Centres of endemism of Noctuidae (Lepidoptera) in the Palaearctic arid mountains: biogeographical and phylogenetic implications

The oreal fauna is connected with orographically limited non-arboreal habitats. Its chorological centres can be recognised by the high species-diversity of numerous typical genera, and by the accumulated occurrence of endemic species and/or subspecies of disjunct species. The oreal fauna is partitioned to the alpine type, as the faunal type of humid high-mountains with strong connections to the tundral zonobiome, and the xeromontane type, as the faunal type of arid high-mountains with close connections to the eremic zonobiomes. As the results of revisions of several Noctuinae genera, species groups and/or sister species were recognised and their distributions were mapped. The restricted areas of allopatric sister species, often described by us as new for science, fulfil the criteria of the “ areas of endemism ”. Core areas of the Palaearctic xeromontane Noctuidae, outlined by the distribution of endemic species, have been proven by the occurrence of allopatric subspecies of polytypic species, and/or by the presence of allopatric sister species. In the revised genera of Noctuidae several types of allopatric speciation have been identified based on the analysis of the areas of endemism and of vicariance patterns. As a result of these analyses, it is proved that allopatric sister species, as elementary monophyletic supraspecific units, are suitable for phylogenetic biogeographical surveys. Although the major part of the xeromontane fauna appears to be range-restricted, a considerable fraction of the species could have expanded into the steppic zonobiome due to adaptive changes of thei r life cycles. High diversity of cold-adapted species originated from the Sino-Himalayan mountains by passing two main filter-corridors. One track of this bifurcation was directed across the “Rhododendron-corridor” to the Holarctic taiga zone while the other one, across the “Xeromontane filter-corridor” to the mountain systems of Central and Inner Asia. This bifurcation becomes apparent from the taxonomic division of the genera, composing both of these main faunal types. Supposedly, the faunal movements of the xeromontanean species in the West Palaearctic had been shaped by the Messinian salinity crisis and, additionally, significantly influenced by the Mid-Pleistocene climatic transition which deeply transformed the zonality of the vegetation by cooling and aridisation of vast areas. als elementare monophyletische supraspezifische Einheiten bestätigt, die sich als geeignet für phylogenetisch-biogeographische Untersuchungen erwiesen. Obwohl der größte Teil der xeromontanen Fauna engbegrenzte Areale aufzuweisen scheint, konnte sich ein beträchtlicher Anteil der Arten weit in die zonalen Steppengebiete durch die adaptiven Veränderungen der Lebenszyklen ausbreiten. Die große Mannigfaltigkeit der im sino-himalayanischen Hochgebirge entstandenen kälteangepassten Arten lässt sich durch das Durchdringen von zwei wichtigen Filter-Korridoren ableiten. Ein Zweig dieser Bifurkation verläuft durch den “Rhododendron-Korridor” in den holarktischen Taiga-Gürtel, während ein anderer Zweig durch den xeromontanen Filter-Korridor in die Gebirgssysteme von Zentral- und Innerasien verläuft. Diese Unterteilung lässt sich durch die unterschiedliche taxonomische Verteilung jener Genera verstehen, aus denen diese beiden grundsätzlichen Faunentypen bestehen. Es ist wahrscheinlich, dass die Ausbreitungsvorgänge der xeromontanen Elemente in der Westpaläarktis durch die Messinischen Salinitätskrise geformt wurden; einen weiteren wichtigen Einfluss besaß die mittelpleistozäne klima-tische Transition, während derer durch großräumige Abkühlung und Austrocknung die zonale Anordnung der Vegetation umgeordnet wurde.


Introduction -Oreal fauna, definition and subdivision
The composition of terrestrial animal assemblages strongly depends on the level of primary production, limited by the available solar energy and water circulation (Beer et al. 2010, Pappas et al. 2016, Simová & Storch 2017. Satellite images clearly reveal that both the Net Primary Production (NPP) and the Gross Primary Production (GPP) are the lowest in the tundral and desert zonobiomes. They are, however, similarly low in orobiomes, i.e. in different high mountains, surrounded by belts with a higher level of primary production (e.g. Rodin & Bazilevich 1966, Prince & Goward 1995, Running et al. 2004). If we simply specify the arboreal biomes (see: de Lattin 1967) as terrestrial habitats characterised by medium or higher level of primary production, the oreal fauna can be identified as the faunal type of the orographically defined non-arboreal biomes (Varga 1997). Its members, as a rule, have insular, endemic or disjunct areas in the Eurasiatic high mountains ( Fig. 1). Their core areas can be recognised by the high species-diversity of numerous typical genera and, thereby, the accumulated occurrence of endemic species and/or subspecies of disjunct species. Parallel with the non-arboreal zonobiomes, the oreal fauna can be subdivided into the alpine type, as the faunal type of humid high-mountains with prevailing glacial weathering and morphology, and with strong connections to the tundral zonobiome, and the xeromontane type, as the faunal type of arid high-mountains with prevailing physical weathering and manifold connections to the eremic zonobiomes. As the dynamics of the alpine faunal type is closely connected with the Quaternary glaciations, its history is characterised by long-distance range translocations and disjunctions, resulting in the diversification of a great number of arctic-alpine species (Varga & Schmitt 2008, Schmitt 2009). On the contrary, the evolutionary history of the xeromontane faunal type was supposedly less disturbed by glaciation-interglaciation cycles. Thus, it has a high potential for speciation in several groups adapted to the seasonally arid conditions as many Noctuinae moths belonging to the genera Euxoa, Dichagyris, Actebia s.l., Chersotis, Rhyacia, Standfussiana, Eugnorisma, Spaelotis, Xenophysa, etc. (Varga 1995, 2010b, Polyommatini blues (e.g. Polyommatus subg. Agrodiaetus, Eckweiler & Häuser 1997, Kandul et al. 2004, 2007, Lukhtanov et al. 2005, and also in some flightless grasshopper groups (e.g. Conophyma, Nocaracris, Nocarodes, see: Li et al. 2011Li et al. , Ünal 2016. Considering the composition of the European alpine Lepidoptera, high species diversity was found in several "Microlepidoptera" families and genera (Gelechiidae: , Huemer & Karsholt 2020Sattleria, Huemer & Hebert 2011, Huemer & Timossi 2014Yponomeutidae: Kessleria, Huemer & Mutanen 2015), certain genera of Geometridae (e.g. Entephria, Psodos, Charissa, Elophos;Huemer 1998, Müller et al. 2019 and Nymphalidae (Boloria, Erebia;Varga 1996, 2003, Varga & Schmitt 2008, however, surprisingly much less in the otherwise highly diverse Noctuidae (Varga 2003). This situation is contrasting with the outstanding taxonomic diversity of the xeromontane faunal assemblages in the Central and Inner Asiatic 1 mountain systems (Fig. 1, Varga 2010b) where numerous genera of Noctuidae (mostly of Noctuini, Hadenini, Xylenini, Apameini, and Oncocnemidinae) feeding on grasses and herbaceous plants predominate not only in the species composition but also in the density of individuals. Following these preliminary considerations, I developed the hypothesis that we can recognise some typically repeated patterns of vicariance in range-restricted sister species and, in parallel, essentially similar disjunct distributions in polytypic species in numerous genera of the xeromontane Noctuidae. Based on the taxonomic revisions of several diverse genera, mostly from Noctuinae, I also hypothesised that these patterns reveal important general trends of allopatric speciation which can be uncovered by phylogenetic biogeographical methods. Furthermore, I found it plausible to search for certain repeatedly appearing biogeographical and phylogenetic connections between the range-restricted xeromontane species and the zonally distributed congeneric steppic species. Finally, I hypothesised that the core areas of the Palaearctic xeromontane fauna are connected with the Central and Inner Asiatic high mountains as a consequence of the cooling and aridisation of these regions following the emergence of the Sino-Himalayan mountain systems and these core areas were modulated by the Mid-Pleistocene climatic transition.

Materials and methods
This biogeographical review is based on generic and suprageneric taxonomic revisions of several Noctuidae genera (mostly Noctuinae: Dichagyris, Actebia, Euxoa, Chersotis, Rhyacia, Standfussiana, Eugnorisma, Goniographa, Spaelotis, Xenophysa). Most of the type specimens of the revised taxa were studied; thousands of genitalia slides were prepared, photographically and graphically documented and electronically stored. In a book series (Varga et al. , 2015(Varga et al. , 2019 and in generic revisions (Varga & Ronkay 1987, 2002Ronkay & Varga 1999;Varga & Gyulai 1999;Varga 1998Varga , 2011aVarga , b, 2014Varga et al. 1989Varga et al. , 2018 numerous new genera, subgenera, species and subspecies were described, subgenera and species groups were separated, pairs or triplets of sister species were stated, and new taxonomical statuses were established. Based on the data of large European public and private collections, and of relevant literature, distribution maps of Noctuinae species were prepared, with special respect to allopatric sister species and disjunct polytypic species. Distributional data matrices of the genera Dichagyris, Chersotis, Rhyacia, Standfussiana, Eugnorisma, Goniographa, Xenophysa were prepared and the species were ordered to some types of distributions (Supplement 1), and they also grouped according to the types of geographical variations (Results 1). In the revised genera the species have been arranged into subgeneric groups, the allopatric sister species/subspecies relations were established, and based on them, area-dendrograms of species groups were constructed (Figs 4,5,13). In the first step, following the principle of the "areas of endemism" (Harold & Mooi 1994), I outlined some core areas by the accumulated occurrence of strictly endemic species (e.g. Kopet-Dagh Mts, parts of Hindukush Mts, the Pamirs and the Tien Shan system, etc. (Results 2). In the next step, these core areas were confirmed by the repeated patterns of disjunct polytypic species, and pairs (eventually triplets) of allopatric sister species. These species-group taxa were considered as elementary monophyletic supraspecific units (Results 3). Principally, this procedure is followed by the analysis of monophyletic species groups within a genus, then by phylogenetic analyses of subgenera and genera etc. It is essential that the whole multi-step procedure runs from the taxonomically lower levels to the higher ones. As the last step, according to the available life cycle data (aestivation, hibernation), some eco-physiological backgrounds of restricted distribution vs expansive area dynamics were uncovered (Figs 7,14,15,18; Results 3 and Discussion 1).

Results 2: The xeromontane fauna: areas of endemism and phylogenetic implications
A major part of the xeromontane fauna appears to be range-restricted with numerous relicl-ike species, especially in certain mountain systems of Central and Inner Asia. The restricted areas of the endemic species and of the allopatric sister-species correspond to the criteria of the "areas of endemism" (Harold & Mooi 1994), therefore, as monophyletic species groups, they are suitable for multi-step phylogenetic-biogeographical studies (see: Methods). As a result of taxonomic revisions, those genera proved to be most suitable for phylogenetic-biogeographical analyses in which (i) there are numerous strictly endemic species, (ii) there are also some polytypic species with disjunct ranges, and (iii) also some expansive species occur with large, seemingly continuous ranges. These highly diverse genera, as e.g. Euxoa, Dichagyris s.l., Actebia s.l. (subg.: Parexarnis, Protexarnis, Hemiexarnis), Chersotis (Varga 1998), Rhyacia, Eugnorisma (Varga, Ronkay & Yela 1990), Xenophysa (Varga 1989a(Varga , 2011 etc. can often be subdivided into some subgenera and/or several groups of species with closer phylogenetic relationships (Supplement 2). Eco-physiological responses to climatic challenges proved to be significant drivers of the ranges. The phylogenetic bifurcation of the Noctuinae genus Xenophysa (Figs 11-13) reflects a close connection with some essentially different climatic belts of the arid high mountains of Eurasia. The range of distribution of the western lineage ("X. junctimacula"-species group) reaches the central part of the Hindukush Mts, and its limitation clearly coincides with the eastern boundary of the sub-Mediterranean equinoctial type of precipitation (Agakhanjants 1981, Agakhanjants & Breckle 1995. The only widespread species pair of this group, X. cacumena and X. afghanorea shows a typical long-distance disjunction (Elburs and Kopet-Dagh Mts vs. Hindukush Mts) with occurrences in the western Hindukush massif (Koh-i-Baba) and the Paghman Mts. The presence of numerous range-restricted species of this group (X. argyrogramma, X. xenogramma, X. monastica, X. poecilogramma) seems to be confined to the central and eastern-south-eastern flank of the Hindukush mountain range (Fig. 11). The range of the eastern lineage ("X. agnostica"-group: X. agnostica, X. naumanni, X. pseudopoecila and the more isolated X. sharhu) extends from the Saravshan and Hissaro-Darwaz Mts to the western part of Mongolia ( Fig. 12), reaching also to the western Tien-Shan and Karakoram Mts. They have a narrow belt of overlapping with the western species group, mostly in the Hissaro-Darwaz range and in the eastern part of the Hindukush Mts. The most closely related sister species, the widespread X. agnostica and the restricted X. naumanni seem to be parapatric, while the distribution of the widespread X. pseudopoecila is partly sympatric with both species. However, X. poecilogramma, which belongs to the western group, locally co-occurs with both latter species in the eastern flank of the Hindukush range (Shandur pass, . The range of distribution of some genera or subgenera of Noctuidae, typical for xerothermic scrub-forests of western Asia (e.g. Ostheldera, see: Ronkay & Varga 1993; species groups of the large and heterogeneous genera Polymixis, Mniotype, Anchoscelis etc.) also extends to the same eastern boundary of the sub-Mediterranean type of precipitation in the Hindukush and western Tien-Shan ranges as the western clade of Xenophysa.
The variety of allo-and sympatric distributions of sister species, combined with disjunctions of the ranges, has been observed in the Eugnorisma chaldaica -E. spodia species group. The widespread E. chaldaica occurs from Central Anatolia and the south-eastern Russian steppes to southern Siberia, with subspeciation from southeastern Turkey to the adjacent areas of North Iran and also in the western Altai Mts (Fig. 14). Between the two major population groups of E. chaldaica a wide disjunction is gapping, from North Iran to the western Altai Mts in north-eastern part of Kazakhstan, which is "filled" by further taxa: E. kristenseni and the stenotopic E. spodia spodia in the Kopet-Dagh Mts, and the widely distributed E. kristenseni and E. spodia psammochrea in southern Kazakhstan and Uzbekistan, respectively (Varga & Ronkay 1987;Varga & et al. , 2015. In the Eugnorisma ignoratum species group (Fig. 15)  The genus Chersotis is rich in pairs and/or groups of closely related species (Tables 3-4, Appendix II). Here those allopatric species-groups are treated which are distinguished by recognised synapomorphies of genitalia. Significant characters of these species-groups are already described and illustrated in taxonomic revisions (Varga 1996b(Varga , 1998 . 5)). The Ch. rectangula-andereggii species group can be regarded as outgroup of this species-group. The next "triplet" of species uniformly shows some characteristic features of genital structures. The westernmost species, Ch. binaloudi occurs in Khorassan (NE Iran), and its sister species are the "twins": Ch. antigrapha (Hindukush Mts) and Ch. argyllographa (W Pamirs range, Fig. 10). We consider the set of the Ch. vicina group and Ch. juvenis group as their sister-group (the taxonomic details and species descriptions are published by Varga 1996bVarga , 1998Varga & Ronkay 1996; Fig. 8). All these species share the acute process of the juxta as synapomorphy. Their probable sister-group is the widely distributed, polytypic R. subdecora. Two further species pairs also belong to this subgenus, the partly sympatric R. psammia* (polytypic, widely distributed) -R. nyctymerina (Tien Shan) and the allopatric twin species R. diplogramma (Tien Shan system) and R. oromys (Pamirs, Hindukush, West Himalayas). All of these species groups share the synapomorphies as follows: the modified clavus densely covered by small spines, the hook-shaped carina of the aedeagus, and the large subbasal and the conical, sclerotised medial diverticula of the vesica (Varga , 2011. In some other cases, there is a partial overlap in the distribution of the closely related species. These overlapping species have a fairly large extended range with subspecific subdivision (e.g. the species pairs Chersotis rectangula and Ch. andereggii, Ch. elegans* and Ch. anatolica*, Ch. fimbriola* and Ch. laeta*, Fig. 16). As all of these species are widely distributed and mostly polytypic*, their distribution patterns should be considered as cases of secondary overlaps, subsequently to the previous allopatric process of speciation. The geographical extension of the allopatric areas of sister species may be rather different.  (Fig. 17).    However, the existence of a few, phylogenetically strongly separated species (e.g. Ch. sjuntensis, Ch. glebosa, Ch. illauta) supports the hypothesis, that in this genus also an older wave of speciation should have proceeded, from which most of the species became extinct. This hypothesis is strongly supported by such cases in which the species-groups consist of one strongly differentiated species, which can be regarded as outgroup of all other members of the species group, e.g. Ch. larixia* (with extremely fragmented range of distribution) for the Ch. elegans-kacem-eberti-anatolica group. All these cases refer to an ancient and a subsequent, more recent wave of the allopatric speciation in the genus Chersotis.   (1), and three closely related ones: G. naumanni (1), G. metafunkei (2), G. funkei (3). The sister species (2) and (3) are strictly allopatric, (1) and (2)

Discussion 1: Climatic constraints in the life cycle evolution of Noctuidae
In Noctuidae there are several types of univoltine life cycles, adapted to the seasonally cold and/or arid, temperate climate by different forms of larval and/ or imago diapause (Ryabov 1956;Sukhareva 1999;Saulich et al. 2017). Such univoltine life cycles probably can be derived from ancestral polycyclic types due to the insertion of diapause periods as hibernation and/or aestivation (Fig. 19). In these cases the larvae can use some resources of short duration by the right timing of active periods, and they can also be adapted to the seasonal insufficiency of resources by the diapause. There are several, in parallel developed different combinations of life cycles, connected to the centres of species diversity, mentioned above. One of the typical responses of noctuid moths to the aridity is the aestivation of the larvae (most often in the prepupal stage, Specht et al. 2013), and/or of the adults, which has evolved in parallel in many different taxonomical groups (Fig. 19), mostly in the subfamily Noctuinae (Euxoa, Dichagyris s.l., Actebia s.l., Chersotis, Rhyacia s.l., Standfussiana, Noctua, Spaelotis etc.). The well-developed abdominal fat bodies obligatorily serve as pre-requisite of the successful aestivation and reproduction after the aestivation of the adults.
In the "capnistis"-group of the genus Chersotis there are four stenotopic endemic species (Fig. 7) in high altitudes of arid high mountains of West and Central Asia, without well-developed fat bodies, while the sister species Ch. capnistis* and Ch. leucostola have large fat-bodies and a life cycle with imaginal aestivation, combined with  a relatively large range of distribution (Varga & Ronkay 1997: 115), extending into moderate altitudes in steppe areas of the southern Ural Mts (Ch. capnistis) or the Dzhungarian Ala-Too (Ch. leucostola). The possibility to gain a "zonal" expansion into the steppe biome seems to be linked to those noctuid moths (mostly Noctuinae) which either emerge at early summer and subsequently have a post-aestivation dispersal period in the late summer or early autumn (Fig. 19) (Figs 14-15). As one of main results of our preliminary taxonomic surveys (e.g. Ronkay & Varga 1999;Varga 1996bVarga , 1998Varga , 2011Varga & Ronkay 1997;, taxonomic groups of the Mediterranean xeromontane fauna appear to have two different main sources of origin. The bulk of genera probably can be derived from the primary bifurcation of the Palaearctic xeromontane faunal complex (Fig. 20). The other group had evolved from diverse Mediterranean xerophilous arboreal groups by adaptation to the aridity in connection with the late Tertiary Messinian Saline Crisis (Varga 1997(Varga , 2010b). This hypothesis is strongly supported by the "macro"taxonomic duality of the Mediterranean xeromontane Noctuidae. Those genera which belong to the subfamily Noctuinae and have "cutworm"-type, terricolous larvae, obviously originated in the continental, western and central Asiatic orobiomes (e.g. Euxoa, Dichagyris, Chersotis, Rhyacia, Standfussiana, etc.). Their Mediterranean representatives regularly belong to some different derived phyletic lines (subgenera) of these polytypic genera. Their East Mediterranean-Anatolian taxa often display western and Central Asiatic connections but only in a few cases a marginal speciation in the Mediterranean ranges. This group of genera correspond to the "saxobiotic ecofaunal complex" of the Palaearctic Orthoptera, described by Bey-Bienko (1948; see also Shumakov 1963, Pravdin & Mishtshenko 1980   Tien-Shan ranges). Their ranges greatly overlap with the core areas of the Noctuidae genera mentioned above.

Discussion 2: Subdivision and history of the Palaearctic xeromontane fauna
In Europe and adjacent areas the majority of xeromontane species occur in the summer-dry Mediterranean mountain systems, from the Atlas Mts to Asia Minor (Stegmann 1938, Voous 1960, Kryzhanovsky 1965, Varga 1997. To the contrary, Central and Northern Europe just holds a few species of this faunal type and were only colonised by xeromontane species possibly during the cold-continental younger glacial and early post-glacial phases only Varga 1996Varga , 2010aVarga , 2010b. A much higher diversity of xeromontane species was observed from the arid high mountains of East Anatolia to the mountain systems of Central and Inner Asia, mostly due to the high species numbers of some typical genera e.g. in Noctuidae (subfamily Noctuinae), Lycaenidae: Polyommatini, Orthoptera: Catantopidae (Varga 1996(Varga , 2010bEckweiler & Häuser 1997;Pravdin & Mishtshenko 1980;Ünal 2016). Thus, the composition of the xeromontane fauna is subdivided into a West Palaearctic (Mediterranean-xeromontane) and a Central-and Inner-Asiatic (Continental) sub-type. The protracted biogeographical confusion in the use of terms Central Asia vs. Middle Asia, or Inner Asia (used historically as "Innermost Asia", or "Died Heart of Asia", see: M. A. Stein 1928) has been clarified by the recent paleo-ecological surveys. They have shown a deep historical split between the more western Central Asiatic and the Inner Asiatic (Mongolian-Tibetan) steppe biota, both in the composition of vegetation, and in the mammalian assemblages (Barbolini et al. 2020). This split is clearly reflected by the differentiation of the faunal composition of the more western Central Asiatic vs. the cold-continental Inner Asiatic mountain systems. While the former regions are significantly influenced by the sub-Mediterranean, equinoctial type of precipitation, the latter show an impoverished version of the continental type with scarce summer precipitation maxima (Agakhanjants 1981, Agakhanjants & Breckle 1995. This subdivision was also shown in the area-cladograms of the genus Xenophysa (Varga 1989a(Varga , 2011. In the xeromontane fauna of the "western" Central Asiatic mountains, some butterfly genera (e.g. Parnassius, Karanasa, Paralasa, subgenera of Polyommatus) and also typical genera of Noctuidae (e.g. Euxoa, Dichagyris, Chersotis, Rhyacia, Eugnorisma, Goniographa; and some oligotypic genera, e.g. Hypsophila, Fergana) predominate. The highest diversity of these genera mostly overlap with the core areas of the "saxobiotic" Orthoptera genera, see: Discussion 1.
Contrarily, in the eastern, Inner Asiatic group of the mountains, the typical butterflies belong to the genera Colias, Oeneis and Boloria, which also have xeromontane connections but penetrated into the tundral zonobiome.
The Noctuidae of the Inner Asiatic xeromontane faunal type consist of several "Mongolian-Tibetan" coldcontinental genera, e.g. Trichosilia, subgenera of Xestia Hence, the connections of the xeromontane fauna must be also historically more manifold and partially more ancient than the oreotundral connections of the alpine faunal type, resulted mostly in the Quaternary climatic fluctuations and area dislocations (Varga 1995(Varga , 1996. Thus, the fauna of the subtropical, monsoonic orobiomes (e.g. in Southern China and the Himalayan region) displays a somewhat less differentiated, hypothetically more ancestral character. This regularity is clearly unfolding from the ranges of Poliina genera/subgenera Tricheurois, Haderonia and Metallopolia displaying the species diversity restricted to the Sino-Himalayan area, as opposed to the more derived genera Polia s. str. and Ctenoceratoda (Varga et al. 2019, Fig. 20). We suppose that a bulk of basal groups of the fauna have evolved, jointly with some groups of Angiospermae, in the Eastern Gondwana (see: theory of Axelrod 1960). They could expand northwards after the earliest collision of the Gondwanian plates with south-eastern Asia (e.g. Southern Tibet and parts of south-western China) forming an important source of the Sino-Himalayan core area of biodiversity as was also shown e.g. in the Passerine birds (Fjeldså 2013;Cai et al. 2019).
A high diversity of cold-adapted ancestral species originating from the seasonally humid, monsoonic southeastern Asiatic mountains have been constrained by passing two main filter-corridors (Fig. 21, modified from Varga 1995).
-The "Rhododendron-corridor", being characteristic for several evergreen Angiospermae, e.g. Vaccinium, Empetrum, Rhododendron incl. Ledum, etc., which now compose a major part of the undergrowth of the Siberian (especially southern Siberian mountain) taiga, typical for a number of "taigabirds" (e.g. predominate. In the second group of the mountains the typical butterflies belong to Colias, Oeneis and Boloria, which supposedly have a xeromontane origin but penetrated into the tundral zonobiomes. The typical genera of this second, more "Siberian", group of (originally) xeromontane Noctuidae are e.g. Trichosilia, Lasionycta, Discestra, Anarta and some species groups of the Oncocnemis-Sympistis generic complex. Based on these biogeographical facts, the connections of the Continental-Inner-Asiatic xeromontane fauna should be historically more manifold and ancient than the "typical" oreotundral connections of the alpine faunal type which can be mostly regarded as a consequence of the younger Quaternary climatic fluctuations and area dislocations.   (Dichagyris) squalidior (Staudinger, 1901)