УДК 595.773.4

FAUNA AND BIOECOLOGY OF SNAIL-KILLING (DIPTERA ) IN THE CENTRAL BLACK SOIL REGION OF RUSSIA

S. P. Gaponov

Voronezh State University Поступила в редакцию 11.01.2016 г.

Аннотация. В результате изучения фауны сциомизид в Центральном Черноземье выявлено 54 вида из 21 рода. 12 видов указывается впервые для региона. Исследованы некоторые особенности биологии сциомизид региона и новые хозяева (жертвы) из числа моллюсков. Ключевые слова: сциомизиды, фауны, хозяева, фенология, Центральное Черноземье

Abstract. Fauna of the snail-killing flies in the Central Black soil Region includes 54 from 21 genera; 15 species have been found in the region for the first time. some peculiarities of biology of the sciomyzidae and their relations with molluscs have been studied Keywords: snail-killing flies, fauna, phenology, Central Black soil Region

Family sciomyzidae (snail-killing flies) with coastal vegetation, seasonal ponds, puddles, and contains about 600 species, of which 100-110 can ditches (Anticheta, albiseta, Renocera, some be found in Russia. Currently, the family includes , ). Larvae of aquatic species of subfamilies salticellinae and sciomyzinae (tribae genera , Hydromya, Dictya, Dictyodes, Ilione, sciomizini, Tetanocerini) [1]. Larvae of snail-killing Pherbellia, Hoplodictya, Neolimnia Dichetophora, flies are predators or parasitoids of molluscs; Sepe- , , , Psacadina and some don knutsoni and Sepedonella nana use freshwater Tetanocera [8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19] oligochaetes for their larval development [1]. species are usually predators and attack aquatic nonoperculate whose larvae attack aquatic molluscs are usually pulmonate snails slightly submerged in shallow and predators feeding on several prey [1, 2, 3]. species slow waters or aquatic vegetation mats. some species whose larvae are connected with semi-terrestrial or attack aquatic snails who expose their bodies on terrestrial snails or slugs are either parasitoids or moist surfaces (, , , predators with some features of parasitoids. Based on Bithyniidae, Bulinidae). The operculum of operculate the degree of environmental connections with water prosobrancheal snails usually prevent attacks by and the hosts (prey), sciomyzidae can be divided snail-killing larvae with some exceptions (several into aquatic, semi-aquatic (semi-terrestrial) and species of Pherbellia, Dictya, Hoplodictya and terrestrial ecological groups. Larvae of Renocera, Neolimnia feed on the snails of this group). some Ilione and Eulimnia feed on fingernail clams aquatic pulmonate snails crept out of (: ) under water [4, 5, 6, 7], water or stranded on shore are used by larvae while larvae of Hedria spp. attack aquatic pulmonate of Atrichomelina, Colobaea, some Pherbellia, snails under water. Many species are connected with and Tetanocera for their development. marshes, swamplands, logged ponds, shallow rivers These larvae almost always attack live prey, but continue feeding on decaying organs after they have © Gaponov s. P., 2016 killed the snail. Larvae of Atrichomelina demonstrate saprophagous habits when invading and consuming

54 ВЕСТНИК ВГУ, СЕРИЯ: ХИМИЯ. БИОЛОГИЯ. ФАРМАЦИЯ, 2016, № 1 Fauna and bioecology of snail-killing flies dead mollusc bodies. semi-terrestrial species such (2.3), Grayvoronsky (2.4), Krasnogvardeysky (2.5), as some Colobaea, Pteromicra and many Pherbellia Novooskolsky (2.6), Prokhorovsky (2.7), Rakityansky develop in snails that live near water and (2.8), Rovensky (2.9), shebekinsky (2.10); 3. Kursk connect with coastal vegetation. Terrestrial species Oblast: regions: Glushkovsky (3.1), Gorshechny of genera , Oidematops, Sciomyza, (3.2), Kostorinsky (3.3), Kursky (3.4), sovetsky , Dichetophora, Trypetoptera, Tetanura, (3.5), solntsevsky (3.6), Timsky (3.7), schigrovsky some Pherbellia, Pteromicra and Euthycera attack (3.8); 4Lipetsk Oblast: regions: Gryazinsky (4.1), terrestrial pulmonate snail and/or slugs [20, 21, 22]. Eletsky (4.2), Zadonsky (4.3), Izmalkovsky (4.4), In particular, larvae of some species of Tetanocera Krasnovsky (4.5), Lebedyansky (4.6), Terboonsky and Euthycera develop in slugs [25]. Exposed eggs (4.7), Khlevensky (4.8.), Chapliginsky (4.9); 5. of molluscs are consumed by larvae of Anticheta spp. Tambov Oblast: Zherdevsky (5.1), Znamensky (5.2), [23, 24]. Kirsanovsky (5.3), Michurinsky (5.4), Mordovsky Partial saprophagy of some snail-killing fly larvae (5.5), Morshansky (5.6), Rasskazovsky (5.7), has been discussed [26]. However, there are species Tambovsky (5.8) (fig. 1). whose I-instar larvae start feeding on dead tissue of the victim and then feed on living organs as predators or parasitoids. On the other hand, larvae of some species consume living tissue of their prey until the III-instar, and then finish their development feeding on tissues of already-dead snails. [1]. In general, species of the tribe sciomyzini are more often parasitoids of terrestrial snails. Puparia of terrestrial and semi-terrestrial sciomizini usually stay inside their shells or in shoreline debris, near water or even in water. aquatic predators prevail in Tetanocerini, but some of them are parasitoids or predators of terrestrial snails or slugs [1]. Fig. 1. Fly collection sites. Fauna, morphology and some elements of ecology of snail-killing flies in Russia's Middle Don Region RESULTS AND DISCUSSION and Central Black soil Region have been studied In the territory of the Middle Don Region, during the last 15 years [27, 28, 29, 30, 31, 32, 33, 34, include the Central Black soil Region, 39 species of 35, 36, 37, 38, 39]. These results can now be updated sciomyzidae have been previously observed. This with the new data presented here. research identifies 15 new species for the region. Subfamily Sciomyzinae MATERIAL AND METHODS Tribe Sciomyzini Imago of snail-killing flies have been collected Colobaea Ztt., 1837 in their typical environment using standard Colobaea bifasciella (Fallen, 1820) entomological nets. Dry material from collections Univoltine species. Imago occur in June near of several authors was also analyzed, and additional shallow ponds and temporary water with slow stream. material was bred from the hosts (Tabl.). Numbers Puparia winter inside the host shell. Larvae are para- representing fly collection sites (Fig) correspond sitoids of nonoperculate snails near water surface to the following areas and locations: 1. Voronezh (Lymnaeidae: stagnalis (L., 1758), Stagnic- Oblast: Voronezh-city (1.1), Novovoronezh ola palustris (Muller, 1774)). (1.2), regions: anninsky (1.3), Bobrovsky (1.4), Distribution: 1.1, 1.2, 1.9, 1.10 [27], 1.6, 1.7,1.9, Bogucharsky (1.5), Borisogledsky (1.6), Gribanovsky 1.12; 2.1, 2.3, 2.10; 3.1, 3.4, 3.7; 4.2, 4.3, 4.6, 4.7; (1.7), Kantemirovsky (1.8), Nizhnedevitsky 5.1, 5.7. 26 ♀♀, 18 ♂♂. (1.9), Novousmansky (1.10), Liskinsky (1.11), Colobaea distinctа (Meigen, 1830) Novohopersky (1.12), Olhovatsky (1.13), Pavlovsky Univoltine species. Imago occur in June-July. Pu- (1.14), Paninsky (1.15), Podgorensky (1.16), paria overwinter in host shells. Larvae are parasitoids Rossoshansky (1.17), semiluksky (1.18), Khoholsky of nonoperculate snails living near water surface (L. (1.19), Ertilsky (1.20); 2. Belgorod Oblast: regions: stagnalis, (Muller, 1774), Peregri- alekseevsky (2.1), Belgorodsky (2.2), Valuysky ana peregra (Muller, 1774)).

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Distribution: 1.10 [30], 1.7, 1.11, 1.12, 1.14, 1.17, parasitoids of nonoperculate snails living near water 1.19, 1.20; 2.1, 2.7, 2.8, 2.9; 3.2, 3.4; 3.6, 3.7, 3.8; surface ( planorbis). 4.1, 4.2, 4.3, 4.5; 5.1, 5.2, 5.4, 5.8. 22 ♀♀, 17 ♂♂. Distribution: 1.4, 1.10, 1.15, 1.16 [29], 1.5; 2.2, Colobaea pectoralis (Zetterstadt, 1847) 2.4, 2.9, 2.10; 3.2, 3.3, 3.4, 3.8; 4.1, 4.2, 4.3, 4.5, 4.6, Univoltine species. Imago occur in June-July near 4.9; 5.1, 5.2, 5.5. 40 ♀♀, 34 ♂♂. shallow and slow ponds, ditches and marshes. Puparia (Fallen, 1820) overwinter inside prey shells. Larvae are predators of Multivoltine species. Imago occur in June-begin- nonoperculate snails living near water surface or with ning of July and in august-beginning of september bodies exposed on moist surfaces. This species is ob- usually at the shoreline of coastal vegetation. Puparia served for the first time in the Region. are overwintering; in the southern parts of Voronezh Distribution: 1.20; 2.5, 2.10; 3.2, 3.3; 4.3. 16 ♀♀, Oblast, imago are overwintering. Larvae are preda- 9 ♂♂. tors of succineidae. Colobaea punctata (Lundbeck 1923) Distribution: 1.1, 1.2, 1.4, 1.9, 1.10 [27, 29], 1.8, Univoltine species. Imago occur in June usually 1.14, 1.18; 2.1, 2,3, 2.6, 2.9, 2.10; 3.3, 3.4; 4.1, 4.2, near shore edges of shallow seasonal water, swamp- 4.3, 4.4, 4.8; 5.3, 5.7. 18 ♀♀, 17 ♂♂. lands and lakes with abundant aquatic vegetation. Pu- Pherbellia dorsata (Zetterstadt, 1846) paria overwinter inside prey shells. Larvae are preda- Univoltine species. Imago occur in august usual- tors of nonoperculate snails living near water surface ly near logged water. Puparia overwinter inside prey (G. truncatula). shells. Larvae are predators of nonoperculate snails Distribution: 1.1, 1.2, 1.9, 1.10, 1.18 [29, 35, 39], living near the water surface (Planorbis, Lymnea). 1.8, 1.15, 1.17; 2.2, 2.5, 2.9; 3.4, 3.6; 4.1, 4.2, 4.3, This species is observed for the first time in the Re- 4.4, 4.9; 5.1, 5.2, 5.4, 5.6, 5.7. 27 ♀♀, 25 ♂♂ gion. Ditaeniella sack, 1939 Distribution: 1.11. 3 ♀♀, 3 ♂♂. Ditaeniella grisescens (Meigen 1830) Pherbellia dubia (Fallen, 1820) Univoltine species. Imago occur in June usu- Univoltine species. Imago occur in July near shore ally near a shore edges of shallow seasonal waters, vegetation. Puparia overwinter inside host shells. swamps and marshes. Puparia overwinter outside Larvae are parasitoids of terrestrial and semi-terres- prey shells. Larvae are predators of nonoperculate trial snails (succineidae, Zonitidae). snails living near water surface. This species is ob- Distribution: 1.1., 1.9, 1.10 [27, 29], 1.3, 1.7, served for the first time in the Region. 1.11, 1.14, 1.17, 1.19, 1.20; 2.1, 2.2, 2.3, 2.6, 2.7, 2.9; Distribution: 1.5, 1.8; 2.1, 2.2, 2.7; 3.1, 3.2; 4.3, 3.1, 3.2, 3.3, 3.8; 4.1, 4.2, 4.3, 4.4, 4.6, 4.7, 4.8, 4.9; 4.8; 5.8. 14 ♀♀, 22 ♂♂. 5.3, 5.4, 5.8. 29 ♀♀, 31 ♂♂. Pherbellia R.-D., 1830 Pherbellia griseola (Fallen, 1820) Pherbellia albocostata (Fll., 1820) Univoltine species. Imago occur in July near the Univoltine species. Imago occur in august usual- shoreline. Puparia overwinter inside prey shells. Lar- ly at river shores. Puparia are in the debris near prey vae are predators of the snails at coastal line. This shells. Larvae are predators of succineidae. species is observed for the first time in the Region. Distribution: 1.9, 1,10, 1.16 [29], 1.19, 1.20; 2.1, Distribution: 1.3, 1.4; 2.2; 3.2, 3.3; 4.3. 5 ♀♀, 7 2.3, 2.5, 2.9, 2.10; 3.1,3.2, 3.6, 3.7; 4.1, 4.2, 4.3, 4.4, ♂♂. 4.7, 4.9; 5.1, 5.3, 5.6. 42 ♀♀, 36 ♂♂. (Fallen, 1820) (Fll., 1826) Multivoltine species. Imago occur in June-sep- Multivoltine species. Imago occur in June and tember near shallow water with dense vegetation, sea- august-september. Puparia are overwintering. Lar- sonal puddles and ditches. Puparia overwinter inside vae are parasitoids of succineidae. This species is prey shells. Larvae are predators of nonoperculate observed for the first time in the Region. snails near water surface. This species is observed for Distribution: 1.6, 1.11; 3.3; 4.3, 4.7; 5.4. 6 ♀♀, the first time in the Region. 3 ♂♂. Distribution: 1.11; 5.1, 5.2. 2 ♀♀, 3 ♂♂. Pherbellia argyra Verbeke 1967 Pherbellia obtusa (Fallen, 1820) Univoltine species. Imago occur in July usu- Univoltine species. Imago occur in July. Puparia ally near shoreline of shallow ponds, such as marsh- overwinter inside prey shells. Larvae are parasitoids/ es, ditches, swamps and logged ponds. Puparia are predators of nonoperculate snails near water surface. overwintering, often outside of host shells. Larvae are Distribution: 1.10 [30].

56 ВЕСТНИК ВГУ, СЕРИЯ: ХИМИЯ. БИОЛОГИЯ. ФАРМАЦИЯ, 2016, № 1 Fauna and bioecology of snail-killing flies

Pherbellia pilosa (Hendel, 1902) near logged water. Puparia are outside of host shells. Imao occur in July. Puparia overwinter inside Larvae are parasitoids of succineidae [13]. prey shells. Larvae are predators of nonoperculate Distribution: 1.1, 1.2, 1.4, 1.10 [29], 1.3, 1.6, 1.7, snails near water surface. 1.11, 1.18; 2.1, 2.3, 2.8, 2.9, 2.10, 3.6, 3.7; 4.2, 4.3, Distribution: 1.10 [30]. 4.8; 5.1, 5.2, 5.6. 28 ♀♀, 30 ♂♂. Pherbellia scutellaris (Roser, 1846) Tetanura Fll., 1820 Larvae are parasitoids of Clausilia. This species Tetanura pallidiventris (Fallen, 1820) is observed for the first time in the Region. Multivoltine species. Imago occur in June-au- Distribution: 1.6, 1.7, 1.11; 4.2, 4.3; 5.1, 5.5. 11 gust in bushy stations and edges of woods. Puparia ♀♀, 14 ♂♂. are outside of host shells. Larvae are parasitoids of Pteromicra Lioy, 1864 terrestrial snails [40]. Pteromicra glabricula (Fallen, 1820) Distribution: 1.1, 1.2, 1.4, 1.10 [29], 1.3, 1.6, 1.7, Univoltine species. Imago occur in May-begin- 1.9, 1.11, 1.12; 2.6, 2.7, 2.10; 3.3, 3.6; 4.1, 4.3, 4.6, ning of June near shores of swamps, marshes and 4.7; 5.7. 24 ♀♀, 28 ♂♂. small rivers with aquatic vegetation. Larvae are pred- Tribe Tetanocerini ators of nonoperculate snails. Anticheta Haliday, 1838 Distribution: 1.4, 1.5, 1.9, 1.10 [29; 30], 1.3, 1.18; Anticheta analis (Meigen, 1830) 4.2, 4.3, 4.6; 5.1, 5.2, 5.5. 24 ♀♀, 26 ♂♂. Univoltine species. Imago occur in June- (staeger, 1845) beginning of July near shallow water. Puparia over- Univoltine species. Imago occur in June near winter inside prey shells. Larvae are predators of logged lakes, ponds and swamps. Puparia are outside exposed snail eggs. of host shells. Larvae are predators of nonoperculate Distribution: 1.1, 1.2, 1.4, 1.10, 1.14 [29], 1.16, snails. 1.17, 1.19, 1.20; 2.1, 2.3, 2.8, 2.9, 2.10; 3.2, 3.5, 3.8; Distribution: 1.5, 1.10, 1.11, 1.17 [29, 31], 1.3, 4.1, 4.2, 4.3, 4.8; 5.4, 5.5, 5.8. 42 ♀♀, 38 ♂♂. 1.4; 2.1, 2.3, 2.4, 2.8, 2.9, 3.3, 3.5, 3.7; 4.1, 4.2, 4.3, Anticheta brevipennis (Zetterstadt, 1846) 4.5, 4.8, 4.9; 5.2, 5.3. 31 ♀♀, 22 ♂♂. Univoltine species. Imago occur in June-July Pteromicra leucopeza (Meigen, 1838) usually in moist areas with grass vegetation not far Univoltine species (?). Imago occur in June-July from water. Puparia are overwintering. Larvae are near logged rivers, lakes and marshes. Larvae are predators of eggs of Lymnaeidae and succineidae. predators of nonoperculate snails. Distribution: 1.4 [29], 1.3; 2.2; 3.4; 5.1. 9 ♀♀, 5 Distribution: 1.10 [30]. ♂♂. Pteromicra oldenbergi (Hendel, 1902) Anticheta atriseta (Loew 1849) Imago occur in July near logged rivers, lakes Univoltine species. Imago occur in June near and marshes. Larvae are predators of nonoperculate shallow water. Puparia are overwintering. Larvae are snails. predators of eggs of snails. This species is observed Distribution: 1.10 [30]. for the first time in the Region. Sciomyza Fll., 1820 Distribution: 1.3, 1.4, 1.7, 1.11; 2.1, 2.3, 2.6, 2.7; 21. Sciomyza simplex (Fallen, 1820) 3.3; 4.2, 4.3. 22 ♀♀, 22 ♂♂. Multivoltine species. Imago occur in July-august Coremacera Rd., 1856 near logged ponds. Puparia are outside of host shells. (Fabricius, 1775) Larvae are parasitoids of succineidae. Univoltine species. Imago occur in May-July Distribution: 1.10 [30], 1.4, 1.12, 1.3, 1.18 [31], usually in relatively dry open conditions at the mead- 1.8, 1.11, 1.14; 2.1, 2.2, 2.3, 2.5, 2.9, 2.10; 3.1, 3.2, ows and the edges of woods. Puparia are overwinter- 3.3, 3.4, 3.6; 4.1, 4.2, 4.3, 4.4, 4.9; 5.2, 5.6, 5.7, 5.8. ing. Larvae are predators of terrestrial snails. 46 ♀♀, 38 ♂♂. Distribution: 1.10 [30], 1.16 [31], 1.5, 1.8, 1.14, Sciomyza lucida (Hendel, 1902) 1.17; 2.3, 2.8, 2.9; 3.6; 4.1, 4.2, 4.3, 4.9; 5.2, 5.8. 33 Multivoltine species. Imago occur in June-July ♀♀, 28 ♂♂. near logged water. Puparia are outside of host shells. Coremacera catenata (Loew 1847) Larvae are parasitoids of Stagnicola. Univoltine species. Imago occur in May-July Distribution: 1.10 [30]. usually in relatively dry open conditions at the mead- Sciomyza testacea Macquart, 1835 ows and the edges of woods. Puparia are overwin- Multivoltine species. Imago occur in June-July tering. Larvae are predators of terrestrial snails [22].

ВЕСТНИК ВГУ, СЕРИЯ: ХИМИЯ. БИОЛОГИЯ. ФАРМАЦИЯ, 2016, № 1 57 Gaponov S. P.

This species is observed for the first time in the Re- Univoltine species. Imago occur in June-July gion. usually at the meadows, wet woods. Puparia are over- Distribution: 1.7, 1.11; 2.3; 4.3. 4 ♀♀, 2 ♂♂. wintering. Larvae are endoparasitoids of slugs. Dichetophora Rd., 1868 Distribution: 1.10 [30]. Dichetophora finlandica Verbeke, 1964 Euthycera stictica (Fabricius, 1805) Univoltine species. Imago occur in June usually Univoltine species. Imago occur in June-July in relatively dry open conditions in meadows and the usually at the meadows, wet woods. Puparia are over- edges of woods. Puparia are overwintering. Larvae wintering. Larvae are probably parasitoids of slugs. are predators/parasitoids of terrestrial snails. This species is observed for the first time in the Re- Distribution: 1.10 [30]. gion. Dictya Mg., 1803 Distribution: 1.5, 1.8. ♂♂. Dictya umbratum (Linnaeus, 1758) Hydromya R.-D., 1830 Multivoltine species. Imago occur in april-May Hydromya dorsalis (Fabricius, 1775) and august near shorelines of shallow and slow riv- Multivoltine species. Imago occur in May and ers, marshes and swamps. Puparia are overwintering. august-september. III-instar larvae are overwintering. Larvae are predators of nonoperculate molluscs liv- Larvae are predators of nonoperculate snails living ing near water surface or nonoperculate and opercu- close to water surface. This species is observed for late submerged snails [41]. the first time in the Region. Distribution: 1.4, 1.10 [29], 1.9, 1.20; 2.3, 2.9; Distribution: 1.7, 1.12; 2.2; 4.3. 7 ♀♀, 5 ♂♂. 3.7; 4.2, 4.3, 4.5, 4.6; 5.1. 17 ♀♀, 16 ♂♂. Ilione Haliday et Curtis, 1837 Elgiva Mg., 1838 (Fallen, 1820) (Linnaeus, 1767) Univoltine species. Imago occur in July-august Multivoltine species. Imago occur in May- near the shoreline. I-instar larvae are overwintering. June and October at shorelines of logged ponds Larvae are aquatic predators/parasitoids of fingernail and marshes. Puparia are overwintering; in the clams (sphaeriidae) [1]. southern parts of Voronezh Oblast imago are Distribution: 1.10 [30]. overwintering. Larvae are aquatic predators of (scopoli, 1763) nonoperculate snails living near water surface – Univoltine species. Imago occur in July near Limneaidae [14]. shorelines of shallow water, including seasonal Distribution: Воронежская обл.: 1.10 [28; 30], ditches and ponds. III-instar larvae are overwintering. 1.3, 1.4, 1.8, 1.16, 1.19; 2.2, 2.3, 2.8, 2.9; 3.8; 4.1, aquatic predators of nonoperculate snails living near 4.2, 4.3, 4.4, 4.9; 5.2, 5.3. 41 ♀♀, 38 ♂♂. water surface [42]. (Harris, 1780) Distribution: 1.10 [30]. Multivoltine species. Imago occur in May-June Ilione rossica (Mayer 1953) and august. Puparia are overwintering; in the south- Univoltine species. Imago occur in July-august ern parts of Voronezh Oblast imago are overwinter- near shorelines of shallow water, including seasonal ing. Larvae are aquatic predators of nonoperculate ditches and ponds. I-instar larvae are overwintering. snails living near water surface. aquatic predators of nonoperculate snails living near Distribution: 1.4, 1.6, 1.10, 1.11, 1.18 [29], 1.7, water surface. 1.8, 1.17, 1.20; 2.1, 2.2, 2.10; 3.5, 3.6, 3.7; 4.2, 4.3, Distribution: 1.4, 1.5, 1.10, 1.18 [29], 1.3, 1.15, 4.9. 36 ♀♀, 47 ♂♂. 1.16, 1.19; 2.1, 2.3, 2.4, 2.7, 2.9; 3.8; 4.1, 4.2, 4.3, Elgiva divisa Loew, 1845 4.6, 4.8; 5.2, 5.4. 24 ♀♀, 19 ♂♂. Multivoltine species. Imago occur in May-June Limnia R.-D., 1830 and July-august at shorelines of logged ponds and Elberg 1965 marshes. Puparia are overwintering; in the southern Univoltine species. Imago occur in July-august parts of Voronezh Oblast imago are overwintering. in wet meadows. Larvae are predators of succineidae Larvae are aquatic predators of nonoperculate snails and slugs. This species is observed for the first time living near water surface. in the Region. Distribution: 1.6 [31], 1.12, 1.19; 2.9, 3.1; 4.3, Distribution: 1.12. 2 ♂♂. 4.4; 5.8. 11 ♀♀, 12 ♂♂. Limnia unguicornis (scopoli, 1763) Euthycera Latr., 1829 Univoltine species. Imago occur in July-august Euthycera chaerophylli (Fabricius, 1798) at river coasts with well-developed vegetation,

58 ВЕСТНИК ВГУ, СЕРИЯ: ХИМИЯ. БИОЛОГИЯ. ФАРМАЦИЯ, 2016, № 1 Fauna and bioecology of snail-killing flies sometimes near swamps. Larvae are predators/ Sepedon spinipes (scopoli, 1763) parasitoids of succineidae and slugs [43]. Multivoltine species. Imago occur in May-sep- Distribution: 1.1, 1.4, 1.6, [29, 31], 1.10 [30], tember near shallow water. Puparia are overwinter- 1.12, 1.15, 1.16, 1.19, 1.20; 2.1, 2.2, 2.3, 2.4, 2.7, 2.8, ing; in the southern parts of Voronezh Oblast imago 2.9, 2.10; 3.1, 3.2, 3.3, 3.7; 4.1, 4.2, 4.3, 4.6, 4.8; 5.2, are overwintering. Larvae are aquatic predators of 5.3, 5.7, 5.8. 46 ♀♀, 43 ♂♂. nonoperculate snails and semi-terrestrial succineidae. Pherbina R.-D., 1830 Distribution: 1.10 [30] (scopoli, 1763) Sepedon sphegea (Fabricius, 1775) Univoltine species. Imago occur in May-June near Multivoltine species. Imago occur in May-sep- shallow water. III-instar larvae are overwintering. tember near shallow water. Puparia are overwinter- Larvae are aquatic predators of nonoperculate snails ing; in the southern parts of Voronezh Oblast imago near water surface. are overwintering. Larvae are aquatic predators of Distribution: 1,9,1.10 [29, 30 ], 1.16, 1.17, 1.19, nonoperculate snails near water surface and semi-ter- 1.20; 2.2, 2.6, 2.7, 2.8; 3.2, 3.4, 3.8; 4.1, 4.2, 4.3, 4.9; restrial succineidae. 5.6, 5.7. 21 ♀♀, 18 ♂♂. Distribution: 1.1, 1.2, 1.4 [27, 29],1.10 [29, 30], Pherbina intermedia Verbeke, 1848 1.12, 1.16, 1.19, 1.20; 2.1, 2.2, 2.3, 2.7, 2.8, 2.9, 2.10; Univoltine species. Imago occur in July near 3.1, 3.2, 2.5, 3.8; 4.1, 4.2, 4.3, 4.6, 4.8; 5.1, 5.4, 5.5. shallow water. III-instar larvae are overwintering. 32 ♀♀, 27 ♂♂. Larvae are aquatic predators of nonoperculate snails Tetanocera Dumeril, 1800 near water surface. Tetanocera elata (Fabricius, 1781) Distribution: 1.10 [30]. Multivoltine species. Imago occur in June-au- Psacadina Enderlein, 1939 gust usually at the edges of woods or wet meadows. Psacadina vittigera (schiner, 1864) Puparia are overwintering. Obligatory predators of Univoltine species. Imago occur in June-sep- slugs with some features of parasitoids. tember near shallow water. III-instar larvae are over- Distribution: 1.1, 1.2, 1.4, 1.12, 1.18 [27, 29], 2.1, wintering. Larvae are aquatic and semi-terrestrial 2.2, 2.3, 2.4, 2.7; 3.3, 3.8; 4.2, 4.3, 4.6, 4.8; 5.1, 5.5, predators of nonoperculate snails near water surface. 5.7, 5.8. 34 ♀♀, 29 ♂♂. This species is observed for the first time in the Re- Tetanocera ferruginea Fallen, 1820 gion. Multivoltine species. Imago occur in May-July Distribution: 1.12. 1 ♂. and september-October usually at the edges of woods Psacadina verbekei Rozkosny 1975 or wet meadows. Puparia are overwintering. Larvae Univoltine species. Imago occur in april-august are aquatic predators of nonoperculate snails whose near shallow water. III-instar larvae are overwinter- bodies are exposed on wet surfaces. ing. Larvae are aquatic and semi-terrestrial predators Distribution: 1.2, 1.4, 1.8 [31], 1.6, 1.7, 1.11; 2.2, of nonoperculate snails near water surface. This spe- 2.5; 3.3, 3.4; 4.1, 4.3, 4.6; 5.8. 22 ♀♀, 26 ♂♂. cies is observed for the first time in the Region. Tetanocera robusta Loew, 1847 Distribution: 1.7. 2 ♂♂. Multivoltine species. Imago occur in May-sep- Psacadina zernyi (Mayer, 1953) tember usually at woods edges or wet meadows. Pu- Univoltine species. Imago occur in May-sep- paria are overwintering. Larvae are predators of slugs tember near shallow water. Larvae are aquatic and and snails Zonitoides, . semi-terrestrial predators of nonoperculate snails Distribution: 1.1, 1.2, 1.4, 1.6, 1.16 [27, 29], 1.12, near water surface. This species is observed for the 1.14, 1.17, 1.19, 1.20; 2.2, 2.3, 2.6, 2.7, 2.9; 3.1, 3.2, first time in the Region. 3.6, 3.7; 4.1, 4.2, 4.3, 4.8; 5.3, 5.5, 5.7. 40 ♀♀, 35 Distribution: 1.6, 1.6, 1.12. ♂♂. ♂♂. Renocera Hendel, 1900 Tryprtoptera Hendel, 1900 Renocera pallidа (Fallen, 1820) scopoli Univoltine species. Imago occur in June-august Univoltine species. Imago occur in May-august near the shoreline. Puparia are overwintering. Larvae usually in bushy stations or at the edges of woods. Pu- are aquatic predators/parasitoids of fingernail clams paria overwinter outside prey shells. Larvae are pred- (sphaeriidae) [5, 6, 7]. ators of terrestrial viviparous snails Lauria cylind- Distribution: 1.10 [30]. racea [44] and sometimes aquatic and semi-aquatic Sepedon Latr., 1804 predators of Lymnea, Vertigo, Zonitoides.

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Table List of Sciomyzidae breeding sciomyzid species Host/prey species 1 Colobaea bifasciella Lymnaeidae: Lymnaea stagnalis (L., 1758), (Muller, 1774) Lymnaeidae: L. stagnalis, Galba truncatula (Muller, 1774), Peregriana peregra 2 Colobaea distinctа (Muller, 1774) 3 Colobaea punctata Planorbidae: Planorbis planorbis [29], Limnaeidae: G. truncatula Planorbidae: Planorbis planorbis (L., 1758); Anisus septemgyratus (Rossmaessler, 4 Pherbellia argyra 1858); Planorbis carinatus (Muller, 1774) 5 Pherbellia cinerella succineidae: Succinea oblonga Drap., L. Pteromicra 6 Lymnaeidae: Galba truncatula; Physidae: fontinalis (L., 1758) angustipennis 7 Sciomyza testacea succineidae: Succinea putris 8 Tetanura pallidiventris Pupilidae: Vertigo antivertigo; Zonitidae: Zonitoides nitidus Muller 9 Coremacera marginata Zonitidae: Zonitoides nitidus 10 Elgiva divisa Limnaeidae: Limnea stagnalis [29, 35, 39] 11 Euthycera chaerophylli Limacidae: Agriolimax agrestis 12 Ilione albiseta Physidae: Physa fontinalis 13 Limnia unguicornis succineidae: Succinea putris 14 Sepedon sphegea Lymnaeidae: Peregriana peregra; Stagnicola palustris 15 Tetanocera ferruginea Lymnaeidae: Lymnaea stagnalis; Galba truncatula 16 Tetanocera elata Limacidae: Agriolimax agrestis (L.); A. reticulatus (Muller) 17 Tetanocera robusta Limacidae: Limax sp. [29, 35, 39] Trypetoptera punctu- 18 Pupillidae: Vertigo antivertigo Drap.; Zonitidae: Zonitoides nitidus lata

Distribution: 1.1, 1.4, 1.6, 1.7, 1.10 [27, 29]; 1.10 yleti, L. paludicola, some Tetanocera. Consumption [30], 1.5, 1.18, 1,19; 2.1, 2.2, 2.4, 2.5, 2.9; 3.1, 3.2, of dead host/prey tissues is usually in addition to the 3.6, 3.7; 4.3, 4.8, 4.9; 5.2, 5.3, 5.7. 38 ♀♀, 28 ♂♂. main feeding on living mollusc organs. The typical aquatic species in the region are CONCLUSION Ilione albiseta, Renocera pallida, Sepedon spinipes, Current fauna of snail-killing flies in the Central S. sphegea and Tetanocera ferruginea. several Black soil Region of Russia comprise 54 species species are semi-terrestrial, including Pherbellia from 21 genera. Of these, 15 species were identified schoehnerri, Sciomyza testacea and Trypetoptera for the first time by this study. punctulata. Terrestrial larvae are characteristic of Breeding data representing 14 sciomyzidae host/ Euthycera chaerophylli, Coremacera marginata, prey species are reported for the first time. In total, Dichetophora, Tetanura pallidiventris, Trypetoptera 18 sciomyzidae host/prey species are known for the punctulata, Pherbellia cinerella, Limnea anguicornis Region. and Tetanocera elata. The majority of species of sciomyzidae in the region are predators at larval stage (42 species); some REFERENCES of them have features of parasitoids. 12 species are 1. Knutson L.V. snail-killing sciomyzidae parasitoids at their larval stage. some occurrences Flies. / L.V. Knutson, J.-C. Valá — Cambridge Univ. of saprophagy on dead molluscs were observed for Press, 2011. — 505 pp. larvae of Ph. cinerella, C. pectoralis, C. punctata, 2. Rozkošný R. a review of Palaearctic D. grisescens, Pteromicra spp., S. simplex, Ph. cor- sciomyzidae (Diptera) / R. Rozkošný // Fol. Facul.

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26. Пантелеева Н.Ю. Новые сведения С.П. // Состояние и проблемы экосистем Средне- о сапрофаги личинок акалиптратных ко- русской лесостепи. Тр. биологического учебно- роткоусых двукрылых (Diptera, , научного центра «Веневитиново» Воронежского acalyptrata) Среднего Подонья. I. Надсемейства госуниверситета. — Воронеж: ВГУ, 2004. — Вып. , , , , 17. — С. 55-63. , и / 36. Gaponov s.P. Fine morphology of Н.Ю. Пантелеева, С.П. Гапонов // Вестник ВГУ, sciomyzidae (Diptera) female head sensory system Серия: Химия. Биология. Фармация. — 2015. — / s.P. Gaponov, L.N. Khitzova, a.s. sergeev // Proc. № 3. — С. 78-84. VI International Congress Dipterology. — Japan, 27. Gaponov s. at the fauna of sciomyzidae 2006. — 98 р. (Diptera) in the Central Black soil Region of Russia / 37. Гапонов С.П. Морфология сенсорной s. Gaponov, V Pershina // abstr. Vol. of 6th European системы головы самок сциомизид (Diptera, Congress of Entomology. — V. 1. — Czeske sciomyzidae) / С.П. Гапонов, А.С. Сергеев, Л.Н. Budeiovici, 1998. — P. 349. Хицова, В.А. Резнева // Состояние и проблемы 28. Силина А.Е. Вылет короткоусых двукры- экосистем Среднерусской лесостепи. – Воронеж: лых (Diptera, Brachycera) из реки Усмань / А.Е. ВГУ, 2005. — Вып. 19. — С. 65-75. Силина, О.Н. Чалая // Проблемы энтомологии в 38. Гапонов С.П. Новые диптерологические России. — СПб., 1997. — T. 2. — С. 119. находки в Центральном Черноземье и морфо- 29. Гапонов С.П. К фауне и биологии мух- логия яиц двукрылых из семейств , сциомизид (Diptera, sciomyzidae) Среднего По- , и (Diptera) / С.П. Га- донья / С.П. Гапонов, В.А. Резнева-Першина // понов // Состояние и проблемы экосистем Сред- Состояние и проблемы экосистем Среднерус- нерусской лесостепи. — Воронеж: ВГУ, 2000, a. ской лесостепи. — Воронеж: ВГУ, 2000. — Вып. — Вып. 14. — С. 63-67. 14. — С. 67-73. 39. Гапонов С.П. Эколого-фаунистическая 30. Пантелеева Н.Ю. К изучению мух се- характеристика сциомизид (Diptera, sciomyzidae) мейства sciomyzidae в Центральном Черноземье Среднего Подонья. / С.П. Гапонов // Экология, / Н.Ю. Пантелеева, С.А. Горбань // Состояние и эволюция и систематика животных. Мат. Всерос. проблемы экосистем среднерусской лесостепи — научн.-практ. конференции с междунар. участием. Вып. 14. — Воронеж: ВГУ, 2000. — С.72-80. — Рязань, 2009. — С. 56-57. 31. Гапонов С.П. К фауне мух-сциомизид 40. Knutson L.V. Biology and immature stages окрестностей Воронежского водохранилища / of Tetanura pallidiventris, a parasitoid of terrestrial С.П. Гапонов, А.С. Сергеев // Алтай: Экология и snails (Diptera, sciomyzidae) / L.V. Knutson // Ent. природопользование. Труды IV Российско-мон- scand. — 1970. — Vol. 1, № 2. — P. 81-89. гольской научной конференции молодых ученых 41. Willomitzer J. Further observation on the и студентов. — Бийск, 2005. — С. 105-108. rearing of sciomyzid larvae (Diptera) for the control of 32. Гапонов С.П. Морфология и эволюцион- intermediate host snails / J. Willomitzer, R. Rozkosny ные преобразования яиц двукрылых (Diptera). / // acta Vet. Brno. — 1977. — Vol. 46. — P. 315-322. С.П. Гапонов. — Воронеж: ВГУ, 2003. — 323 с. 42. Lindsay W. Biology of the snail-killing fly 33. Гапонов С.П. Биология и стадия яйца дву- Ilione albiseta (scopoli) (Diptera: sciomyzidae) / W. крылых из семейства sciomyzidae (Diptera) Цен- Lindsay, R.J. McDonnel, C.D. Williams, L. Knutson, трального Черноземья / С.П. Гапонов. — М., 2002. M.J. Gormally // studia dipterol. — 2011. — Vol. 16. - Деп. в ВИНИТИ 13.03.02, № 466 — В2002. — 36 с. — P. 245-307. 34. Гапонов С.П. Морфология яиц, особенно- 43. Vala J.-C. stades immature et biologie de сти развития сциомизид (Diptera, sciomyzidae) и Limnia anguicornis (scopoli), Diptėre sciomyzidae система семейства / С.П. Гапонов // Мат. II Все- prėdateur de mollusques / J.-C. Vala, L.V. Knutson. российского симпозиума по амфибиотическим // ann. soc. Ent. Fr. (N.s.). — 1990. — Vol. 26, № и водным насекомым: фауна, вопросы экологии, 3. — P. 443-450. морфологии и эволюции амфибиотических и во- 44. Vala J.-C. Description des stades larvaires дных насекомых России. — Воронеж: ВГУ, 2004. et donnėes sur la biologie et la phėnologie de — С. 18-28. Trypetoptera punctulata (Diptera, sciomyzidae) / J.- 35. Гапонов С.П. Эколого-морфологические C. Vala // ann. sco. Ent. Fr. (N.s.). — 1986. — Vol. особенности яиц sciomyzidae (Diptera) / Гапонов 22, № 1. — P. 67-77.

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Воронежский государственный университет Voronezh State University Гапонов С. П., доктор биологических наук, Gaponov S. P., PhD, DSci, Full Professor, Head профессор, заведующий кафедрой зоологии и па- of the Department of Zoology and Parasitology разитологии

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