CZU: 595.78:591.9

ZOOGEOGRAPHIC FEATURES OF BLACK FAUNA (DIPTERA; SIMILIIDAE) OF PALAEARTIC

1VASILEVICH F.I., 2KAPLICH V.M., 3SUKHOMLIM YE.B

1 Moscow State Academy of Veterinary Medicine and Biotechnology named of K.I.Skrabin 2 Belorussian State Technological University 3 Eastern European National University named after Lecya Ukrainka

Abstract. The representatitives of the sub-families Prosimuliinae and Similiinae which are widespread in Palaearctic have holarctic, holarctic-oriental, palaearctic, palaearctic-oriental, oriental- easternpalearctic and palaearctic-oriental-afrotropical-Australian prevalence. The core of Palaearctic simuliidofauna consists of the palaearctic habitats. Key words: Black , Fauna, Palaearctic, Simuliidae, Zoogeography.

INTRODUCTION According to the reports of P.H. Adler R.W. Crosskey (2012) there are 725 species of Simuliidae from 38 genera, which comprises 33-38 per cent species of the world fauna black flies. The composition of the genera in the zoogeographic area is much more diverse (38 genera from 81 world faunas or 47%) than in the other ones.

MATERIALS AND METHODS The materials from the Zoological Research Institute (Russian Academy of Sciences, S.- Petersburg), GNPO “NPTS NAN of Belorussia on Bioresources” (Minsk), Veterinary Institute in the town of Schwerin (Bezirksinstitut fur Veterinarwesen Schwerin) in 1991, later re-named the State Veterinary Agency (Landesamt MV LVLUA), Donetsk National University named after Lecya Ukrainka (Lutsk). The habitat classification is given after K.B. Gorodkov with additions.

RESULTS AND DISCUTIONS The subfamily Prosimuliinae is represented by two tribes: Gymnopadini Rubtsov, 1954 and Prosimuliini Enderlein, 1921. The black flies of this sub-family inhabit Holarctic (Yankovsky A.V., 2002), the largest part of neotropical and afrotropical kingdoms (Adler P.H. et all., 2004). But they are not encountered in the Oriental or Australian ones (Crosskey R.W., Chichester U.K., 1990). The tribe Gymnopaidini comprises three genera: Gymnopais Stone, 1949 (7 species), Twinnia Stone et Jamnback, 1955 (7 species), Levitinia Chubareva et Petrova, 1981 (3 species). The black flies of all three genera inhabit the disjunctive mountain areas (Gorodkov K.B., 1983): Gymnopais-Siberian-amphipacific (Rubtsov I.A., 1964); Twinnia-Holarctic (Rubtsov I.A., 1959); Levitinia-Mediterranian-Central Asiatic (Chubareva L.A., Petrova N.A., 1981); i.e. they are widespread, but their spreading is mosaic. To A.V. Yankovsky’s opinion (2002) the modern disjunctive areas of tribe species are the remnants of the entire past areas that arose on the borders of the former glaciations or they are the result of colonizations in the Pleistocene period of mountain and the circumpolar biotopes during glacier recession. The tribe Gymnopaidini includes the fossil species Kovalevimyia lacrimosa Kalugina, 1991, dated from Malm. It was well preserved on the scar found in the chalk deposits of the South of Siberia (Chitinskaya oblast). There are no Kovalevimyia species in the recent fauna, but their scars were found within up- to-date spreading Gymnopaidini tribe representatives.

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The tribe Prosimuliini comprises three genera: Prosimulinium Roubad, 1907 (37 species), Helodon Enderlein, 1921 (20 species) (Yankovsky A.V., 2002), Urosimulium Contini, 1963 (3 species) (Adler P.H. et all., 2004). The black flies of this tribe inhabit Holarctic and they are not encountered in other zoogeographical areas. The Holarctic habitat is characteristic of the representatives of the genera Prosimulium and Helodon. The exception is the species of Urosimulium genus which are only spread within the limits of Palaearctic and have a local Mediterranian habitat (inhabit Italy, Spain and the Balearic Islands (Adler P.H. et all., 2004)). To Prosimuliini tribe belongs the fossil species Simulimima grandis, Kalugina, 1985, which is encountered in the deposits of lias/middle Jurassic, and also Baisomia incognita Kalugina, 1991 and Prosimulium brevirostris spread in the lower cretaceous period, but Mesasimulium lahaigounense – in the upper cretaceous period. The black flies are well preserved in the scars found in the South of Siberia (Buryatia, Tuva). There are no representatives of these genera in the Siberian recent fauna, but their scars were found within the limits of up-to-date spreading Prosimuliini tribe. The Simuliinae subfamily includes five tribes: Stegopternini Enderlein, 1930, Ectemniini Enderlein, 1930, Nevermanniini Enderlein, 1921, Wilhelmiini Baranov, 1926 and Newman, 1834. The subfamily species are spread almost in all zoogeographical areas (Yankovsky A.V., 2002; Adler P.H., 2012). To the tribe Stegopternini belong two genera: Stegopterna Enderlein, 1930 (11 species) and Greniera Doby et David, 1959 (7 species). The black flies of this tribe inhabit Holarctic, the northern part of neotropic kingdom (Yankovsky A.V., 2002). The representatives of Stegopterna genus have a holarctic habitat and they are encountered in North America, in the North of Europe and Asia and in Japan (Adler P.H. et all., 2004). The boreal amphiatlantic habitat spreading down to North Africa (Algeria, Morocco) is characteristic of the genus Greniera. The genus Greniera includes three fossil species: G. affinis Meunier, 1904, G. importuna Meunier, 1904, G. pulchella Meunier, 1904 (oligocene) observed in the Baltic amber. They confirm the boreal amphiatlantic habitat of recent species. The fossil species Gydarina karabonica Kalugina, 1991 which is close to the genus Greniera was described by the scar found in Chitinskaya oblast and it must fall under the category of the tribe Stegopternini. The location of finding the scar is within the limits of holarctic spreading the tribe, but extends it to the south within the limits of Siberia (Kalugina N.S., 1991). There are three genera in the tribe Ectemniini. They are Cnephia Enderlein, 1921 (6 species), Metacnephia Crosskey, 1969 (53 species) and Sulcicnephia Rubtsov, 1971 (21 species) (Adler P.H. et all., 2004). The representatives of the tribe inhabit Holarctic, but some genera are encountered in Neotropic (Yankovsky A.V., 2002). Holarctic type of the habitat is characteristic of the black flies of the genus Metacnephia and of those of Cnephia is flat northern – holarctic one (Yankovsky A.V., 2002). The genus Sulcicnephia representatives are Palaearctic endemics and they have a disjunctive mountain central-eastern-palaearctic habitat. They inhabit Central Asia, the south of Siberia, Pamir, Mongolia, Kamchatka (Rubtsov I.A., 1955; 1982). Researchers revealed different reasons for disjunctive habitats of the tribe species. Mosaic spreading of Cnephia is connected with pollution of long plain rivers with industrial sewage in Central and Northern Europe, North America (Rubtsov I.A., 1984), but Sulcicnephia black flies inhabit large mountain rivers, which results in their geographical isolation (Rubtsov I.A., 1956). Two fossil species belong to the tribe Ectemniini: Ectemnia Cerberus, Enderlein, 1921 and Ec. lithuanica Yankovsky et Bernotiene, 2005. Both finds were described by the Baltic amber and dated from the oligocene. It should be noted that modern representatives of the genus Ectemnia inhabit only Neoarctic. The spreading of fossil species shows, that in the past the habitat of the genus was Holarctic and a modern habitat arose due to its descreasing. 112

The tribe Nevermanniini has a lot of genera and species: Hellichiella Rivosecchi, 1925 (4 species), Psilocnetha, 1921 (82 species), Byssodon Enderlein, 1925 (4 species), Psilocnetha Enderlein, 1935 (2 species), Cnetha Enderlein, 1921 (82 species), Nevermulium Rouband, 1907 (29 species), Schoenbaueria Enderlein, 1921 (19 species), Gomphostilbia Enderlein, 1921 (11 species), Morops Enderlein, 1930 (1 species), Montisimulium Rubtsov, 1974 (42 species), (Adler P.H. et all., 2004). After A.V. Yankovsky (2002) and R.W. Crosskey (1990), the black flies of this tribe inhabit almost all the land and have palaearctic-oriental-afrotropic-australian type of the habitat (Nevermannia), Holarctic-oriental habitat is characteristic of the simuliidae of the genera Eusimulium and Cnetha. Besides Holarctic, their representatives are encountered on the Philippine Islands, in Indonesia, Myanmas, India (Crosskey R.W., Chichester U.K., 1990). Hellichiella Byssodon, Schoenbaueria black flies, with the holarctic habitat inhabit the rivers of North America, the north and centre of Eurasia. The black flies of the genus Psilocnetha are in the disjunctive palaearctic-oriental habitat. They were observed in Kazakhstan (Rubtsov I.A., 1982), Egypt and on the island of Ceylon (Adler P.H. et all., 2004). Few representatives of the genera Gomphostilbia, Montisimulium and Moros with oriental- easternpalaearctic habitat are attributed to the mountain chains of the Pacific Ocean coast (Crosskey R.W., Chichester U.K., 1990). To the tribe Nevermanniini belongs the fossil species Hellichiella oligocenicum Rubtsov, 1936. This find is dated from the oligocene period and described from the Baltic amber. The habitat of the fossil species confirms the holarctic habitat of modern species of this genus. The tribe Wilhelmiini in Palearctic is represented by one genus Wilhelmia Enderlein, 1921 (29 species) (Adler P.H. et all., 2004). The black flies of this genus are spread only in Palaearctic. Mediterranian and Central Asian fauna is especially rich in the species of this tribe. The largest tribe Simuliini in terms of species and genera includes Boophthora Enderlein, 1921 (4 species), Psilozia Enderlein, 1936 (1 species), Cleitosimulium Seguy et Dorier, 1936 (4 species), Obuchovia Rubtsov, 1974 (15 species), Paragnus Rubtsov et Yankovsky, 1982 (3 species), Parabyssodon Rubtsov, 1964 (1 species), Archesimulium Rubtsov et Yankovsky, 1982 (22 species), Striatosimulium Rubtsov et Yankovsky, 1982 (15 species), Argentisimulium Rubtcov et Yankovsky, 1982 (8 species), Tetisimulium Rubtcov, 1963 (14 species), Phoretodagmia Rubtcov, 1940 (5 species), Odagmia Enderlein, 1921 (54 species), Gnus Rubtsov, 1940 (28 species), Simulium Latreille, 1802 (53 species). After A.V. Yankovsky (2002) and R.W. Crosskey (1990) the black flies of the tribe inhabit the largest part of the globe and they are absent only in the Australian kingdom. Holarctic-oriental type of the habitat is characteristic of the representatives of the genera Simulium and Archesimulium. Besides Holarctic they are spread in Thailand, Malaysia, Idonesia, India (Adler P.H. et all., 2004). Psilozia, Parabyssodon, Argentisimulium and Gnus with the holarctic habitat inhabit the rivers of North America and Eurasia. The representatives of the genera Boophthora, Tetisimulium, Obuchovia (mediterranian- centralasian-southsiberian type of the habitat), Cleitosimulium (disjunctive mountain mediterranian centralasian), Paragnus (mountain eastern-mediterranian-centralasian) and Phoretodaginia (mountain centralasian) are encountered only in Palaearctic. The genus Odagmia representatives have a palaearctic-oriental habitat. The oriental-easternpalaearctic type of the habitat is characteristic of the black flies of the genus Striatosimulium that was found in Kazakhstan, the Far East, Japan, Thailand, Malasia, India (Rubtsov I.A., 1984). Thus the representatives of two subfamilies (Prosimuliinae, Simuliinae) spread in Palaearctic have holarctic, holarctic-oriental, palaearctic, palaearctic-oriental, oriental- easternpalaearctic and palaearctic- oriental-afrotropical-australian spreading. The core of the sumuliidofauna of Palaearctic consists of the black flies with holarctic and palaearctic habitats.

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The Holarctic type of the habitat is typical to 14 genera (Gymnopai; Twinnia; Prosimulium; Helodon; Stegopterna; Greniera; Cnephia; Metacnephia; Hellichiella; Byssodon; Psilozia; Parabyssodon; Argentisimulium; Gnus), among them disjunctive mountain-holoarctic habitats being characteristic of the genera Gymnopais, Twinnia, but the plain nothern-holarctic one - of the genera Cnephia. The black flies of 4 genera (Cnetha, Eusimulium, Archesimulium, Simulium) have the holarctic-oriental habitat. Simuliidae of 11 genera (Levitinia, Urosimulium, Sulcicnephia, Schoenbaueria, Withelmia, Boophthora, Tetisimulium, Obuchovia, Cleitosimulium, Paragnus, Phoretodagmia) are characterized by palaearctic spreading, among them the black flies of the genus Levitinia have a disjunctive mountain-palaearctic habitat. The palaearctic-oriental habitat is typical to Odagmia black flies. Simuliidae of 4 genera (Gomphostilbia, Morops, Montisimulium, Striatosimulium) have oriental-easternpalaearctic areas. Psilocnetha black flies are characterized by a unique afrotropical-oriental-southern- palaearctiic habitat. They inhabit Central and North Africa, Asia, Kazakhstan and Ceylon. A peculiar palaearctic-oriental-afrotropical-australian habitat is characteristic of the genus Nevermannia.

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