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The Catalogue contains the basic taxonomic, nomenclatorial and
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BIOGEOGRAPHY AND EVOLUTION OF ORE ALLEPIDOPTERA IN THE PALAEARCTIC
D Griment of ZOOlOD and Evolution; Lossuth L. Universi H 0J0 Dehreceri, Huriga FThe concept of the oreat fauna can be defined as that typis ying orographi catly caused non-arboreal biomes. It is considered here as a major biogeographical unit in iis own right. not ora lyas a component of the oreOtundi at fauna. The oreat fauna is correlated with orographi catly determined non-arboreal ecosystems and iis members have, as a rute, insular, osten strictlyendemic or disjunci areas os distribution. Iis chorological centres can be regarded mostly ason ly potentiat centres of dispersal. They can be recogni sed by accumulated occurrerace of
stenochorous species and by the hi gli species-diversi ty os sonae typical genera. The orealfauna can he subdivided into an a ine type, as the faunal type of humid hi gli motintains illi prevat ling glaciat morphology and with strong connections to the tundrai Zonobio me. and Xeromontane type, as the faunal type os arid hi gli motintains with prevat ling physical weathering and with manifold connections to the eremic Zonobiome. The formation of the alpine faunal type is closely connected to the Quaternary glaciations. Iis history can be characteri sed by long-distance translocations and disjunctions. resulting in a great number Ofarctic-alpine species. On the contrary, the Xersmontane faunal type displays a more continu-ous evolutionary history extending far back into preglaciat times and also demonstrates ahigh potentiat for spectation in such groups whicli are adapted to the cold-arid conditiosas. Amajor part of the xeromontane fauna appears to have been stationary with a great number of relict- like species, especialty in Some core areas of Central and Inner Asia. Xero montane species could only populate Central and Northeria Europe during the extreme continentallate-glacial and early post-glacial phases. In Europe and adjacent areas. INOSt xero montane species occur in the summer-dry Mediterranean high mountains frona the Atlas to Asia Minor. The xero montane fauna can be subdivi ded into a W Palaearctic Mediterranean-xero- montane) and a Central - and Inner-Asiatic Continental) subtype. This bifurcation arosethrough the influence of a 'xero montane stillet V on a set os ancesti ai species coming mostlyhom seasonalty humid. solitheasteria Asiatic mountains. Thus, the fauna of subtropical. mon-soonic orobio mes se. g. in solitheria China and the Himalaya region) displays a sonaewhat intermediate. less disserentiated sancestral) character. The core areas of allopatric spectation dispersal and evolutionary history of Palaearctic xeromontane Noctu idae are considereil. Key wordso biogeography, oreat fauna, core areas, Alpine type, Xero montane type, Palae- arctic xero montane Noctuidae
OREAL FAUNA: CONCEPTS. PRINCIPI ES AND DEFINITIONS
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In lowland s the biotic composition of the most recent faunal range extensi oras predominate interacting with the prevat ling Zonobiomes . Even minor differences in the reli ef, e. g. canyon- like, deep, rocky ValleyS in table-land s, gorgesor step- like escarpments of plateaux can result in a heterogenei ty of the generalfaunal picture. The compositionat homogenei ty of the prevat ling Zonobio mehrealis down in response to narrow- scale en Vironmental factors whicli promote the furvivat os relict- like populations. We observe essentialty the fame phenomena in montainous regi Ons, where the macroclimaticalty influenced Zonat arrangement of major hi omes is disturbed by orographic factors. These conditions areexpressed in the high biotic diversi ty with in a limited area and in the coexistenceos different faunal types. Some of them are relicis of more ancient faunal in Vasions, which have furvived in restricted refuges. They are, as u rule, Osten members of relict- like communities or communi ty fragments that were more widely distributed in sonae former phases of the Quaternary climatic fluctuations. During climatic oscillations the more successi ut fraction of such previ ous ly restricted populations can irradiate, hence mountains also osten serve as Centres of disper- Sal. Thus, Severat components of orobiomes could be transformed during the ire volutionary history into components of Zonobiomes. These facts provide thebasis for severat ideas concerning the origin and history of tundi a talga and
The concePI Ofthe orealsauria The fauna specific to high mountains is osten referred io as the 'orecit una ' in the Zoogeographical literature, although mostly without a satisfactorydefinition of this notion. I wili try to place this concept with in the contexi of the hasic of the biomes division. Biomes not only represent types of ecoSystems, but also types of primaryproduction. Perhaps the most strii ing feature of the biosphere, shown on the map
This paper discusses the generat Datures of distribution and faunal historyof Palaearctic oreat Lepidoptera. A detailed analysis of the patierias of spectation
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in xeromontane Noctu idae with a sui vey of their biogeographical types wili bepresented in a SubSequent paper. Oretii ianu tun rest munia Here I recogni se the oreat fauna as a main Zoogeographical unit, not Only asa component of the 'ore tu drias' one scf. DE LATTIN l 96J). although a very intimate geographical and historical connection belween the tundrai and orealhiomes is evident. Their interaction can he seen during the entire Quaternaryperiod as a vast faunal mixing in the peri glaciat belis stili continuing at presentalong the motintain ranges of Easteria Siberia and Pacific North America Fig. l). On the other hanci, one also has to consider the main differences bet weentundi al and oreat faunas. The tundi at fauna be longs to one of the major ZOnobiomes, and most os iis members have a circumpo lar or amphi-Beringian distribution the possibility of an amphi-Atlantic connection is rejected, See KONONENKO et est l989. LA FONTA INE & WOOD l 988. MIRMOLA l98 P. MIMMOLAet til l99 l). The oreat fauna, in contrast, is essentialty not of a ZOnal nature. Its members are connected to numerous Orographicatly determined ecosystems and they Occur as a rute, in Spot- like, 'insularV, osten hight y endemic or widely separated areas, with an essentialty higher diversity of areal types si .e. 'faunal elementsV in the German tradition) than the members of the generat ly more uniformiundi at fauna the exceptional position of Beringia is discus sed in severat recent papers of KONONEN O et ah l 989. LA FONTA INE & WOOD l 988 and MIRROLAet ah l99 l. see abo ve). The frequently discussed arctic-alpine sor tundi o-alpine)distribution is only a specific case of formerly expansive, but recently restrictedand disrupted distribution ranges, occurring equat ly well IS con Vergent area regressions in species of oreal and tundi at origin. or somelimes having closely re
There are, howeVer, extended Oreal biomes without any connecti On or any compositional or physiognomical similari ty to the tundi at Ones, e .g. the Puria vegetation. the tali Compositae-Euphorbiaceae formation in East African hi glimOuntains, the xerophytic 'pol sterV-scrub vegetation in the motintains of the
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xerophytic formation S). Thus, the widely accepted definition of the oreat hi omes 'Diomes above the timherline ' is unsatisfactory for similar reasons VARGA
diversi ty of an iso lated molintain range depends in sonae cases on the distance frona the ''source of the coloni sation see also interaction belween distance, Sur-
tera , Orthoptera, passerine hird S etc. while the lalter are represented by numerous torrenticolous in secis se. g. the trichopteran genera DruSuS. PleCI OCHemici
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BI EOGRAPHY AND EVOLUTION OF ΟREAL LEPIDOPTERA
293 Rhyacophilia); by the petrophilotis Carabidae and land snails, e .g. the limestone-rock inhabiting Alopia species in the East Carpathians BOTOSANEANU l 962.
- refugiat iSoliation connected with evolving vicariant laxa by area regressionand disjunction: - isolation limeracit 'post-dispersat V. more exactly 'propterV-dispersal)connected with the peripheric fluctuation of the range VARGA l9Jl). Faunisti l and Diogeogri hical Suru of Palaearctic Noctuidae The facts on whicli the following discussions and conclusions are based
The taxonomy and distribution of the European Noctu idae is relative lywell-known. We have numerous check-lisis frona different paris of Europe
Data frona litige areas of Asia are much more incomplete and scattered. Many Species are known only frona a few type specimens collected during the last de- cades of the l9th century and preserved in sonae 'classic'' collections se. g. STAUD INGER. PUNGELER and BANG-HAAS collections in the Zoological Museum of the Humboldi University in Berlin. many famous collections in The Natural History Museum. London, the collection of the Grand Duke NIKOLAI MICHAILO VICH ROMANOFF in St. Petersburg. the CORTI collection in the Natural HistoryMuseum of Basel etc.). Aster a long period of very scattered collecting in the first half of this century the sui Veys of Palaearctic Noctu idae were intensifled during the l960s and the l9J0s by systematic collecting trips with more modern toois sportable gener- utorS, mercury Vapo Ur lampS, fluorescent iubes) by severat hight y quali fled ento- mologi sis in different paris of Western and Central Asia Tui key, Armenia, Iran. Afghanistan. Pakistan. Mongolia e .g. by G. EBERT, G. FRIEDEL. F. KASY, Z.
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KASZAB. C. M. NAUMANN. R. PINMER. J. PLANTE, E VARTIAN) and by the immense taxonomicat achievements of CH. BOURSIN. However, the majori ty of this materiat could only be evaluated during the last two decades. after the death of
BOURSIN. Because the collection and taxonomic notes of this author tinfortunate-ly have beco me practi catly inaccessi bie in the meanwhile, the specialisls whotri ed to continue his work were forced to re- study ali the type materiais originalty
examined and photographicatly documented by him.
These difficulties hecame, however, the staning hlock for new revision swith a more tip-to-date attitude and with more sophisticated methods se. g. thenew information coming froin the 'lock-and- eyV structu res of the everted in ner
Due to these recent faunistic investigations carried out in many. hitherio Very poorly Stud ted paris of Eurasia, a vast amount of ne w information on thetaxonomic composition and geographical distribution of the Palaearctic noctu id fauna was obtain ed. The furvey of these severat tens of thou sancis of noctu id specimens resulted in the description os severat hundred new species and thousantis of data on the distribution of hitherio incomplete ly known species. This descriptive and data collecting work occurred during a very intensive phase s see: References). Due to the description of a very large number of new species and by the study of many taxonomicalty incomplete ly known or mi Sinterpreted species sonae monographic studies have been stat ted on taxonomicalty complicated and highly diverse genera of Noctu idae. Thanks to these monogruplis, we Cun nowunderstand hetter the evolutionary processes and phylogeographic relations in many groups of Noctu idae. A new, Ρραyt-BOURSINV generation os noctu id speciali sis has emerged. for whom team-work and close cooperation is naturai. Thisis the reason why a large part of the distributionat data of Noctu idae materia is preserved in different museunas and private collections could he taken into consideration for the purpOSes of the preSent paper. J Xonomic rei'iSioris of Palaearctic ciuidae The extraordinary richness of the new noctu id materiai, and the favoti rabie conditions for a close cooperation in taxonomicat furveys, made possibi e thepublication. during the last ten to fifteen years, of a large number of longer re Vi Sional papers on many complicated and taxonomical ly-phylogeneticalty import-
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BI E RAPHY AND EVOLUTION OF OREAL LEPIDOPTERA
Some of these papers contain maps of distribution, biogeographical and phylogenetic considerations which could be used for the areographic and faunal historic Surveys of this paper. Further monographic works are imminent or are in a very intensive phase of preparation EuXOct. DichagyriS, CherastiS, RhyaCici, Dici rata, CardieStra, CardePici, DiSCeStra, Hadula, Thargetici, Odontelia, Ha deronia, Cteriscerat odia, Tricheu miS, Polla, SamgOSSa, SideridiS, ConiSania, Perigr hia, HarutiaeGogresPha, OrthsSia, Mythimna, Xylomota etc.) and will hepublished during the nexi law years. Because of these monographic wOrkS many hitherio uia solved phylogenetic problenas and biogeographical questions can beatas ered se. g. the major phylogenetic division os Noctuinae. Cuculliinae and
The delailed resulis of the phylogenetic and biogeographical analyses offeVerat xeromontane noctu id genera and specie S grOups, based on these revisionat works, will be discussed in a subsequent paper. Here, I can Only preSentsonae generat ideas abolit the taxonomic and hiogeographic diversity of xeromontane Noctu idae, whicli is obvious ly one of the most ancient and important so urces of the Palaearctic fuUnu. Chorological Princoles 'core areaS V in oreat his nesIn the Zoogeographical literature the term 'core areaSV in Germano Areal-
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boreat latina osten ite in hilly or montainous regions cf. DE LATTIN l 949. l 952 l9 P. t 96J. REI NIG l 93J. l 938. l 950). Faunal richness and high levet os ende-mism of European huiternies in motintainous regions of Southern Europe has also been clearly demonstrated by DENNIS et ah sl 99 l) and D ENNIS l 993). It is widely accepted that during the last glaciations the main sui vivat centres of the
western Palaearctic arboreat fauna were restricted to e. g. SOme paris of the Mediterranean peninsulas. In this phase of the genesis of the fauna, the arborealbion es were disconnected into severat larger or smaller PislandsV. Due to the post-glaciat re-afforestation, however the zonat character of these biomes couldbe restituted. combined with the reorgani sation os forested biocenoses.
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not) represent centre S of dispersal, as opposed to the arboreal and eremial ones hecati se the postglaciat expansion of the arboreat hi omes has fragmen ted these habitais into patches which could rema in sui table for the biota of the orobio me s. Ne Vertheles S, they may beco me expansi Ve, e . g. in the case of a future glaciationor also by aridi sation, which would lead to a retreat of the arboreal biomes. Frona these generat principies it cata be concluded that the core areas of theoreat fauna in this phase of our work can be defined main ly asi . centre S Os occurrence of a large number of stenochorous Vende mic V)speCieS, ii. centres with an accumulated occurre iace of disjunci subspecies of widely dispersed species with scattered distribution. ii l. centres with higher than average representation Os genera, whicli are typicat for Orobi OmeS.It seems qui te evident that the fame basic patierias are repeated in the distribution of the monophyletic, very clo Sely related allopatric SpecieS grou pS and
Fig. 1. Paleogeographical conditiosas during the last maxima os WUrm glaciations in Eurasia. Legend l α main ice shields, 2 α inland brackish water, 3 α major forest refugia. 4 α periglaciat tundrahelis. pseudoperi glaciat colu steppe and forest-steppe belis. 6 α Mediterranean weStern part), T continental desert and semi-desert areas