Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv

The 35 anthropophilic species are analyzed. The main tendancies of adaptations tending the transition of bloodsucking mosquito to synanthropic existence are considered. It is shown, that certain part of bloodsucking mosquitoes species inhabiting natural biocoenoses have been already adapted before t...

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1. Verfasser: Kilochytska, N.P.
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Zitieren:Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv / N.P. Kilochytska // Вестник зоологии. — 2012. — Т. 46, № 5. — С. 439–444. — Бібліогр.: 14 назв. — англ.

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spelling irk-123456789-1094652016-11-30T03:02:20Z Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv Kilochytska, N.P. Экология The 35 anthropophilic species are analyzed. The main tendancies of adaptations tending the transition of bloodsucking mosquito to synanthropic existence are considered. It is shown, that certain part of bloodsucking mosquitoes species inhabiting natural biocoenoses have been already adapted before the assimilation into the anthropogenic landscapes. Проанализировано 35 антропофильных видов комаров. Рассмотрены основные направления адаптаций, способствующих переходу кровососущих комаров к синантропному существованию. Показано, что опредёленная часть видов комаров, населяющих природные ценозы, уже преадаптирована к освоению антропогенных ландшафтов. 2012 Article Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv / N.P. Kilochytska // Вестник зоологии. — 2012. — Т. 46, № 5. — С. 439–444. — Бібліогр.: 14 назв. — англ. 0084-5604 DOI 10.2478/v10058-012-0035-8 http://dspace.nbuv.gov.ua/handle/123456789/109465 576.895.771 en Вестник зоологии Інститут зоології ім. І.І. Шмальгаузена НАН України
institution Digital Library of Periodicals of National Academy of Sciences of Ukraine
collection DSpace DC
language English
topic Экология
Экология
spellingShingle Экология
Экология
Kilochytska, N.P.
Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv
Вестник зоологии
description The 35 anthropophilic species are analyzed. The main tendancies of adaptations tending the transition of bloodsucking mosquito to synanthropic existence are considered. It is shown, that certain part of bloodsucking mosquitoes species inhabiting natural biocoenoses have been already adapted before the assimilation into the anthropogenic landscapes.
format Article
author Kilochytska, N.P.
author_facet Kilochytska, N.P.
author_sort Kilochytska, N.P.
title Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv
title_short Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv
title_full Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv
title_fullStr Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv
title_full_unstemmed Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv
title_sort synanthropy of bloodsucking mosquitoes (diptera, culicidae) under conditions of kyiv
publisher Інститут зоології ім. І.І. Шмальгаузена НАН України
publishDate 2012
topic_facet Экология
url http://dspace.nbuv.gov.ua/handle/123456789/109465
citation_txt Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv / N.P. Kilochytska // Вестник зоологии. — 2012. — Т. 46, № 5. — С. 439–444. — Бібліогр.: 14 назв. — англ.
series Вестник зоологии
work_keys_str_mv AT kilochytskanp synanthropyofbloodsuckingmosquitoesdipteraculicidaeunderconditionsofkyiv
first_indexed 2025-07-07T23:08:45Z
last_indexed 2025-07-07T23:08:45Z
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fulltext UDC 576.895.771 SYNANTHROPY OF BLOODSUCKING MOSQUITOES (DIPTERA, CULICIDAE) UNDER CONDITIONS OF KYIV N. P. Kilochytska Taras Shevchenko National University of Kyiv Volodymyrska str., 64, Kyiv, 01601 Ukraine E-mail: kilochytska@ukr.net Received 5 March 2012 Accepted 14 September 2012 Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv. Kilochitskaya N. P. – The 35 anthropophilic species are analyzed. The main tendancies of adaptations tending the transition of bloodsucking mosquito to synanthropic existence are considered. It is shown, that certain part of blood- sucking mosquitoes species inhabiting natural biocoenoses have been already adapted before the assim- ilation into the anthropogenic landscapes. Ke y wo r d s: bloodsucking mosquitoes, Culicidae, synanthropy, Kyiv. Ñèíàíòðîïèÿ êðîâîñîñóùèõ êîìàðîâ (Diptera, Culicidae) â óñëîâèÿõ Êèåâà. Êèëî÷èöêàÿ Í. Ï. – Ïðîàíàëèçèðîâàíî 35 àíòðîïîôèëüíûõ âèäîâ êîìàðîâ. Ðàññìîòðåíû îñíîâíûå íàïðàâëåíèÿ àäàïòàöèé, ñïîñîáñòâóþùèõ ïåðåõîäó êðîâîñîñóùèõ êîìàðîâ ê ñèíàíòðîïíîìó ñóùåñòâîâàíèþ. Ïîêàçàíî, ÷òî îïðåä¸ëåííàÿ ÷àñòü âèäîâ êîìàðîâ, íàñåëÿþùèõ ïðèðîäíûå öåíîçû, óæå ïðåàäàï- òèðîâàíà ê îñâîåíèþ àíòðîïîãåííûõ ëàíäøàôòîâ. Êëþ÷åâûå ñ ëîâ à: êðîâîñîñóùèå êîìàðû, Culicidae, ñèíàíòðîïèÿ, Êèåâ. Introduction The global warming accompanied by radical changes in temperature and humidity could lead to signif- icant implications for species composition and increase the quantity and rate of reproduction of bloodsucking mosquitoes, the carrier agents of vector-borne diseases and, as a consequence, to rise the circulation intensi- ty of such causative agents. At that, the expansion of tropical disease distribution to more moderate latitudes accompanied by change of their distribution features is probable. Under conditions of urbanization of landscapes, the tendency to replacement of bloodsucking mosquitoes coherent evolution by phases of incoherent evolution associated with deep disturbance of cenotical structure is observed. One of the responses to the increase of the urbanization rate is a transition of some part of “wild” species to the facultative or obligatory existence in agrocoenoses. The problem of the distinct factors hierar- chy is especially topical during the evaluation of mosquitoes adaptation in anthropogenous landscapes where a significant rise of part of endophilic and polycyclic species is observed. In this situation, the most significant target is to identify the main tendencies of species diversity changes and mosquitoes ecology in urbanized land- scapes. Kyiv is located in the flood plains of numerous rivers and creeks. The majority of them (Hlybochytsya, Lybid’, Yurkovitsya, Kiyanka, Setoml’, Syrets’, Nyvka, Pochayna, Syrets’ Creek) are currently either complete- ly or partly embedded in sewage system pipes. The floods of Dnieper and Lybid’ rivers only are 17.6% of the city territory. At the same time by 2001 the Kyiv built-up area (landscape and architecture systems) was approx- imately 30% (Stetsyuk et al., 2001). Abundance of waterways along with the hilly terrain is adequate for the formation of different kinds of circulating and inflowing water reservoirs (bays of Dnieper and Desenka rivers, reclamation canals, lakes, ponds, departmental water reservoirs). Only the scheduled bonification by the City Sanitary and Epidemiological Services within the city area exceeds 176 water reservoirs which total water table area is equal to 1735 ha (from 0.9 to 175 ha). It is easy to estimate the hatching of malarial mosquitous larvae if their density in different water reser- voirs varies from 0.8 to 3.0 agents/m2 (according to City Sanitary and Epidemiological Services data). According to monitoring studies 35 anthropophilous species and subspecies of bloodsucking mosquitoes were recorded in Kyiv (Kilochytska, 2008; Kilochytska, Kilochytskiy, 2009). The certain changes in species Vestnik zoologii, 46(5): e-15—e-20, 2012 DOI 10.2478/v10058-012-0035-8 Unauthenticated Download Date | 11/28/16 9:40 PM composition and quantity of mosquitoes over 30 years have been recorded, since the latest consolidated report on mosquitoes (inhabiting Kyiv area) was published (Sheremet, 1978). This paper is based on the results of analysis of bloodsucking mosquito collection for 2003—2010, col- lection materials of the Laboratory of ecology and toxicology of Taras Shevchenko National University of Kyiv and collection of adult female mosquitoes during the same period of time submitted for analysis by City Sanitary and Epidemiological Services (by the “ñollecting from oneself” method). The terms used in this work (coherent evolution, incoherent evolution, anthropophilous species, acom- municative species, communicative species, eurygamy, stenogamy, eusynanthropic, hemisynanthropic, asynan- thropic, endophilic, exophilic, communicative, non-communicative, polyphagous, polystationality) are adopt- ed from Artamonov (2003). Results Synanthropic species are trophically and topically associated with the human and the human biological environment and are the components of anthropocoenosis. ²n order to evaluate potential possibilities of bloodsucking mosquitoes transition to synanthropic existence in city we first of all paid attention to requirements for the habitat necessary for imago and preimaginal phases of development (egg, larva, pupa) of different mosquitoes species. One of the limiting factors is the absence of water reservoirs suitable for develop- ment of mosquito preimaginal phases in the city. Female mosquitos of the genera Culex, Culiseta and Coquillettidia oviposit boat-like clutches of eggs glued together on the water surface. Anopheles females oviposit separately eggs supplied by floats on the water sur- face as well. Females of Aedes and Ochlerotatus oviposit eggs in moist soil separately. Their eggs, in the state of diapause, can survive from several months to several years in moist soil. Consequently, the essential condition for mosquito development from the first four genera is the presence of long-term permanent or semi-permanent water reservoirs, where- as representatives of the two other genera can occur in temporary water reservoirs. In urban areas, the lack of natural water reservoirs (overgrown with higher water plants coastal zone of the lakes, ponds, tanks, ditches and underfloodings) is compensated by artificial ones (pools devoid of vegetation, fire water ponds, barrels, containers, etc.). Larvae of most mosquito genera (with the exception of Coquillettidia) develop in dif- ferent kinds of water reservoirs and quite often they are the significant part of the neuston fauna. The main factors limiting larval lifetime are the duration of water reser- voirs existence and the presence of predators. The limiting factor for the Coquillettidia genus larvae which belong to periphyton is the absence of the higher vascular plants in the permanent water reservoirs. Comparing the participation degree of different stages in the Diptera adaptive evo- lution, we agree with opinion of most dipterists that mosquito adults are more conser- vative than larvae and thus the “driven” phase. The role of imago stipulates the expan- sion and reproduction (nectarophagous, hematophagous, search for substrate and water reservoir, its evaluation and oviposition). Biological markers, which directly or indirectly lead to the success in adapting to an urbanocoenosis, and are found in different species of mosquitoes separately are as fol- lows: the ability to autogenous oviposition; the imago’s need for carbohydrates nutrition (nectarophagous); anthropophyly; the presence of diapause (larval – in representatives of the genera Anopheles, Orthopodomyia, Coquillettidia, and Culiseta, and imaginal – in Anopheles, Culex, and Culiseta); swarming and copulation in a large space – eurygamy, and inability for swarming and copulation in a small space – stenogamy. Based on the assessment of the representation degree of different species in the col- lection (based on Sauhword, 1978) among 35 mosquitoes species attacking human, one super-dominant (An. maculipennis); two dominants (Cx. pipiens and Ae. vexans) and 6 rece- dent species with index of dominance less than 5 % but higher than 1 % were detected (see the table 1). e-16 N. P. Kilochytska Unauthenticated Download Date | 11/28/16 9:40 PM For the classification, of the synanthropic representation degree we apply the sys- tem suggested by Gregor, Povolony (1958) applicable to the higher dipterans, as modi- fied by Mihalyi (1967). According to this system, three groups of synanthropic species are recognized: a) eusynanthropic: endophilic (attributed to residential and utility buil - ding) and exophilic (living outside); b) hemisynanthropic: communicative (directly con- tacting with human), non-communicative (incontacting) and c) asynanthropic. Since the authors applied as a fundamental criterion, an ability of some species of dipterans to infect food and human skin and animals fell without taking into account the changes in species variety of such dipterans under conditions of anthropogenic influence on ecosystems we found useful to amend in the systems following Artamonov’s suggestions (2003). The mosquitoes which lose connection with natural coenosis and are characterized by strong ecological relations with human as a source of food for imago and for larvae indirectly should be referred to the first group (eusynanthropus). Among 35 anthropophilic species only Culex pipiens Linnaeus (f. molestus), which populations distinctly timed to urbanocoenosis could be referred to eusynanthropus with some approximations. This species is characterized by the following: the ability to autogenous oviposition, the absence of diapauses and the need for nectarophagy, stenogamia and (especially in the cold season) is the distinctly expressed endophily. Such biological features allow Cx. pipiens f. molestus to develop year-round in the flooded basement and to attack a human for the bloodsuck- ing, inside the premises. Formation of Cx. pipiens f. molestus synanthropy was originally recorded in 1775 in Cairo and Alexandria (Egypt). The original description of this taxon indicated that it was characterized by the high aggression to human and reached particularly high numbers in urban areas. For the first time Cx. pipiens f. molestus was observed in Europe in 1921 (London) and then it spread throughout Western Europe by 1950. In Ukraine, the mosquito was recorded in 1926 (Dnipropetrovsk), and by 1990 it became ubiquitous in more than 300 cities of the former USSR in all landscape and climatic zones (Vinogradova, 2004). During this period changes of biotopes inhabited by the immature stages of Cx. pip- iens f. molestus occurred. In temperate climatic regions, the breeding sites of its larvae in urban areas were the coastal shallows, pits, ditches, bogs, fire barrels. Then they were almost entirely replaced by the filtration fields. However, in 1940—50s Cx. pipiens f. moles- tus hatching in the basements of Leningrad and Moscow was reported for the first time, followed by accelerated growth of its population and the range expansion (Bogdanova et al., 1992). In 1964 the WHO International Workshop established that the biological progress and the spreading of Cx. pipiens f. molestus were closely linked with the urbanization pro- cesses (in particular, with the construction of buildings with district heat supply), which is a general trend in the world (Seminar..., 1965). Currently, the process of progressive urbanization of the other anthropofilous species such as Culiseta longiareolata Macquart, e-17Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv Ta b l e 1. Species of mosquito, which dominate in attacking on human in Kyiv Ò à á ëèö à 1. Âèäû êîìàðîâ, äîìèíèðóþùèå ïðè íàïàäåíèè íà ÷åëîâåêà â ã. Êèåâ Species Index of dominance, % range values, 1992—2008 average Anopheles maculipennis Meigen (complex) 2.3—66.0 35.6 Culex pipiens Linnaeus (complex) 2.5—50.9 18.8 Aedes vexans (Meigen) (complex) 1.8—41.6 14.6 Ochlerotatus sticticus Meigen 1.8—21.6 9.0 Aedes cinereus Meigen 1.4—6.4 3.6 Culiseta annulata (Schrank) 0.7—5.9 2.7 Ochlerotatus cataphylla Dyar 0.1—6.9 2.7 Coquillettidia richiardii (Ficalbi) 0.1—4.6 2.4 Anopheles ñlaviger (Meigen) 0.1—2.8 1.3 Ochlerotatus caspius (Pallas) (complex) 0.1—3.8 1.1 Unauthenticated Download Date | 11/28/16 9:40 PM overwintering as the egg stage, the larvae and fertilized females are also observed. This species occurred initially in a spring lake and then in artificial ponds, barrels of water, pools, etc.; they were recorded by E.the flooded basements in Javan (Tajikistan). The second group (hemisynanthropes) consists of mosquitoes of greater ecological flexibility: their population is equally capable of occupying both urbanized and natural habitats. By food specialization along with Cx. pipiens f. molestus “pure” anthropophilic mosquito is Coq. richiardii. In Kyiv environment its larvae live in a few reservoirs over- grown with higher vascular vegetation, where they overwinter as a diapausing larval stage. A similar pattern have been recorded earlier in Moscow and Leningrad (Sheremet, 1978). Coq. richiardii with good reason could be categorized as communication exophilic hemisynanthrope, which forms a connection with human primary on the trophic basis (hematophagous), and the status of their populations directly depends on the availabil- ity of breeding sites for the larvae. An. maculipennis is a casually anthropophilic bloodsucker, which prefers to attack the cattle out of city (Gutsevich et al., 1970), while by the intensity of attacking humans in Kyiv it takes the superdominant position. In the city, its larvae can develop in a var- ious permanent ponds. The tendency of adults is to hide for the day and survive the win- ter (imaginal diapause) in various outbuildings (unheated rooms, barns, cellars, basements, tunnels and even subways) and lack of alternative feed source in urbanocoenoces con- tributes to more aggressive attack the human. In fact, An. maculipennis is a typical com- municative endophilic hemisynanthrope in Kyiv. In natural coenoses, polycyclic Ae. vexans as well as An. maculipennis is an option- al anthropophilious bloodsucker. Its larvae are produced in a variety of open preferably floodplain ponds. Perhaps it is the clue for explaining the reasons of high numbers of this species in Kyiv. Ae. vexans of Kyiv population should be attributed to the communica- tive exophilic hemisynanthrope. Long-term population dynamics (and the attack activity) of phenological summer polyseasonal species (O. sticticus, O. caspius, Ae. cinereus, Cs. annulata and An. claviger) correlates directly with the presence and seasonal heavy rains. Larvae produced by early spring species O. cataphylla are among the first to appear in the melt water, mostly in open temporary ponds, and thus they have time (as opposed to other spring species) to complete preimaginal development before breeding sites dry up. Significant anthro- pophyly of hematophagous adults and the ability of immature stages to develop in urbanized landscape attribute O. sticticus, O. caspius, O. cataphylla, Ae. cinereus, Cs. annu- lata and An. claviger to the category of communicative exophilic hemisynanthropes. The last 25 species of mosquitoes which intensity of which to attack a human does not exceed 1.0% in the whole complex of bloodsuckers were referred to a group of option- al communicative exophilic hemisynanthropes. Optional anthropophily of these species is explained by the fact that in the environment Ochlerotatus excrucians Walker and Aedes cinereus Meigen mainly attack domestic animals; Cs. annulata, Cx. pipiens f. pipiens Linnaeus and O. caspius are ornitofilous species; Culex territans Walker attacks more fre- quently reptiles and amphibians (Gutsevich et al., 1970). Synanthropic fauna is formed by means of the penetration and retention of a part of populations occupying adjacent natural ecosystems in the anthropocoenosis. Direction and speed of this process are poorly predictable especially when it is associated with mosquitoes of high flight activity and rapid adaptation. ²nitially, 50—60 years ago the num- ber of species, which inhabited urban habitats was less than the number of species liv- ing in their neighborhoods. Currently, a clear tendency towards increasing the species diver- sity of mosquitoes in urban areas is observed regardless of the climatic zone, the size and age of the city. Thus, 32 species of mosquitoes were registered in the vicinity of Moscow in 1962—1969, while 21 species were recorded within its boundaries. In 1983—1985 the number of mosquitoes species reached 26 in the city. 30 species of mosquitoes were record- e-18 N. P. Kilochytska Unauthenticated Download Date | 11/28/16 9:40 PM ed in Leningrad and its environs in 1928—1958 while only five of them in the city. Observations of 1983—1987 showed the presence of 22 mosquito species in Leningrad (Bogdanova et al., 1992). In the city of Kyiv considering the minor quantitative changes, one should note cer- tain qualitative changes in mosquito species diversity in 1978—2008. Two species of mosquitoes of the genus Culiseta (Cs. glaphyroptera Schiner and Cs. morsitans Theobald) and four representatives of the genus Ochlerotatus were added to the list of species. The elimination of Ochlerotatus pulchritarsis Rondani bred in hollows and six other species of this genus (O. annulipes Meigen, O. behningi Martelli, O. intrudens Dyar, O. cataphyl- la Dyar and O. leucomelas Meigen), whose larvae are propagated in grass-covered and enriched with leaf litter temporary forest ponds and their edges under natural conditions could be explained by the progressive urbanization (ordering/regulation) of the urban area accompanied by the disappearance of corresponding types of reservoirs (Kilochytska, Kilochytskiy, 2009). V. M. Beklemishev (1952) observed earlier this tendency, which is the human development of the territory directly worsens the environment for the exis- tence of mosquitoes of the genera Aedes and Ochlerotatus deteriorate. Observation of Ochlerotatus euedes Howard, Dyar et Knab, O. cyprius Ludlow, O. d³an- taeus Howard, Dyar et Knab and O. pullatus Coquillett in Kyiv is obviously related to the presence of such artificial ponds without vegetation such as pits, trenches, potholes or ruts, etc. in the areas of capital construction during the spring season. Understanding the process of synanthropic fauna formation is facilitated by the use of the “preadaptation” concept (the process of rising the character in species, which orig- inally does not have any adaptive value and becomes adaptive only after changing the living conditions of animals and, if the need arises, it is used afterwards in future gener- ations). In particular, E. Mayr (1974) considered the species were preadapted if it was able to occupy a new habitat. The analysis of larval habitats in Kyiv shows that most of the mosquito species domes- ticate the ponds, which are not different in their characteristics from those natural con- ditions. Eight species develop only in ponds and lakes (in the coastal heavily overgrown area), the most numerous of them, An. maculipennis, Cx. pipiens f. pipiens and Ae. cinereus, are propagated. Fourteen species are the most abundant in floodplains, flooding of rivers and streams such as Ae. vexans, An. maculipennis, O. cantans, O. excrucians, O. caspius, O. flavescens and O. behningi O. cataphylla, O. diantaeus and Cx. thailary are propagat- ed in roadside ditches. Cx. pipiens f. pipiens and An. maculipennis predominate in pools, tanks and barrels while Cx. pipiens f. molestus predominate in basements. Larvae of Anopheles plumbeus Stephens, Aedes geniculatus Olivier and O. pulchritarsis are propagated in the hollows. A regular appearance of saline ponds (as a result of melting snow sprinkled with mix- ture of sand and NaCl or KCl) in the city leads there to the primary development of a number of such historically preadapted to the existence in water of different salinity such species as An. maculipennis, An. claviger, O. caspius caspius, O. detritus, O. caspius dor- salis, O. excrucians, O. leucomelas, Ae. vexans, Ae. cinereus and Cx. theileri. Perhaps the mineral pollution of water results in preferential conditions for the development of some acommunicative species of mosquitoes in the city. Summarizing the results, it can be stated that the main complex of preadaptations of bloodsucking mosquitoes enabling their inflow of species into the synanthropic fauna from adjacent ecosystems includes the following factors. – Polyphagous adults. Such species could most likely shift for feeding on humans and commensal animals since there are no vitally important imaginal trophic differences for these hematophagous. – Polystationality. It is common for almost all representatives of all genera (except Coquillettidia), which manifested in the absence of strict determinism in the choice of habitats (reservoirs) for preimaginal development. e-19Synanthropy of Bloodsucking Mosquitoes (Diptera, Culicidae) under Conditions of Kyiv Unauthenticated Download Date | 11/28/16 9:40 PM – Tolerance to the fluctuations of climatic factors. Increasing the number of species, which are trophically and topically related to the human, does not depend on the cli- matic zone, the size and age of the city. – There are neither predators or parasites of anthropogenic origin in ponds. – Significant expansion of the distributional radius of adults. Such a benefit espe- cially effectively appears in Kyiv during the migration of mosquitoes in the valleys of numer- ous rivers (especially the Dnieper), along the railways and roads. The author thanks O. M. Pogorelova (chief entomologist of Kyiv Sanitary Epidemiological Service) for the granted materials. Artamonov S. D. The formation of adaptation to synanthropic in the example dipterous families Sarcophagidae and Calliphoridae (Insecta, Diptera). Kurentsova – 2003. – 14. – P. 14—20. – Russian : Àðòàìîíîâ Ñ. Ä. Ôîðìèðîâàíèå àäàïòàöèé ê ñèíàíòðîïèçìó íà ïðèìåðå äâóêðûëûõ ñåìåéñòâ Sarcophagidae è Calliphoridae (Insecta, Diptera). Beklemishev V. N. The fight against midges. Problems of anti-malarial organization in the new stage // Med. parasitil. i parasitarn. bol. – 1952. – 21, N 6. – P. 487—492. – Russian : Áîðüáà ñ ãíóñîì. Î çàäà- ÷àõ ïðîòèâîìàëÿðèéíîé îðãàíèçàöèè íà íîâîì ýòàïå. Bogdanova E. N., Smyslova Ò. Î., Ostanin G. I. et al. Dynamics the development of urbanization coenosis mosquitoes // Blood-sucking and zoophilous dipterans (Insecta, Diptera) : Materialy V Wsesouznogo dipterologicheskoho symposiuma (14—17.08.1990, Novosibirsk). – Novosibirsk, 1992. – P. 22—25. – Russian : Áîãäàíîâà Å. Í., Ñìûñëîâà Ò. Î., Îñòàíèí Ã. È. è äð. Äèíàìèêà îñâîåíèÿ êîìàðàìè óðáà- íîöåíîçîâ. Gutsevich À. V., Monchadskiy À. S., Stackelberg À. À. Mosquitoes (Family Culicidae) // Insecta Diptera. – Leningrad : Nauka, 1970. – 384 p. – (Fauna of the USSR. New ser. 100; Vol. 3. Fasc. 4). – Russian : Ãóöåâè÷ À. Â., Ìîí÷àäñêèé À. Ñ., Øòàêåëüáåðã À. À. Êîìàðû (Ñåìåéñòâî Culicidae). Kilochytska N. P. Short key of blood-sucking mosquitoes fauna of Ukraine. – Êyiv : Geoprint, 2008. – 90 p. – Ukrainian : ʳëî÷èöüêà Í. Ï. Êîðîòêèé âèçíà÷íèê êðîâîñèñíèõ êîìàð³â ôàóíè Óêðà¿íè. Kilochytska N. P., Kilochytskiy P. Y. 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