장음표시 사용
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also in the subspecific divisions of many widely distributed polytypic species with disjunci areas. A detailed chorological survey and the phylogeographical analysis of Some monophyletic Species groups, and polytypic species of the Palaearctic Noctui dae mostly Noctuinae and Haden inae) will be publis hed in a
Most biogeographical and taxonomic publications deat with the description mapping and analysis of ranges of species. AS 'Dunal elementa , species sor sub- Species of polytypic Species. See DE LATTIN l 96J) are usualty considered as theuntis of biogeographical typisation. It appears to he evident that the ranges of species with some geographical congruences can be accepted as 'imperfeci repetitions , considered as basic objects of ΡGeographical EcologyV MACARTHURl9J3 P). There are numerous closely related concepis in the biogeographicalliterature whicli reflect essentialty the sanae regulari ty the partiat overlap of a large number of ranges in well-defined regions see the review of UDVARDYl98 l). KULCZYNSKI l 923) demonstrated the regularities of Overlap in a large number of boreal and arctic-alpine plant species. His method was ''systematiged
areiaXV. They unite those species areas that reflect more Or less concentric radiation hom so me core areas wherein ali areas of a certain geographical patiernoVerlap. He considered these centres as glaciat refugia hom whicli 'progressive species irradiate , while 'centrant'' species remain with in or near these refugia.
Practicatly the fame principie was nearly simultaneotis ly applied by REI NIG l93 P. t 950). who ouilined a number of climaticalty favoui able areas of the Ho-larctic region as glaciat refugia. STEGMANN sl 938) designated the set os species having 'aequi format V areas as a Dunal type, a term whicli has been used by severat authors se. g. DE JONG l9 2. VOOUS l 960. l 963). I consider this term to bepracticatly synonymous with DE LATTIN's 'Faunen reiXV whicli refers to the set os species sin polytypic species: Subspecies) having the sanae ''dispersat centreV
ΡAuShreiiungSzentrum ). He emphasi Zes the continuous series of dispersat capacity of species whicli connect the ''Stationa V species, restricted to the core areas to those whicli Could irradiate, i. e. the 'evansive'' members of the fame
faunal type. VARGA l9J a. h. l9JJ) suggested a simple modet for the centrifugat decrease of species belonging to a gi ven faunal type, describing it as an inverse function of the logistic equation. generat ly used in the population biology
Based on this mode l. the vicariotis stationary species and the disjunci sub- species of polytypic Species were characteri sed by a 'refugiat ' type of spectation. On the other hancl. the ' OSI-dimereas ' marginal iso lates of the expansive species
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299were contrasted with the former ones as cases of peripheric spectation and sub- spectation, respective ly. In addition the necessi ty of the analysis of the core areas of species-ricli genera was potiated out, especialty in those ones whicli consist ofa large number of 'stationary V species se. g. in the case of Lepidoptera species connected to orobio mes). The fame conclusion can he drawn by the analysis offlenochorous hi gh mountain species of Orthoptera LA GRECA l9JJ. LA GRECA
The idea that supraspecific taxa can he considered as uniis of hiogeographical analysis, is also possibie. KOSTRO ICKI l 969) considered genera as usefultoo is for statistical compari sons to ouiline faunal types and faunal regions asweli. Other authors emphasi Zed the phylogenetic significance of supraspecifictaxa. KRYZHANOVSKY l 965) argued that genera are uniis of Zoogeographica lanalysis hy 'common origin V, by essentialty the fame morphology and ecologyof species be longing to the sanae genus. PRAUDIN & MIS HI SHENKO l 980) ex
Fig. 2. Main areas of distribution and area connections of alpine and xeromontaneous biota. Legendo i m major areas of alpine biota horiZontalty halched with fuit lines). alpine and tundro-alpinearea connections black arrows: 2 α major areas of xeromontane biota verticalty Stripped), xero- montane areat connections hroken arrows
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methods. Units of the areographical analysis are the ranges Os species. Core areasdrawn by the distribution of stenochorous 'endemic ) species can he affirmed by the allopatric subspecies of polytypic Species, and by allopatric Semi-Species or sibi ing species of superspecies. Superspecies consisting os allopatric taxa of the Species-group are considered as the elementary monophyletic supraspecificuniis whicli are at ready adequale for phylogenetic analysis. ThiS procedure canbe continued by the phylogenetic analysis of monophyletic species groups withina genus, then by phylogenetic analyses of Subgenera and genera etc. ThOSe genera proved to be the most valvabie for the phylogenetic-biogeographical analysis in whicli there are numerous strictly stenochorous 'endemicV) Species, polytypic species with disjunci ranges, and also some expansive ones with large, continu- Ous range . Such genera can osten be subdivided into severat groups of species with closest phylogenetic relationships. It is essentiat that the whole multi-st Procedure ruriS fmm the taxonomicatly lower leveis to the higher ones and not
Generiai concePIS and termini in DiogeograPhyBecause a considerable part of the biogeographical literature is dispersed indifferent national or locat periodicats, publislied in many different languages Severat nearly Synonymous Scientific terins os biogeography have been introduced in parallel and used in the Anglo-American, French. German and Russianetc. literature. Hence, a comparative interpretation of some terins seems to heneceSSary Table l).
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Τahle 1. Comparative interpretation os sonae biogeographical ternas Biogeographicat entity consisting of taxa species or Subspecies)sharing the sanae Core uren
Set os laxa species or Subspecies)expanding Dona the fame core area
Taxonomic uni t smostly Species or subSpecies)belonging to a gi ven faunal type faunal element E: Only Faunenelement, rarely used in Englisti Geoelement G)biogeographical l iterature)Taxa with a wide range of distributione tent grOUp, expansive Arten G)widespread speciues E)Taxa of very limited range,
restricted to the com area endemic, stenochorous stenochore Arten, stationare
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DIVISION AND BIOGEOGRAPHICAL CONNECTIONS OF THE OREAL FAUNA
dominates in lower and moderate altitudes, and the snow-line is relatively low. The alpine and higher helis are characteri Zed by the well-known alpine geomorphological features, e .g. Sharp ridges, glacial cirques and long glacier longues reaching into the deep valleys susualty with a Zone- inversion), somelimes clown to the forested belis, especialty in the regions os expressed oceanic climate. Theselligh-lying open biomes are hordered, as a rute, by a timberline, and in the V m froneV of arboreal and non-arboreal biomes, the Zonation Os a Scrub-CΟmmunity ΡΚrummhola V scrvb-like Pinus, IunmeruS, Betulia, Rhododendron and different Ericaceae) is characteristic. Seasonat humidity, solifluction, the occurrence of peat-hogs and the regular snow-cover os long duration resemble conditions in the tundrai Zonobiome. Hence, numerous members of alpine faunal complexes hecame pre-adapted for dispersal into tundral si .e. periglacial) belis by the retreat os foresis during glacial phases. Os course, upward migration os tundrat species into alpine elevations was equat ly possibie, brought about by extensive inter- and post-glaciat re-afforestation, whicli has led to the formation os characteristic oreo-tundrai disjunctions. The alpine s. l.) faunal type was subjected to the most extensive migrations and area translocations during the oscillating glacial and interglacial phases. This faunal type is, on the other hand. at present most restricted in iis dynamics by ar
though in a more restricted sense fas ' Paleomediterran-Xemmontiane '). In the oreat helis of Xerom niane type, PhySical Neathering prOCeSSeS pre- dominate: thermal- inci . frost) fluctuation, leading to the formation os vastinasses of clastic rocks and graveis subjected to gravitational movemenis and tothe establishment of gravet pedimenis, ravitae pyramids etc. It is also important tonote that Xeromontane his nes osten do not couer inhole mountains, but forna mo-saic- like patierns of habitats. osten surrounded by alpine belis, e . g. in severat Eu-
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BI E RAPHY AND EVOLUTION OF OREAL LEPIDOPTERA
ropean or Solithern Siberian high mountains. On mountains sor paris of mountains) of xero montane character the snow-line lies essentialty higher than in comparabie molintains of the alpine type cf. MANI l 968 28), and the snow-cover isonen irregular and sparse . Primary production is inhibited not only by the ostenextremely cold winter season. but also by the seasonat aridi ty. Hence. the levet ofhioproductivity is os ten lower here than in the mountains of the alpine type, and their vegetation is semidesert- or desert- like se. g. Pamir plateati, Gobi Altri Mis paris of the Tibetan plateau etc.. WALTER & BOX l 983. WALTER & BRECK E
nobi Omes, their ecosystems are not. or only insignificantly de limited by the timberline. Heiace their elements are not inhibited frona dispersat in to the Zonal step- pes and deseris. This siluation was essentialty similar during the whole Quater- nary less influenced by the alternation of the cold and temperate phases. Thus
the xeromontane fauna could serve as an important fource for the genesis of the eremiC One. We Can Observe a nearly continuous transition of xero montane anderemic patierias of distributions in numerous genera of Noctuidae, being typicalfor arid belis in Central and Inner Asia.
Ovesternary dynamicS in the Xeromontane faunia Therefore. in the xeromontane fauna the range fluctuations and translocutions have been, as a rute, not as extreme as in the alpine One. Only during the extremely continental late glacial and earliest postglacial Ρkryoxerotic ) phases
bes ore the major postglaciat re-afforestation) a major fraction os continental elemenis, connected to open habitais, could spread into Europe, and the most Mediterranean-xei Omontane species could populate only the typical summer-dryMediterranean mountainous regions Atlas, arid paris of the Iberian peninsula solitheria Bal kan peninsula. Asia minor etc.). During the postglaciat vegetation history of Central Europe. there was not a single climatic phase which could have
Scarce representatives in Central and Northeria Europe are mostly 'pre-Litorina relicis, Occurring either above the tree-line in xeric habitais se. g. Central Alps) orin the uia forested habitats of moderate and low altitudes, determined by extremeedaphic andior microclimatic conditions. However the largest and obvious ly richest xeromontane hi omes of the Palaearctic occur, mostly in the mountainous massifs of Central and Inner Asia. Thehighlancis of Inner Anatolia and Armenia atready represent a fraction of this fascinating worid. but the greatest part of the Tien-Shan mountain system, the Panair-highland the magnificent chains of H indukush, Lai akoram and Trans-Hi
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BIOCE RAPHY AND EVOLUTION OF OREAL LEPIDOPTERA
ciat interstadiat periods and also in the Holocene; resulted, in these latitudes, in agenerat regression and disjunction of the oreal biomes and also of the distributionareas of their inhabitanis. The expansion of the arboreal biomes caused a wide diversification of siges and sh apes in the ranges of their faunal elements with anearly continuous Vari ance. Hence, it is osten not simple to recogniZe the station-ary and expansive extremes of the fame faunal type cf. DE LATTIN l9Jβ. l 964.l96J), e. g. Stationary MandZhurian species V s. trans-Palaearctic Eurosi heri an OnOS, Or stenoch Orous Mediterranean elements vs. widely distributed Mediter
On the other hand. the regressive disjunctions caused superficiat ly similarshapes of geographical ranges in oreal and oreo-tundrat species of different origin. Thus, the revision of the stenochorous species, or of the genera ricli insuch species, may be hight y significant for the analysis of the oreat fauna. The phylogenetic-biogeographical analysis of severat suci, cases will be attempted ina subsequent paper. Chorologiciat review of the alPine faunia Because the alpine latina is generat ly better known and carestilly analysed in iis delatis; I will present only a sholet. thesis- like review of state ments on iis
main Datures and tendencies here. l. The tundro-alpine disjunction, analysed in numerous publication S se. g.
NEC E l9β9). is a generat phenomenon in vascular planis and also in most majorgi ou pS of animal s. 2. The tundi o-alpine type of disjunction, and the Eurasian or even Holarctic extension os distribution show a strong positive correlation whicli confirms thathoth features must have a common historical background VARGA l9 β h. l 989c.l 995a) , i .e. capable these species were of Sprea ling Zonalty in to the peri glacialbelis during sonae more recent glacial phases smostly Riss andior Wiurna: Illinois
and wiscons in glaciation S, reSp.).3. The pure ly alpine s. l.) distribution is, on the Other hand, strongly correlated with regional se. g. European) or sub- regional siZe in range sonen endemicin sonae hi gli molintains with extended alpine eleVation S, C. g. PyreneOS, SO INCparis of the Alps; the highest massis s of the Bathan peninsula or - in the easternparis of the Palaearctic the litige molintain systems of the Altai and S an see VARGA l 989c). These species could survive the glaciat periods in molliatainous refugia at the margin of the motantain glaciations se. g. 'massiis de refuge , in thel ess glacialed alpine belis se. g. in the hi gli motantains of the Mediterranean penin
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sula) or - in speciat cases - in refugiat patches with in the major glaciations i. e. VnunatakV refuges, JANETSCHE K l9 6).4. Numerous widely distribu ted alpine species do not occur in the Northeria Calcareotis Alps or in the Northeria Carpathians. These mountains also Seem tobe poor in endemic high mountain species. The relative ly smali and scattered extension of the alpine bells. combined with un favori rabie climatic conditions ''Nord stati lage ) in these molintains can explain this fact. Numero us species groti ps whicli are mostly distributed in solitheria European hi gh mountains of at pine type, are osten represented by Vicariant species patrs in solithwestern and SotltheaStern Europe, respective ly, separated by this 'gapV. The presence of this
N Alpine - N Carpathian 'gapV gives iis the possibili ty for basic regionali sationof the European oreat fauna of the alpine type Fig. 4).
Fig. 4. Glaciat evenis as factors of regionat division in the western Palaearctic oreat fauna. t m Territories directly influenced by glaciations: Only alpine and arctic-alpine species. 2 Areas of sui vivat os relict-like alpine species displaying a western-eastern European Subdivision. osten with vicariant sister species subspecies: 3 m Main areas os distribution of xeromontane species