In such a variety of environmental conditions may exist a variety of mosquito species. The total number of species is 53. Among these, six members of the genus Anopheles, six — sort of Culiseta, 31 belong to the genus Aedes. eight — to the genus Culex and one — to leave Uranotaenia and Mansonia.
Analysis of the species composition and ecology of mosquito fauna of Kazakhstan allowed include composing it to the four types of eco-faunistic complexes: the forest, steppe, steppe and desert. The basis of allocation of the principles of geographic homogeneity, uniformity of distribution and ecological similarities of species.
In these groups sufficiently characterize the concept of "fauna complex" in the definition of G. Nicholas (1947), although it does not reflect the nature of the considered groups. This term the author means a group of related community to its geographical origin to a particular area, to the conditions which the adapted species that make up the complex. We add to this definition are such criteria as environmental similarities. To mosquitoes not quite apply the principle of self-titled zoogeographical analysis of association of a region, so successfully applied by VV Shevchenko (1961) in a study of horseflies. The reason is probably the fact that the habitats of mosquitoes are generally too numerous to be specific to a small region, since the divergence of their slower and species endemism is weak. At present, these interconnecting applies to broad categories such as geographic region, as is done Sazonova O. (1962).
To forest faunistic complex include cold-, monocyclic species Aedes and polycyclic, oligotermnye views Culiseta. Almost all members of this complex extended to the north of the USSR. They pinch out to the south island on the relic forests and floodplains to the steppe and semi-desert areas, which is entirely characteristic of Kazakhstan. Its territory forest industry representatives are quite numerous in the steppe zone, covering the northern areas of the country. As we move to the south spread their acquired mosaicism. Limited areas of their natural habitat are found in the high forests of Central Kazakhstan and the floodplains of large and small rivers in the western part. Such zoning is marked up to 48-47 °. sh. Their adaptation to these conditions is expressed in earlier than the forest-steppe zone of the development, a brief period of activity and strong affinity to typical habitats.
On the slopes of the Tien Shan individual members of this complex (Ae. leucomelas, Ae. Cataphylla) reach the South Kazakhstan and neighboring regions of Kyrgyzstan (43-42 ° c. lat.). In the mountainous parts of the south and south-east of Kazakhstan, depending on the altitude they are either dominant or subdominant. Dominate in the belt of forests and subalpine meadows and less numerous in the area of mountain steppes. Sometimes in the cold mountain river floodplains penetrate the semi-desert, and even in a desert area (near Ili), that is observed in the vertical distribution of the same pattern as in the plains of Kazakhstan. The most cold-representatives of this complex (Ae. pullatus, Ae. Intrudens) rise to 3000 m n. a. m, which corresponds to the zone of cold mountain deserts (syrty). The spread in the mountains of the genus Culiseta limited area of mountain steppes and lower open areas of the forest zone.
It's not clear ecological status of three recently described species — Ae. simanini (Gutsevich, 1966) Ae. kasachstanicus (Gutsevich, 1962) Ae. montchadskyi (Dubitsky, 1968). Morphological and ecological features first should probably refer to the forest complex, and the second and third — to the forest-steppe. Currently solved this question is not possible, because the ranges of these species are unknown beyond the eastern limits of Kazakhstan.
Representatives of the steppe faunal assemblage psychrophilic less than the previous species. Wider range of plasticity that allows them to equally exist in the forest and steppe zones. Some of them (Ae. excrucians, Ae. Flavescens) penetrate to the north to 68-69 ° c. sh. (Rumsh, 1948). Members of this complex along with some steppe species are dominant in the corresponding zonal areas of North and East Kazakhstan. Sometimes there is a development of the second (letneg-e) generation of a general monocyclic species.
Individual representatives observed further south than forest species. For example, Ae. flavescens occurs throughout the length of the lower Syr Darya and Ili (43-44 ° c. lat.). Some types of (Ae. cyprius, Ae. Subdiversus) some do not reach this milestone, and Ae. behningi. Ae. excrucians and Ae. cantans prevalent in Kazakhstan to about 48 °. sh. To the south of all comes Ae. claviger, but it is found only in the cool mountain and foothill habitats.
In the mountainous and foothill parts of Kazakhstan kinds of complex distributed unevenly. In the foothills of the Altai and the Zaisan basin numerous Ae. excrucians, Ae. cantans, Ae. cyprius and Ae. flavescens, and in the Tian Shan Mountains — only Ae. flavescens, which is found here in the mountain steppe zone and the lower portions of the forest belt. To the south it comes to northern Kyrgyzstan (Konurbaev, 1965), but does not reach Tajikistan (Keshishyan, 1941).
Species included in the steppe complex, yet more plastic than forest and forest-steppe, and therefore, as a rule, have a large spatial extension areas. Suffice to say, this group includes Ae. vexans and Ae. cinereus, ranges that extend from the desert to the south of the taiga zone, and Cx. pipiens through its commensal comes farther north (Rumsh, 1948). Establish zonal affiliation of these species can only be on the steppe habitat preferred by extrazonal and wedging in the desert and steppe zones.
If you compare the representatives of the faunal assemblage from the previous ones, it is striking increase in the number of Culex and Anopheles. Group Aedes presented evribiontnymi polycyclic species, depending on climatic conditions that might give one or more generations. In the southern parts of their ranges of development and hatching of these species are confined to the well-warmed waters with the surrounding dense vegetation, and in the north, they penetrate only to the shaded and heavily overgrown ponds. Extrazonal habitats in the first case are the floodplains of the southern rivers, and in the second — open space with well warmed temporary ponds.
In mountainous areas there are all kinds of this group, except An. hyrcanus, M. richiardii, Cs. annulata. For them, probably not warmed mountain reservoirs. All other species inhabit the zone of mountain steppes and downstream areas. Ae. with. dorsalis occurs in areas of sparse forest belt of mountainous areas, and sometimes in a well-heated areas of sub-alpine meadows.
Representatives of the steppe faunal assemblage common in Kazakhstan is quite uneven. The five most evriplas-particle species (Sh. pipiens, Cs. Annulata, M. richiardii, Ae. Vexans, Ae. Cinereus) meet not only all over Kazakhstan, but also beyond. All other Culex and An. hyrcanus, covering the southern areas of the country, reach 48-50 ° c. sh. Areas of Ae. with. dorsalis and An. so messeae occupy the northern part of Kazakhstan, reaching 46-48 ° c. sh., under favorable conditions, pinch out for this milestone. To the south they are replaced by similar forms (Ae. with. Caspius, An. Tons sacharovi).
The bulk of the steppe complex in either direction extends from Kazakhstan. Penetration An. so messeae north to the southern boundary of the taiga, most authors (Beklemishev Zhelohovtsev, 1945 Rumsh, 1948 Monchadsky, 1951) explain the large commensal this species and the concurrent development of unshaded, well-warmed waters and wintering sites — to various rooms. Many of the more extreme points of its discovery is the result of delivery of transport and promotion for the man during the clearing of the taiga (Rumsh, 1948).
In the desert fauna complex includes species, the main part of the area which lies beyond the southern limits of Kazakhstan. As representatives of the previous set, all polycyclic, but are highly suhoustoychivostyu. This allows them to live in dry, arid areas, is what populates them zone. The group in question is not uniform over the thermophilic: there are as strict politermofily (An. pulcherrimus, An. Superpictus. An. Tons sacharovi, Ae. Pulchritarsis, Cx. Pusillus), and heuristic-tslastichnye representatives of this category (Sh. modestus, Cs. Longiareolata, Ae. Since. Caspius, Ur. Unguiculata). The same complex is a group of three types: Cx. theileri.Cx. hortensis. An. algeriensis, tending to develop in the cool waters of the mountain and foothill habitats. This discrepancy between the desire for cool habitats and distribution in dry hot regions VN Beklemishev Zhelohovtsev and AN (1945) for example An. algeriensis explain the need for a long warm season, which makes it possible to carry out in cool habitats two generations needed to preserve the species.
All representatives of the desert fauna complex distributed south of 48 ° c. sh. Only two types of (Cs. longiareolata and Cx. modestus) than desert steppe zone and cover found in open areas of the forest-steppe. In contrast, the areas of the two most thermophilic species — An. pulcherrimus and An. superpictus — Go to only in the southern regions of Kazakhstan, to about the Karatau ridge. In the mountains, no view No high rises. Ae. with. caspius and Cx. modestus marked in the most low-lying, well-warmed ground belt mountain steppes. Ae. detritus, Cx. pusillus and Ur. unguiculata do not occur in the mountains: the first two, possibly due to the lack of specific mountains solo novodnyh water, and a third on the territory of Kazakhstan, as already mentioned, has only been in floodplains.
Analyzing areas of all the above groups, we can see that the spread on the plains of Kazakhstan the majority of the representatives of the forest faunal assemblage does not go south of 50 ° N lat. sh., and representatives of forest-steppe — 47-48 ° c. sh. However, by 47-48 ° c. sh. approaching the northern border areas of the desert, and the majority does not get steppe north of 50 ° N lat. sh. This indicates the presence in Kazakhstan two climatic boundaries: about 48 ° and about 50 °. sh.
The observed phenomenon is not accidental. Somewhere around 50 °. sh. is the major axis of the continent, which is likely to be the border of the two major climatic regions: north and south. The area lying south of the axis, differ dryness and more completely continental climate than the area to the north of it. The main importance is probably the amount of summer temperatures, which is north of 50 "s. Lat. Decreases sharply, enabling approach to security is oligotermofilam north and to the south — mezotermofilam.
Second line extending from about 47-48 °. sh., characterized by the abrupt transition from high summer of positive to negative balances winter temperatures. Warm and sunny summer creates favorable conditions for the spread of many heat-loving animals, including desert species of mosquitoes, while low winter temperatures limit the ability of their experiences in the winter. However, the last factor can probably at biotopes extrazonal penetrate to this latitude to some forest-steppe species.
Environmental prerequisite for extrazonal available in Kazakhstan steppe species complex is a large amount of solar radiation, which, even in the northern regions of the country in the summer close to that of peculiar to 15 °. sh. Accordingly, in the desert regions of the country dedicated to the development of the cool shaded habitats, and in the forest-steppe zone, on the contrary, to open, well-warmed. This passage reflects the basic principle of the principle of changing habitats (Bay Bien Co., 1959).
From all this it follows that the basic patterns of distribution and ecology of mosquitoes in Kazakhstan is largely consistent landscape-climatic characteristics of various natural zones of the country. A prerequisite is the history of the formation of modern fauna of mosquitoes, which is quite clearly seen the presence of Euro-Siberian, Mediterranean, South Asian and Palearchaearctic representatives.
The primary mosquito fauna of Kazakhstan is probably connected with the tropical and subtropical members of the genus Culex, There was, according to Edwards (Edwards, 1932), already in the Jurassic, and formed, probably in the Triassic period. This is confirmed by local paleontological data. At the beginning of the Tertiary period in large parts of Kazakhstan arid climate is attributable VM Sinica-tion (1962) and Lavrenko (1962) to the type of wet savannas. As in the present, with the arid, warm conditions at the time were probably related species of the genera Culex and Anopheles, with the center of their speciation tropical zone of the Old World. Photographed fossil instance mosquito is also likely to belong to the genus Culex. This can be judged by the convexity of the mesonotum, a relatively small length of the legs and the presence of bands on the abdomen, which distinguishes it from the genus Anopheles.
Spread them throughout the territory of modern Kazakhstan contributed notes in Paleogene advance north to the boreal elements of tropical landscape. By this time, OL Kryzhanovsky (1965) refers to the migration of the Kazakh territory of some genera of grasshoppers, beetles and jewel having family ties with the deserts of the southern hemisphere. Among the earliest members Culicidae should, apparently, include species that have crept into the present, far to the north (Sh. pipiens, Cx. Modestus. An. Tons messeae. An. Claviger). Promotion drugihvidovbylo probably limited matured during the Oligocene general cooling of climate and the retreat of tropical flora and fauna in the area of the southern latitudes. Substitution in the Neogene subtropical savannas of boreal analog — step further limit the optimum life of the thermophilic forms. The existence of such conditions could accommodate a species, are coming to the north and largely adapted to the colder climate. Thermophilic representatives of these groups (An. superpictus. An. Algeriensis. An. Pulcherrimus, Cx. Pusillus) and were within the modern desert zone. The probability of such a population of North American temperate zone species Culex suggests and Ross (Roes, 1964).
By the middle of the Tertiary period can apparently be attributed to the emergence of Kazakhstan boreal species of mosquitoes belonging to the genera Aedes (Subgenus Ochlerotatus) and Culiseta. Their extrapolation occurs in conjunction with the change of savannah landscape of broad-leaved forests. Most of these species — forest dwellers, and the area of their repeated distribution of deciduous flora. Then dominant wet gentle climate resettle many moderate forms of vegetation.
Then began the dry continental climate halt the south boreal flora and due to continued cold weather disappeared tropical and subtropical plants. Desertification is progressing in the Neogene and Quaternary (Lavrenko, 1962). Forest thinning and break up into separate arrays, separated prairie associations. Together with forest vegetation pushed boreal species of mosquitoes. Some of their representatives are preserved in extrazonal forest habitats for upland and floodplain. Hence, and notes now atypical blowout fauna of these northern species to southern Kazakhstan, where they are no more than a boreal relicts.
Along with the advance of deserts went further spread of Representatives Culex and Anopheles. In this case, the present fauna of Kazakhstan was in a somewhat better position, since not all members of these genera have moved so far to the north. Central Asian fauna Culex and Anopheles more abundant.
Thus, the mosquito fauna of Kazakhstan was formed by the northern and southern components. Ways of penetration does not entirely correspond to a pole, there is a zonal migration. From the north, as already indicated, the representatives moved forest faunal assemblage. Therefore, almost all species of mosquito found North of the USSR in the North and East Kazakhstan.
In the same direction, but with a bias to zone migration was probably the current population and the representatives of Kazakhstan steppe complex. This is supported by their significant thinning in the forest, and in the arid zone and the absence of some of them (Ae. behningi, Ae. Beklemischevi) in the eastern part of the forest-steppe zone.
Foci for the spread and the very marked the epicenter can be concluded that the speciation of three representatives of the two sets (Ae. subdiversus, Ae. Kasachstanicus, Ae., Montchadskyi) was probably in Kazakhstan or in adjacent areas Dzungaria. This is indicated by the relatively high number of Ae. subdiversus Turgay dining in the country, and Ae. kasachstanicus and Ae. montchadskyi — In the pool, and notes or expansion to neighboring territories. Single copies Ae. subdiversus found in Saratov, Orenburg, Guriev and Karaganda regions. Along floodplains Urals Irgiz Turgay he probably entered the semi-desert zone, and on the floodplain of the Tobol and its tributaries — in Western Siberia.
Obvious and complex relationship with the fauna of the forest-steppe zone of the RSFSR respective bordering Kazakhstan. All species of this complex are found in the vicinity of Saratov and Orenburg (Martini, 1926, 1928), located at the southern boundary of the forest steppe.
More dismembered way to enter the territory Kazakhstan representatives of the steppe and desert fauna complexes: the western, southern and eastern. From the west, apparently penetrated Ae. detritus, common to the Mediterranean. The main part of the way Culex, Although held in the south-west of the Ethiopian region, but it was through the border regions of Central Asia. There we meet not come down to Kazakhstan Cx. tritaeniorhynekus. Went through the same distribution, and some Mediterranean species Anopheles (An. superpictus. An. Algeriensis). Oriental species moved in two ways: the east, to the Pamir-Tien Shan mountain system, and the West. In Kazakhstan, they are closed, as judged by the occurrence in the south-east of the two geographical races Ae. vexans (Ae. v. vexans, Ae. v. nipponii). Moreover, a mixture of these forms is observed only in the lower bordering China, areas.
Based on all the above we can conclude that the fauna of Kazakhstan, geographically combining a variety of landscape and climate, is a composite character. This also explains its relatively rich diversity of species. The main ways in which its formation occurred, served south — for Culex and Anopheles and north-west — for Aedes and Culiseta. The process of becoming fauna is probably not over. This is indicated by the discovery of some new species (Ae. kasachstanicus, Ae. Montchadskyi, Ae. Stramineus) and expansion. In the future, we should expect some penetration in southern Kazakhstan (Sh. univittatus) and mountain (An. lindesayi. An. Marteri) species already surviving Tajikistan (Keshishyan, 1941).
In the future, the species composition of mosquitoes Kazakhstan can have minor recharged both by identifying previously encountered this species, and as a result of the process of formation.