several boreo-montane species areography and phylogeography has not been established by post-glacial ‘Sibe- The Carpathian basin belongs to the regions of rian’ invasion, but has been formed by South Europe, which have the highest biodiversity European montane refugia. Results of selected (Williams et al. 1999, 2000). Due to its transi- lepidopterological studies are summarized here. tional position during the Quaternary climatic fluctuations, the overlap and accumulation of the The Illyrian-Dacian vicariant taxa: the impor- floristic and faunistic elements of contrasting tance of the southwest-southeast ‘pincer’ habitats took place here. The overlapping of dif- Because the Carpathian basin had occupied a ferent climatic provinces, enhanced by the varied transitional position between the Balkanic refugia relief, edaphic and hydrographic conditions, has and the cold-continental periglacial loess steppe resulted in suitable conditions to survive for a zone during the glacial periods, the postglacial large number of species, belonging to different repopulation of the Carpathian basin has proce- core areas and displaying various patterns of ded both 1. by long-distance dispersal from the long-distance and short-distance re-populations. more remote (atlanto- and ponto-) Mediterranean The ‘molecular toolbox’ of phylogeography and southern continental refugia, and 2. from (Roderick 1996) have facilitated the recent re- some adjacent local survival areas, for instance visitation of many taxonomical and biogeogra- from Northwest Balkanic (‘Illyrian’) versus phical problems. ‘Hard’ evidences gained by South Transylvanian (‘Dacian’) Arboreal refugia. molecular studies could support, complete or In such cases the ‘arrows’ of the northward dis- correct some of the previous biogeographical persal of the southwestern and southeastern popu-

98 Varga - Post-glacial dispersal of Orthoptera and Lepidoptera in Europe and Carpathian basin lations surround the mostly arid central part of the type (Varga 1977). The ‘Siberian’ species are basin. These components of the flora and fauna widespread in temperate Eurasia (‘Euro-Sibe- extend northwards on the one hand through the rian’ or trans-palaearctic type of area), and usual- foothills of the eastern Alps and Southwest- ly exhibit a wide range of morphological varia- Pannonian hilly regions, and through the hilly tion and ecological valence (e.g. the butterfly regions of the Banat area and the western foothill species Maculinea arion Linnaeus, Melitaea of the Transylvanian ‘Island’ mountains (Apus¸ eni phoebe Denis & Schiffermüller, Mellicta athalia mountain), on the other. These areas are also cha- Rottemburg, Euphydryas aurinia Rottemburg). racterised by some secondary precipitation maxi- Their Mediterranean vicarious taxa (sibling spe- ma of sub-Mediterranean type during the ‘aequi- cies or semispecies) are usually confined to one noctial’ periods of the year. of the Mediterranean ‘secondary refugia’ (De In some cases the populations of the south- Lattin 1957, 1967): M. a. ligurica Wagner to the western and southeastern ‘strains’ do not display Adriato-Mediterranean, M. ph. telona Fruhstor- any significant taxonomical differentiation, for fer to the ponto-Mediterranean, P. m. malvoides instance in the silver lime (Tilia tomentosa) or Elwes & Edwards, M. a. celadussa Fruhstorfer some butterflies and moths (Pyronia tithonus and E. a. desfontainei Boisduval to the atlanto- Linnaeus, Maculinea arion ligurica Wagner, Mediterranean refugia. Aplasta ononaria Fuessly, Idaea nitidata Her- They also often have a considerable range of vari- rich-Schaeffer, Zanclognatha tenuialis Rebel). ation, in some cases due to the incomplete repro- Much more evident is the repopulation of the ductive isolation and hybridisation with the Carpathian basin from different directions in the ‘Siberian’ taxa. Habitats, food-plants and pheno- cases of vicarious pairs of closely related species logical traits of the Siberian-Mediterranean vicar- or in subspecies of polycentric species. Such ious pairs are often different. Maculinea a. arion, cases can be mentioned mostly for land gastro- for instance, is connected to short-grass swards pods, like Pomatias elegans O.F. Müller – with cushions of Thymus species (Thomas 1995), P. rivulare Eichwald, Chilostoma illyricum – which is the initial larval foodplant (exceptional- C. banaticum Rossmaessler, or in flighless insect ly also Origanum vulgare). On the contrary, the groups, like short-winged Orthoptera: Odonto- initial foodplant of Maculinea a. ligurica is podisma schmidti Fieber – O. rubripes Ramme, exclusively Origanum vulgare. The imaginal Isophya modestior Brunner von Wattenwyl – I. period is synchronised with the flowering period stysi Cejchan. In the latter pair of species, a clear of Thymus and Origanum, respectively, because differentiation in the stridulation and in allozyme the females usually deposit the eggs on flower pattern has been observed (Orci et al. 2001, capitula of the initial food-plant. Thus, sympatric Pecsenye et al. in prep.). In addition, some vica- populations of both are ecologically and sea- riant atlanto- versus ponto-Mediterranean sibling sonally isolated. The Siberian species, Melitaea pairs can also be observed in the more mobile phoebe is euryoecious, polyphagous (with food- insect groups, as in the butterflies Spialia serto- plants: Centaurea, Carduus, Cirsium species etc.) rius Hoffmannsegg and S. orbifer Hübner. and widespread in the Carpathian basin. In contrast, the Ponto-Mediterranean sister (semi-) The Mediterranean-Siberian polycentricity species M. telona is restricted to forest-steppe in the Carpathian basin mosaics with semi-dry swards, where its larval There are numerous cases of vicarious ‘Siberian’ food-plant, Cirsium pannonicum is abundant. (s.l.) – Mediterranean/arboreal and ‘Siberian’ Melitaea phoebe is regularly bivoltinous, while (s.l.) – Mediterranean/oreal sibling taxa. By M. telona only exceptionally produces a very ‘Siberian’ we understand all continental arboreal scarce second brood in the Carpathian basin. species dispersed north of the steppe zone and In the case of the Siberian – Ponto-Mediterranean which did not pass into the Mediterranean oreal pair of heath butterflies Coenonympha tullia region. They have formed the ultimate peripheric Müller and C. rhodopensis Elwes the isolation is isolates in the high mountains of this area, which complete, as the vicarious taxa occupy only topo- represent a ‘fluctuation belt’ of Siberian faunal graphically, but not ecologically overlapping

99 Proceedings 13th international colloquium European Invertebrate Survey, Leiden, 2-5 September 2001

from Monte Baldo (southern Alps; E. medusa hippomedusa Ochsenheimer) shows the most extreme differences. Erebia medusa psodea Hübner, which is externally the most different subspecies, occurring in the South Carpathians and East Balkans, was not studied electropho- retically. The highest level of genetic variation has been found in the East-Central European group, which can probably be derived from a large, extended East European (and Siberian?) Figure 3 refugial population of the last glacial period. The Sampling localities of Parnassius mnemosyne popula- observed genetic differences strongly support the tions. 1. Gilvánfa, 2. Misina (Mecsek mountains), hypothesis, that all subspecies have survived at 3. Hárskút (Bakony mountains), 4. Vérteskozma (Vértes least the last glaciation in different refugia. mountains), 5. Nadap (Velencei mountains), Nagyszénás (Budai mountains), 7. Pilisszentkereszt (Pilis moun- Genetic differentiation and metapopulation tains), 8. Magas-Tax (Börzsöny mountains), 9. Bán- structures in some butterflies in the yahegy (Bükk mountains), 10. Lusta valley (Bükk moun- Carpathian basin tains), 11. Gyertyán valley (Bükk mountains), Populations of some butterfly species have been 12. Hollóstetö (Bükk mountains), 13. Kékmezö (Bükk studied in the Carpathian basin at different mountains), 14. Sajólád, 15. Nagyoldal (Aggtelek karst), geographical scales. Populations of Parnassius 16. Nagy-Milic (Zemplén mountains), 17. Szokácsi mnemosyne Linnaeus and Euphydryas maturna forest (West-Transsylvanian hills), 18. Békés. Linnaeus have been compared both in larger and smaller regional scales, while populations of Aricia artaxerxes Fabricius and Maculinea alcon areas in the Dinaric karst area. Similar trends Denis & Schiffermüller – M. rebeli Hirschke have also been observed in the polymorphic ly- have only been studied in a smaller region. caenid Aricia artaxerxes Fabricius – inhonora Jachontov – allous Geyer – montensis Verity Parnassius mnemosyne: Populations from 11 superspecies, consisting of Siberian, European geographical regions (Transdanubian Middle (sub-)alpine and Mediterranean-oreal taxa. The Range: Bakony, Vértes, Velencei, Budai and Pilis phylogeographical analysis of such supraspecific mountains; Mecsek mountains; Northern Middle groups can explain some basic trends of the Quaternary faunal history in the Palaearctic. In this context, the results of Schmitt (Schmitt 1999; Schmitt & Seitz 2001b) proved to be especially relevant. He subdivided the European populations of Erebia medusa Denis & Schiffer- müller into five main groups, based on the study of 19 enzyme loci in 53 samples. The ‘West- Central European’ group (E. medusa brigobanna Fruhstorfer) consists of populations from France and Germany and one single sample from the Czech Erz mountains. The large ‘East-Central European’ group (E. m. medusa Denis & Schif- fermüller) includes all populations from Czechia, Slovakia and North Hungary. The third group of Figure 4 populations is formed by those of West Hungary UPGMA dendrogram of Parnassius mnemosyne in (and probably also West Balkans; E. medusa Hungary, constructed on the basis of Cavalli-Sforza narona Fruhstorfer), while the isolated deme and Edwards chord genetic distances.

100 Varga - Post-glacial dispersal of Orthoptera and Lepidoptera in Europe and Carpathian basin

A B

C Figure 5 Simplified geographical patterns of the species studied in Northeast Hungary. A. Hierarchical patterns in Parnassius mnemosyne samples. B. Hierarchical patterns in Euphydryas maturna samples. C. Hierarchical patterns in Aricia artaxerxes samples.

Range: Börzsöny, Bükk, Aggtelek karst, Zem- species, almost all the other investigated popula- plén mountains; lower foothills of the West- tions (except for Zemplén mountains and the Transylvanian mountains) and three lowland iso- Körös valley) should be also grouped to the lates (Sajolád, Dráva valley, Black Körös valley) same subspecies. The Zemplén and Körös valley were sampled in Hungary and the neighbouring populations proved to be genetically distinct. part of Rumania (fig. 3). The UPGMA dendro- Meglécz et al. (1997) demonstrated that the gram constructed on the basis of Cavalli-Sforza Bükk populations exhibit metapopulation struc- and Edwards genetic distances did not indicate a ture. Only occasional differentiation was ob- clear geographic pattern (fig. 3, 4, 5). The served between Bükk populations, while signifi- branching pattern among the populations from cant genetic differentiation was found between the Börzsöny mountains and Transdanubia did almost all pairs of populations from different not correlate with the geographical distances be- regions. Even the small, recently isolated tween sampling localities. Populations of the Sajolad population was found to be genetically Bükk mountains were clustered together, and different from the Bükk region. they also showed genetic similarity to the Sajolád isolate, which is only separated by 15 Euphydryas maturna: Most of our samples origi- km of unsuitable habitat from the Bükk moun- nated from three regions of the eastern part of the tains. These results support Eisner’s statement Pannonian lowland (Bereg-Szatmár lowland, (1954) who grouped most of the Transdanubian Nyírség and Black Körös valley), but also from populations into one subspecies (P. m. litavia). some populations in southern Transdanubia and When populations from Northeast Hungary the North Hungarian middle range. The isolated (except for the Zemplen population) were also population of the Sajólád forest was considered added to the Transdanubian data set, the Fst as a distinct region in this case as well. The value hardly increased. If we accept that the UPGMA dendrogram constructed on the basis of populations of the Vértes, Budai, Bakony and Cavalli-Sforza and Edwards genetic distances Velencei mountains are all members of one sub- shows that – as opposed to P. mnemosyne – most

101 Proceedings 13th international colloquium European Invertebrate Survey, Leiden, 2-5 September 2001 of the genetic variation could be observed Fst values at the PGM, GOT, EST, ME, ADH, among the populations within the regions, while ACON, GDH loci, and the Fit values at the GOT, the differentiation according to regions merely LDH, EST, ME, ADH, GDH loci. shows a lower level. Within all regions, all popu- The UPGMA dendrogram constructed on the lations showed significantly different Fst values. basis of Nei genetic distances, for both years Thus, no evident geographical differentiation has 1999 and 2000, showed a peculiar geographical been established (fig. 5). pattern also within the Karst region populations. The sample from the higher Zadielska planina Aricia artaxerxes: Two larger isolates of this (Hacava)^ was clearly separated from the other species, described as A. artaxerxes issekutzi ones, and the pattern of differentiation of popu- I. Balogh occur in the North Hungarian-Slova- lations essentially reflected the vertical subdivi- kian karst region and in the Bükk mountains. sion (higher versus lower karst plateaux) of the The number of alleles pro loci was higher in the region (fig. 5, 6). Karst region than in the Bükk mountains. Some rare alleles (e.g. Est, Got, Mdh, Pgi, Pgm) Summarizing conclusions: observed in the ‘Karst’ populations were lacking biodiversity and conservation from the samples of the Bükk mountains. On the priorities in the Carpathian basin other hand, the frequency of the heterozygotes The Carpathian basin, and within it Hungary, was not significantly lower in the Bükk moun- shows a high level of biodiversity, despite the tains. In addition, the genetic differentiation of fact that in Hungary there are no high mountains all samples proved to be significant. It means nor any littoral habitats. As an explanation of this that the level of genetic differentiation shows a richness, the transitional biogeographical posi- stronger correlation with the frequency distribution of the Carpathian basin can be mentioned tion of the more frequent alleles than with the (see paragraph 2). The highest level of biodiver- presence/absence of rare alleles. sity has been observed in some marginal areas Significant deviations from the Hardy-Weinberg where the overlapping of several different faunal equilibrium were found at several loci (Adh-1, components has taken place, like the western and Adh-2, Aldox, Est, Idh). The Fisher’s exact test southwestern parts of Transdanubia with the has shown a significant heterozygote deficiency overlap of Pannonian, Illyrean and Alpine influ- in both regions, which was mostly expressed in ences and also the northeastern edges of the the Aldox, Est and Me loci. These results are con- country with the overlap of Pannonian, Carpa- firmed also by the F-statistics. The populations thian and Dacian influences. Accumulation of showed a considerable level of genetic variance diverse faunal elements also took place at the (Fit), in which the ‘within populations’ fraction colline altitudes of the Hungarian middle range, proved to be larger than the ‘between popula- where the overlapping of Mediterranean (Holo- tions’ fraction (Fis>Fst). The Fis values were and Ponto-Mediterranean, Balkanic and Anato- significant at the LDH, EST, ME loci, while the lian) elements has succeeded. Conservation centres of relict species – often as endemic species/subspecies – are typical for the edaphically Nei’s Distance extreme habitats which could resist against the 0,1 0,05 0,0 postglacial re-forestation and could preserve the

Hacava^ elements of earlier climatic periods under speci- Haragistya al ecological constraints. These patterns of spe- Szelce cies diversity are well demonstrated by the mani- Tohonya Szögliget fold composition of insect assemblages of vege- Kecskelábrét tation complexes consisting of mosaic-like pat- Zsidórét terns of forest, skirt and grassland compart- Figure 6 ments. A nested hierarchy of diversity is present UPGMA dendrogram of the North Hungarian / South at levels of populations, communities, communi- Slovakian populations of Aricia artaxerxes. ty-complexes (sigmata) and landscapes.

102 Varga - Post-glacial dispersal of Orthoptera and Lepidoptera in Europe and Carpathian basin

Recently, the biogeographical division of Europe of their populations and metapopulation struc- has been completed by the addition of five tures at the landscape level. The solution of these further biogeographical regions: Arctic, Panno- problems has outstanding importance for the pro- nian, Steppic, Black Sea and Anatolian (Natura tection of the natural heritage of the Carpathian 2000, directive 92/43/CEE, 1997). It means, that basin, but also for Europe. due to its peculiar biogeographical character, the Pannonian region has been recognized as equiva- References lent to the further 11 large regions of Europe. The Assmann, T., O. Nolte & H. Reuter 1994. Postglacial ‘Areas of Special Conservation Interests’ was colonisation of middle Europe by Carabus auroni- completed with the list of threatened types of tens as revealed by population genetics. – In: Desen- natural habitats which require special conserva- der, K., M. Dufrene, M. Loreau, M. Luff & J.-M. tion measures (1998). In the Pannonian region at Maelfait (Eds.), Carabid Beetles: Ecology and Evo- least 36 of these habitat types occur, and some of lution. Kluwer Academic Publishers, Dordrecht: 3-9. the most significant European stands of them can Avise, J.C. 1989. Gene trees and organismal histories: a be preserved in this region. Studies on the bio- phylogenetic approach to population biology. – geographical composition of the flora and fauna Evolution 43: 1192-1208. of the Carpathian basin exhibit also some impor- Avise, J.C. 1994. Molecular Markers, Natural History tant applied aspects. and Evolution. – Chapman and Hall, New York - The indicative value of species largely depends London. on their biogeographical character. Thus, the Avise, J.C. 1999. Phylogeography: The History and habitat lists and the species lists of the European Formation of Species. – Harvard University Press, nature conservation information systems and Cambridge. conventions (like the CORINE-system, the Avise, J.C. & J.L. Hamrick 1996. Conservation Habitat Directive and Natura 2000) must be Genetics: Case Histories from Nature. – Chapman revised, according to biogeographical points of and Hall, New York - London. view. Threatened species with scattered occur- Bennett, K.D., P.C. Tzedakis & K.J. Willis 1991. rence in Central Europe are often widely distri- Quaternary refugia of north European trees. – buted in Eastern Europe. We have numerous spe- Journal of Biogeography 18: 103-115. cies, vanishing or being in decline in Central and Cooper, S.J., K.M. Ibrahim & G.M. Hewitt 1995. western Europe, which are represented in our Postglacial expansion and genome subdivision in region at least partly by strong populations. the European grasshopper Chorthippus parallelus. Strong, viable populations of such species can be – Molecular Ecology 4: 49-60. successfully protected mostly in these regions, Council of Europe 1998. The EMERALD Network – a inter alia, in the Carpathian basin (Euphydryas network of areas of special conservation interests. maturna, Lycaena dispar rutila, Maculinea telei- Strasbourg, 17. February 1998, Council of Europe, us etc.). In addition, the typical ecosystems of the pp. 1-39. Pannonian region (loess, sandy and alkali grass- Cracraft, J. 1983. Cladistic analysis and vicariance bio- lands, rupicolous swards, white oak scrub forests geography. – American Scientist 71: 273-281. etc.) exhibit numerous characteristic invertebrate Cracraft, J. & R.O. Prum 1988. Patterns and Processes of species which do not occur more westwards than Diversification: Speciation and Historical Con- the Hungarian middle range, or in some cases the gruence in Some Neotropical Birds. – Evolution 42: foothills of the Vienna Basin, South Slovakia and 603-620. Moravia. Numerous characteristic species of the Demesure, B., B. Comps & R.-J. Petit 1996. Chloroplast Pannonian region are connected with special DNA phylogeography of the common beach (Fagus habitat structures, like traditionally managed sylvatica L.) in Europe. – Evolution 50: 1515-1520. ecosystems, structured grasslands or forest- Dumoulin-Lapègue, S., B. Demesure, S. Fineschi, V. le skirt/grassland complexes of the Pannonian Corré & R.-J. Petit 1997. Phylogeographic structure forest-steppe (like many grasshopper and butter- of the white oaks throughout the European conti- fly species). Effective conservation of them is nent. – Genetics 146: 1475-1487. possible only by preserving of the connectivity Eisner, C. 1954. Kritische Revision der Gattung Parnas-

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Z. Varga Department of Zoology and Evolution, University L. Kossuth Egyetem-ter 1 H-4010 Debrecen, Hungary [email protected]

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