Review of the genus Ilione (Diptera, Sciomyzidae) Текст научной статьи по специальности «Биологические науки»

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Амурский зоологический журнал, 2024, т. XVI, № 2

Amurian Zoological Journal, 2024, vol. XVI, no. 2

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https://www.doi.org/10.33910/2686-9519-2024-16-2-310-325

https://zoobank.org/References/59BFC186-A536-4037-9CFA-47BD310C5A2D

UDC 595.773.4

Review of the genus Ilione (Diptera, Sciomyzidae)

N. E. VikhrevH, M. O. Yanbulat

Zoological Museum of Moscow University, 2 Bolshaya Nikitskaya Str., 125009, Moscow, Russia

Authors Nikita E. Vikhrev

E-mail: nikita6510@yandex.ru

SPIN: 1266-1140

Scopus Author ID: 32467511100

Maria O. Yanbulat

E-mail: mairynia@yandex.ru

SPIN: 4159-3415

Abstract. A review of a mostly West Palaearctic genus Ilione is offered. It includes eight species of Ilione regarded as valid after the last revision of the genus (Verbeke 1964) and one here described species from North India (I. bindata sp. nov.). The publication is divided into two parts. Part 1 is devoted to five common species which comprise about 99% specimens of Ilione in insect collections. We examined representative material of these species and provide new illustrations and a revised identification key for them. Part 2 is devoted to three rare sub-Mediterranean species of Ilione and one newly described species. In in each part we offer descriptions of available specimens and taxonomic comments.

Copyright: © The Authors (2024). Published by Herzen State Pedagogical University of Russia. Open access under CC BY-NC License 4.0.

Keywords: Diptera, Sciomyzidae, Ilione, review, identification key

Обзор рода Лют (Diptera, Sciomyzidae)

Н. Е. Вихревн, М. О. Янбулат

Зоологический музей МГУ им. М. В. Ломоносова, ул. Большая Никитская, д. 2, 125009, г. Москва, Россия

Сведения об авторах Вихрев Никита Евгеньевич E-mail: nikita6510@yandex.ru SPIN-код: 1266-1140 Scopus Author ID: 32467511100

Янбулат Мария Олеговна E-mail: mairynia@yandex.ru SPIN-код: 4159-3415

Аннотация. Предложен обзор преимущественно западно-палеарктического рода Шопе. Наш обзор включает все восемь видов Шопе, признанных валидными после последней ревизии рода (УегЬеке 1964), и один описанный здесь вид из Северной Индии (I. ЫпйМа sp. по^). Публикация разделена на две части. Часть I посвящена пяти распространенным видам, которые составляют около 99% экземпляров Шопе в коллекциях насекомых и репрезентативный материал которых мы изучили. Мы предлагаем новые иллюстрации и пересмотренный идентификационный ключ для этих видов. Часть II посвящена трем малоизвестным субсредиземноморским видам Шопе и одному описываемому здесь виду. В каждом случае мы предлагаем описание имеющихся экземпляров и даем таксономические комментарии.

Права: © Авторы (2024). Опубликовано Российским государственным педагогическим университетом им.

А. И. Герцена. Открытый доступ на Ключевые слова: Diptera, Sciomyzidae, Шопе, обзор, определительный условиях лицензии СС БУ-ЫС 4.0. ключ

Introduction

Ilione Haliday, 1837 is a genus of the tribe Tetanocerini. Ilione is endemic to the Palae-arctic and clearly originated from the West Palaearctic. The genus was also known as Knutsonia Verbeke, 1964. The reasons for changing its name to Ilione were discussed in Steyskal et al. (Steyskal et al. 2003). The synonymy of the species of Ilione is given in accordance with the catalog (Rozkosny, Elberg 1984).

Ilione is characterised by the following characters: two pairs of orbital setae; pedicel rectangular, postpedicel triangular, both elongated; arista with short, dense, entirely or partly white hairs; anepisternum and an-epimeron setulose; subalar setae usually present; scutellum with two pairs of setae; wing without dark pattern, but crossveins may be darkened and dark spots along M1+2 may be present, hind coxa with setulae on inner posterior margin.

The phylogenetic relationship of Ilione to other Tetanocerini is unclear. According to the molecular analysis (Chapman et al. 2012), I. albiseta is related to the Nearctic Hedrina mixta Steyskal, 1954. According to Tothova et al., Ilione is a paraphyletic genus partly related to Dichetophora (I. lineata) and partly to

Limnia + Pherbina + Trypetoptera (Tothova et al. 2012). Another reason to suspect that the genus Ilione is not monophyletic, is the biology of I. lineata larvae. In contrast with other Ilione and vast majority of Sciomyzi-dae, it parasitises not in Gastropoda but Bi-valvia (Sphaeriidae) molluscs (Foote, Knutson 1970). All these possibilities may have sense, but here we consider Ilione in the old classical volume offered by Verbeke (Verbeke 1964).

In our opinion, there are two interesting and original sources on the taxonomy of Ilione. The genus was revised by Ver-beke, who established several synonymies, illustrated the remaining eight species, and offered keys for them (a detailed key in French, in which each couplet has descriptive notes, and a shorter key in English) (Verbeke 1964).

The second important source is Rozkosny, who gave original and very comprehensive drawings of the male terminalia of the Scandinavian species of Ilione (Rozkosny 1987: 222-224). Rozkosny also offered a short key for the genus Ilione, which is well-known among those working with Sciomyzidae. In order to save the readers from a search for illustrations from the above-mentioned publications, we partly reproduced them below (Figs 1, 2, 3).

Fig. 1. Male terminalia of Ilione from Verbeke (Verbeke 1964: 9-16): a — albiseta; b — lineata; c — trifaria; d — turcestanica; e — rossica; f — truquii; g — corcyrensis; h — unipunctata

Рис. 1. Терминалии самцов Ilione по Verbeke (Verbeke 1964: 9-16): a — albiseta; b — lineata; c — trifaria; d — turcestanica; e — rossica; f — truquii; g — corcyrensis; h — unipunctata

Fig. 2. Male sternite 4 of Ilione from Verbeke (Verbeke 1964: 17, 18, 20, 24): a — albiseta; b — turcestanica; c — lineata; d — rossica

Рис. 2. Стернит 4 самцов Ilione по Verbeke (Verbeke 1964: 17, 18, 20, 24): a — albiseta; b — turcestanica; c — lineata; d — rossica

We had some difficulties in identifying Ilione, therefore we decided that it would be useful to make a new review of the genus. While working on the review we found significant variability of non-genitalic characters, so to compose an identification key was not an easy task. Another problem was the lack of Western European material. After some hesitation we decided to divide the present publication in two parts. Part 1 is devoted to the five common species of Ilione of which we examined a representative material, namely:

Ilione albiseta Scopoli, 1763;

I. lineata Fallen, 1820;

I. rossica Mayer, 1953;

I. trifaria Loew, 1847;

I. turcestanica Hendel, 1903.

These five species cover about 99% of Ilione specimens in insect collections. We offer new illustrations and a revised identification key for them.

Part 2 is devoted to the three rare sub-Mediterranean species and one newly described Indian species, namely:

I. unipunctata Macquart, 1849, examined;

I. corcyrensis Verbeke, 1964, 1$ examined;

I. truquii Rondani, 1863, no specimens examined;

Ilione bindata sp. nov., described from a single male.

In each case we offer the description of available specimens and our taxonomic comments.

Fig. 3. Male terminalia of Ilione from RozkoSny (RozkoSny 1987: 222-224): a — albiseta; b — lineata; c — rossica

Рис. 3. Терминалии самцов Ilione по RozkoSny (RozkoSny 1987: 222-224): a — albiseta; b — lineata; c — rossica

Material and methods

Localities are given as follows: country, region/state/province (in italics), and geographical coordinates in decimal-degree format. The full names of regions of Russian administrative subdivisions are an entangled result of political and historical events of no interest for zoology, so they are listed as a name and the word 'region.

Illustrations are original unless otherwise credited. When referring to figures, to avoid confusion we capitalise the first letter (Fig. or Figs.) for those appearing in this paper and use lowercase (fig. or figs.) for those published elsewhere.

The specimens examined are deposited in the following museums:

ZIN — Zoological Institute, Saint Petersburg, Russia;

ZMUM — Zoological Museum of Moscow University, Russia (not indicated in the text).

The following generally accepted abbreviations for morphological structures are used: f1, t1, f2, t2, f3, t3 = fore-, mid- , hind- femur or tibia respectively; ac — acrostichal setae; dc — dorsocentral setae; prst — presutural; post — postsutural; a, p, d, v = anterior, posterior, dorsal, ventral seta(e). Other abbreviations: L. — lake; R. — river; Reg. — region; vill. — village.

Part I. Five common species of Ilione

Annotated list of examined material with distributional data, taxonomic remarks and identification key

Ilione albiseta Scopoli, 1763 Figs. 1a, 2a, 3a, 4, 12, 15, 19 Musca albiseta Scopoli, 1763 Musca crocus Harris, 1780 Musca aratoria Fabricius 1794 Tetanocera interstincta Fallen, 1820 Chione communis Robineau-Desvoidy, 1830 Chione sepedonoidea Robineau-Desvoidy, 1830 Material examined: AZERBAIJAN: 10 km N to Yashny [unclear locality], salt marshes, 26.05.1972, V. Rikhter, 1$; Astara [38.45°N, 48.85°E], 09.05.1911, L. Mlokosevich, 1$ (all ZIN); BELARUS, Brest Reg.: Pinsk u., Minsk. g., Gaidamashka [now Pinsk, « 52.12°N, 26.08°E], 02.06.1905: A. Bartenev, 2$, 1$; B. Gindtse, 1$; Zvanets Res., Novo-selki env., 52.066°N, 24.834°E, 21.06.2018, P. Prokhorchyk, A. Semeniak, 1$; ESTONIA, Saaremaa Isl., Viidumae Nature Reserve [58.31°N, 22.08°E], 21.09.1975, K. Gorod-kov, 2$, 2$ (ZIN); HUNGARY: 45 km S of Budapest, Domsod, Apajpuszta [47.09°N, 19.11°E], 25.06.1970, K. Gorodkov, 1$, 2$; Monor [47.38°N, 19.41°E], 1900, Schluter, 8$, 4$ (all ZIN); KAZAKHSTAN: Akmo-la Reg., Shortandy [51.687°N, 70.990°E],

Figs 4-6. Ilione: 4 — < albiseta; 5 — < trifaria (photo: Paco Moreno); 6 — prosternum of lineata

Рис. 4-6. Ilione: 4 — < albiseta; 5 — S trifaria (фото: Paco Moreno); 6 — простернум lineata

Figs. 7-8. Males Ilione: 7 — lineata; 8 — turcestanica Рис. 7-8. Самцы Ilione: 7 — lineata; 8 — turcestanica

24.06.1952, B. Kuzin, 1$; East-Kazakhstan Reg.: Altay Mts, Katon-Karagay env., middle course of Bukhtarma R., 2000 m [49.124°N, 85.972°E], 03.09.1970, V. Tanasiychuk, 1? (ZIN); Kyzylkum on Irtysh R., 30 km S of Sa-marskoe [48.72°N, 83.41°E], 03-04.08.1978, E. Narchuk, 1$ (ZIN); Karaganda Reg., Atasu [48.67°N, 71.62°E], 07.08.1959, Keleynikova, 1?; West Kazakhstan Reg. [Zelenovsky Dis-tr.], Yanvartsevo env., floodplain of Ural R. [51.44°N, 52.22°E], 18.07.1949, K. Romadina, 2$, 1? (ZIN); KYRGYZSTAN: Issyk-Kul Reg., S of Rybachje (= Balykchy), 1609 m, shore of

Issyk-Kul L. [42.43°N, 76.19°E], 16.08.1969, E. Narchuk, 2$, 3? (ZIN); Osh reg., Irkeshtam [39.68°N, 73.90°E], 20.05.1924, N. Filippov, 1$; 03.06.1924, N. Filippov; 1$. LITHUANIA, [70 km NW of] Kowno [now Kaunas], Georgenburg [now Jurbarkas, 55.08°N, 22.79°E], 15.06.1903, P. Winogradoff[-Niki-tin], (ZIN); MONGOLIA: Khovd aimak, ur. Elkhon, 20 km SE of Altay on Bodonch [« 45.684°N, 92.448°E], 27.07.1970, E. Narchuk, 1?; Dornogovi aimak, Barun-Bayan, 20 km ESE Tala-Khongoryn-Khuduk [unclear locality, presumably Airag, 45.8°N, 109.3°E],

Figs 9-10. Ilione, anepisternum: 9 — rossica, with weak setulae; 10 — turcestanica, with stronger setulae

Рис. 9-10. Ilione, анэпистернум: 9 — rossica, со слабыми щетинками; 10 — turcestanica, с сильными щетинками_

Figs 11-14. Ilione, hairing of arista: 11 — corcyrensis; 12 — albiseta; 13 — turcestanica; 14 — rossica

Рис. 11-14. Ilione, опушение аристы: 11 — corcyrensis; 12 — albiseta; 13 — turcestanica; 14 — rossica

29.06.1971: M. Kozlov, 2$; B. Kerzhner, 3$ (all ZIN); NETHERLANDS, Nijmegen, Ooi-jpolder [51.862°N, 5.908°E], AC 189-429, 08.06.2008, G. Pennards, 1?; Wageningen [51.957°N, 5.684°E]: 15.06.2001, G. Pennards, 2$; 09.06.2002, G. Pennards, 1?; RUSSIA: Krasnodar Reg.: Adler env., 43.4766°N, 39.9067°E, 29.10.2009, D. Gavryushin, 1?; Dakhovskaya env., valley of Belaya R., 44.199°N, 40.170°E, « 465 m, 18-31.08.2009, K. Tomkovich, 1$; Crimea Reg., Feodosia [45.1°N, 35.4°E], 25.08.1927, Wiolowitsh, 1$; Kursk Reg., Central Chernozemny NR, Psyol R. [51.191°N, 36.313°E], 21.07.2007, N. Vikhrev, 1?; Mordovia Reg., Smolny National Park,

16 km NE of Kemlya, 54.76°N, 45.47°E: 25.07.2021, G. Semishin, 1$; 15.08.2022, G. Semishin, 1?; Moscow Reg.: Golitsyno [55.62°N, 36.99°E], 20.08.1977, A. Shatalkin, 1?; Dmitrov Distr., Kostino env. [56.31°N, 37.75°E], 20-27.06.2007, N. Vikhrev, 3?, 1$; Kosino [now Moscow, 55.717°N, 37.950°E], 1926, B. Dodonov, 1?; Moscow, Izmaylovo [55.79°N, 37.78°E]: 28.05.1983, A. Ozerov, 1$; 05.06.1983, A. Ozerov, 1$; 18.06.1983, 1$; 02.07.1983, A. Ozerov, 1$; Naro-Fominsk [55.39°N, 36.73°E], 03.07.2007, D. Gavryushin, 1?; Shakhovskoy Distr., Stepankovo env., 55.996°N, 35.620°E, 03.06.2007, A. Ozerov, 2$, 2?; Serpukhov Distr., Zibrovo [54.846°N,

Figs 15-18. Sternite 4 of males Ilione: 15 — albiseta; 16 — turcestanica; 17 — lineata; 18 rossica

Рис. 15-18. Стернит 4 самцов Ilione: 15 — albiseta; 16 — turcestanica; 17 — lineata; 18 rossica

37.71°E], 31.05.1903, P. Datskevich, 1$; Novosibirsk reg., Zaeltsovsky Park, Ob R. left bank [55.046°N, 82.840°E], 15.09.2007, O. Kosterin, 2$, 1$; Orenburg Reg., Zhe-tykol L., 25 km NNE of Svetly vill. [51.045°N, 60.856°E], 10-12.08.2004: V. Krivokhat-sky, O. Ovchinnikova, 1$, 2$; E. Narchuk, 3$ (all ZIN); Ryazan Reg., Ryazan env., Khrapovo [54.60°N, 39.64°E], grove near a river bank, poplars, elms, alders, 06.07.1975, V. Kovalev, 1$; Samara Reg., Samara (53.24°N, 50.12°E), 13.08.1910, 2$; Samars-kiy u. [now Samara env., « 53.24°N, 50.12°E], 13.08.1910, V. Bostanzhoglo, 1$; Volgograd Reg.: Sarepta [48.518°N, 44.510°E], 1866, A. Bekker, 3$; 1868, A. Bekker, 3$; Tsatsa [48.20°N, 44.67°E], lake shore, 16.06.2005, E. Narchuk, 1$; Upper Balykley [49.53°N, 45.16°E], meadow, 09.08.2012, D. Astak-hov, 2$ (all ZIN); Voronezh Reg.: Khop-ersky Nat. Res., "Dubovaya Khata" cordon [51.24°N, 41.80°E], 06.08.2022, K. Tom-kovich, 1$; Ramon [51.916°N, 39.350°E], 12-15.09.1978, A. Shatalkin, 3$; Yakutia Reg.: Yakutsk [62.0°N, 129.73°E], V. Zlobin, 21.07.1987, 1$; Yakutsk env., Chochur-Mu-ran L., swampy shore [62.016°N, 129.605°E], 20.07.1974, K. Gorodkov, 2$, 1$; Turan-nakh, path towards Amga R. [61.99°N, 130.73°E], 18.07.1925, V. Bianki, 1$; Olyok-

minsk Distr.: Biryuk R. near mouth of Melichan R. [60.5°N, 119.4°E], 17.07.2008, A. Ovchinnikov, 1$, 1$; Olyokma mouth [60.371°N, 120.685°E], floodplain meadow, 03.08.1974, E. Narchuk, 1$; Olyokminsk airport [60.4°N, 120.5°E], 31.08.1988, K. Gorodkov, 1$ (all ZIN); Yaroslavl Reg., Berditsyno [57.46°N, 40.12°E], A. Yakov-leva: 01.07.1906, 1$; 29.08-08.09.1906, 8$, 2$; 01-05.10.1907, 2$, 1$ (all ZIN); TURKEY, Sakarya Reg., Karasu env. [41.08°N, 30.74°E], 27.08.2009, N. Vikhrev, 7$, 1$. UK: South East England Reg., Oxford, 05-20.09.1998, A. Ozerov & M. Krivoshei-na, 1$; UKRAINE: Odessa Reg., Odessa, 13.08.1978, Yu. Verves, 1$, 2$ (ZIN); Poltava Reg., Poltava env., 1894, V. N. Rodzy-anko, 1$; UZBEKISTAN, Samarkand Reg., Kattakurgan Distr., Kumak [39.99°N, 66.12°E], L. Zimin: 07-20.05.1929, 7$, 8$; 10.07.1929, 1$ (all ZIN).

Distribution. The Palaearctic, except for the Far East. Most common in Europe; Asian records are sporadic: Kazakhstan from west to east, Kyrgyzstan, Mongolia, Russia: Orenburg, Novosibirsk, and Yakutia regions (the easternmost record is at 130°E). Recorded south of 60°N and north of 40°N.

Was also reported by Li et al. from Chinese provinces Xinjiang and Inner Mongolia, but

Fig. 20. I. lineata, postabdomen: lateral, lateral, and semiventral Рис. 20. I. lineata, постабдомен: сбоку; сбоку; снизу и сбоку

without specifying the exact localities in these vast regions (Li et al. 2019).

Ilione lineata Fallen, 1820 Figs 1b, 2c, 3b, 6, 7, 17, 20

Material examined: BELARUS: Gomel Reg., Mozyr env., Knyaz L. [now Chervonoe L., 52.38°N, 28.045°E], 19.06.1905, B. Gindtse, 1$, 1$; Minsk Reg., Barisaw Distr., Velikaje Stachava env., 54.149°N, 28.636°E, 07.07.2013, D. Gavryushin, 1$; Vitebsk Reg., Lepel Distr., Kraytsy [54.67°N, 28.287°E], 09.08.1969, A. Antonova, 1$; ESTONIA, Peedu [59.19°N, 25.66°E], A. Stackelberg, 06-24.08.1951, 16$, 3$ (ZIN); KAZAKHSTAN, Akmola Reg., Borovoe [53.08°N, 70.32°E], 17-25.07.1932, V. Popov, 2$, 1$ (ZIN); RUSSIA: Arkhangelsk Reg.: Arkhangelsk, 64.546°N, 40.568°E, 04.08.2011, D. Gavryushin, 1$; Solvychegodsk, 61.342°N, 46.914°E, 13.08.2010, D. Gavryushin, 1$; Bashkortostan Reg., Abzakovo-Murakaevo env., Kryktytau Mts, steppe, birch [« 53.56°N, 58.39°E], 02-8.08.2008, K. Tom-kovich, 1$, 1$; Chelyabinsk Reg., near Zlatoust, Taganay Mts [« 55.25°N, 59.77°E], 1824.07.2008, K. Tomkovich, 1$; Karelia Reg., 30 km E of Chupa [66.3°N, 33.6°E], 23.07. 1975, K. Gorodkov, 2$, 1$ (ZIN); Mordovia Reg., Purdoshki env., 54.689°N, 43.533°E, 6.09.2020, N. Vikhrev, 1$; Moscow Reg.: Katuar [56.1°N, 37.5°E], 21.07.1954, B. Rodendorf, 1$; 29.08.1954, B. Rodendorf, 1$ (all ZIN); Dm-itrov Distr., Kostino env. [56.31°N, 37.75°E],

N. Vikhrev: 01.07.2007, 1$; 16.07.2007, 4$, 1$; 14.08.2007, 1$; Shakhovskoy Distr. Burtsevo env. [55.981°N, 35.607°E], 01.08.2007, A. Oze-rov, 1$; Odintsovo Distr., Golitsyno [55.62°N, 36.99°E], A. Shatalkin: 09.07.1977, 1$; 20.07.1977, 3$; 06-09.08.1977, 3$; 04.10.1981, 5$; 13.08.1983, 1$; 05.08.1988, 1$; Mamon-tovka [Pushkino, 56.00°N, 37.82°E], 05.07.1956, E. Smirnov, 1$; Stupino Distr., Sokolova Pustyn [54.842°N, 38.04°E], 04.07.1937, B. Rodendorf, 1$; Murmansk Reg., Khibiny railway station [67.67°N, 33.22°E], 27.08.1928, N. Cheburova, 10$, 3$ (ZIN); Saint Petersburg Reg., Luga distr., Yashchera [59.15°N, 29.91°E], A. Stackelberg: 09-27.08.1957, 14$, 3$; 27.08.1957, 1$; 26.08.1960, 1$ (all ZIN); Yaroslavl Reg., Koz-modemyansk [57.498°N, 39.694°E], 14.08.1977, A. Shatalkin, 2$, 1$.

Distribution. Common in Europe, present in the Urals. One Asian record only: Kazakhstan (70°E). In Eastern Europe known from cold (north to 67°N) and temperate (south till 52°N) latitudes.

Ilione rossica Mayer, 1953 Figs 1e, 2d, 3c, 9, 14, 18, 21 Material examined: BELARUS, Vitebsk Reg., Ambrosovichi [55.19°N, 29.64°E], 01.08.1924, Birulya, 1$ (ZIN); ESTONIA, Peedu [59.19°N, 25.66°E], A. Stackelberg, 23.08.1951, 2$ (ZIN); KAZAKHSTAN: North Kazakhstan Reg., Pe-tropavlovsk, 54.93°N, 69.22°E, 27.06.2015, O. Kosterin, 1$; Kokshetau Reg., Akmola R.

Fig. 21. I. rossica, lateral views: of separated surstylus from different angles, of dissected postabdomen, of bent postabdomen of an intact specimen

Рис. 21. I. rossica, вид сбоку: отделенный сурстиль под разными углами зрения; препарированный постабдомен; изогнутый постабдомен на интактном экземпляре

[53.26°N, 69.45°E], V. Popov, 3$, 6? (ZIN); West-Kazakhstan Reg., Solyanka R., left tributary of Ural R. [51.266°N, 52.348°E], 04.07.1949, L. Arnoldi, 1? (ZIN); RUSSIA: Bashkortostan Reg., Beloretsk Distr., Makhmutovo env., Belaya R., 550 m, 54.330°N, 58.807°E, 15.07.2015, D. Gavryushin, 2$; Chelyabinsk Reg., S Ural, Miass, SW Turgoyak L. [55.138°N, 60.032°E], 26-31.07.2008, K. Tomkovich, 1?; Ekaterinburg Reg., Uktus [56.76°N, 60.64°E], 15.06.1910, G. Yakobson, 1$ (ZIN); Irkutsk Reg. [Ust-Udin Distr.], Molka [53.91°N, 103.35°E], 29.06.1931, I. Rubtsov, 1$ (ZIN); Karelia Reg., 30 km E to Chupa, Medvejia Guba [66.26°N, 33.73°E], 23.07.1975, K. Gorodkov, 1$ (ZIN); Kemerovo Reg., Tisul [55.75°N, 88.28°E], 16.06.1911, A. Gorchakovsky, 1$ (ZIN); Mordovia Reg.: Smolny National Park, 16 km NE of Kemlya, 54.76°N, 45.47°E: 26.07.2021, G. Semishin, 1$; 07-15.08.2022, G. Semishin, 1$, 3?; Alatyr, 54.740°N, 45.377°E, 14.07.2022, K. Tomkovich, 1?; Moscow Reg.: Kalistovo, 08.07.1935, N. Filippov, 1?; [Odintsovo Distr.] Golitsyno [55.62°N, 36.99°E], A. Shatalkin: 04.10.1981, 3$; 26.06.1982, 1$, 2?; 09.07.1983, 1$; Novosibirsk Reg., Istikim, Koynikha R. [54.57°N, 83.26°E], 13.07.1977, K. Gorodkov, 6$, 2? (ZIN); Omsk Reg., Omsk, left bank of Irtysh R.

[55.03°N, 73.19°E], 22.07.1912, N. Vydrina, 1$; Severnaya, near Rybnoe L. [54.96°N, 71.90°E], 22.07.1922, A. Reykhardt, 1? (all ZIN); Orenburg Reg.: Troitsk [50.69°N, 54.62°E], 05.07.1902, Ya. P. Shchelkanovtsev, 1?; 25 km N of Svetly, Zhetykol L. [51.1°N, 60.9°E], 10-12.08.2004, E. Narchuk, 1$ (ZIN); Perm Reg., Kishert [(now Ust-Kishert, 57.375°N, 57.235°E], swampy shore ofSylva R., 27.07.1997, K. Gorodkov, 1$ (ZIN); Ryazan Reg., Ranova R., 53.72°N, 39.93°E, 06.06.2012, N. Vikhrev, 2$; Samara Reg., Samara, 13.08.1910, 1?; Saratov Reg., Dyakovka [50.72°N, 46.78°E]: 23-24.06.2003, V. Krivokhatsky, O. Ovchin-nikova, 1$, 2?; 27.06.2012, D. Astakhov, 2? (all ZIN); Saint-Petersburg Reg., Luga Distr., Yashchera [59.15°N, 29.91°E], A. Stackelberg: 06-15.08.1959, 15$, 4?; 29.08.1966, 1$; 09.08.1967, 1$, 1? (all ZIN); Volgograd Reg., Sarepta [48.518°N, 44.510°E], 1868, A. Bek-ker, 2$ (ZIN); Voronezh Reg.: Borisoglebsk env., Tellerman forestry [51.353°N, 42.037°E], 03.08.1964, V. Kovalev, 1?; Ramon [51.916°N, 39.350°E], 12-15.09.1978, A. Shatalkin, 6$, 2?; Yakutia Reg.: Olyokminsk Distr.: Biry-uk R. near mouth of Melichan R. [60.5°N, 119.4°E], 15-17.07.2008, A. Ovchinnikov, 2?; Olyokma mouth, steppe slope [60.371°N,

Fig. 22. Postabdomen of I. trifaria: ventral, semi-lateral and lateral Рис. 22. Постабдомен I. trifaria: снизу; снизу и сбоку; снизу

120.685°E], 03.08.1974, E. Narchuk, 1?; Yakutsk [62.0°N, 129.7°E]: 08.09.1927, I. Moskvin, 1$; 03.08.2008, A. Ovchinnikov, 1$; Olyokminsk airport [60.4°N, 120.5°E], 31.08.1988, K. Gorodkov, 2$ (all ZIN); Yaroslavl Reg., Berditsyno [57.46°N, 40.12°E], A. Yakovlev: 01-5.07.1906, 3?; 05-08.09.1906, 1?; 17.06.1907, 1?; 13.06.1908, 1? (all ZIN); UKRAINE: Poltava Reg. [Shishatsky Distr.], Yareski [49.85°N, 33.91°E], 05.08.1928, L. Gildebrandt, 1$, 4? (ZIN).

Distribution. From E Europe to E Siberia recorded from 49°N to 66°N.

Ilione trifaria Loew, 1847 Figs 1c, 5, 22, 23 Tetanocera trifaria Loew, 1847 Elgiva lateritia Rondani, 1863 Elgiva trivittata Strobl, 1899

Material examined: FRANCE, Var depart, Hyeres [43.1°N, 6.1°E], E. Abeille de Per-in, 2$, 1 ? (Museum National d'Histoire Naturelle, Paris); MOROCCO, Azrou, 33.43°N, 5.215°W, 1400 m, 18.05.2021, O. Kosterin, 1$, 1?.

Distribution. Africa, Maghreb and S Europe, to the east till Serbia.

Ilione turcestanica Hendel, 1903 Figs 1d, 2b, 8, 10, 13, 16, 24, 29

Material examined: ARMENIA, Para-kar [40.17°N, 44.41°E, 890 m], 09.10.1963, V. Rikhter, 12$, 2$ (ZIN); IRAN: Markazi Prov., Arak env. [34.03°N, 49.75°E], 2000 m, 18.05.2017, O. Kosterin, 2$; Tegeran, N. V. Bogoyavlen[sky], 1$; KYRGYZSTAN: Issyk-Kul Reg., Choktal env., 42.58°N, 76.75°E, 1600 m, 19-22.09.2013, N. Vikhrev, 1$; Osh Reg., Uzgen [40.8°N, 73.3°E], 2.07.1924, N. Filippov, 1$; RUSSIA: Dagestan Reg., 45 km S to Makhachkala [« 42.7°N, 47.7°E], 13.07.1983, E. Narchuk, 8$, 1$ (ZIN); Upper Gakvari vill. env., 42.54°N, 46.03°E, 2000 m, 06.08.2023, O. Kosterin, 1$; Gergebil Reservoir, 42.45°N, 47.03°E, 795 m, 12.09.2022, O. Kosterin, 1$; Samur Delta forest, 41.85°N, 48.55°E, 23-28.04.2023, N. Vikhrev, 1$; North Ossetia Reg., Alagir env. [43.030°N, 44.236°E], A. Ozerov: 02.07.1988, 1$; 18.05.1989, 1$; TADJIKISTAN: Stalinabad [now Dushanbe,

Fig. 23. I. trifaria: the same surstylus from different angles of view Рис. 23. I. trifaria: один и тот же сурстиль под разными углами зрения

Fig. 24. Postabdomen of I. turcestanica: ventral, semi-lateral and lateral Рис. 24. Постабдомен I. turcestanica: снизу; снизу и сбоку; сбоку

38.5°N, 68.8°E], Botanical Garden, A. Stackelberg: 06.05.1943, 8$, 3?; 28.12.1944, 3$; Du-shanbinka (= Varzob) R. valley, 06-13.05.1943, A. Stackelberg, 8$, 5? (all ZIN); Romit env., 38.709°N, 69.296°E, 1175 m, 14-16.06.2010, K. Tomkovich, 1?; TURKEY, Bolu Reg., 1200 m, Kibricik env., 40.42°N, 31.854°E, 01.09.2009, N. Vikhrev, 1?; UZBEKISTAN: Fergana Reg., Fergana, 24.08.1955, A. Zhelok-hovtsev, 1?; Shakhimardan [40.0°N, 71.8°E], 06.10.1955, V. Sychevskaya, 1?; Namangan Reg., Namangan, 02.06.1924, N. Filippov, 1?; Navoiy Reg., 25 km WSW of Langar [40.37°N, 65.73°E, 800 m], 27.05.1984, V. Tanasiychuk, 1? (ZIN); Samarkand Reg., Samarkand, 20-25.06.1927, N. Filippov, 2$; Tashkent, 22.04.1925, F. Dobzhansky, 4$, 3? (ZIN).

Distribution. From the Balkans to the Caucasus, Turkey, Iran, and Central Asia. A rather narrow range of latitudes from 43°N to 38°N. Was also reported from China, Xinjiang province (Li et al. 2019).

Discussion

1. Variability. The genus Ilione is characterised by a remarkably high level of intraspe-cific variability in all species. The wing pattern is a good example here. I. lineata usually has wings with characteristic brownish longitudinal stripes as shown in Fig. 7, but some specimens miss them. Similarly, I. rossica normally has hyaline wings, but some specimens have wings with brownish longitudinal stripes. I. albiseta has a very variable wing pattern in the median section of vein M1+2 (between

crossveins): there are 0 or 1 or 2 dark spots in this section. Moreover, these spots can be usual or they surround 1 or 2 short stump veins. Moreover, right and left wings of the same specimen are often quite different. Even the most reliable diagnostic character of I. al-biseta — the dark spot in the apical section of vein M1+2 — sometimes may be absent: one male among the examined material has this spot missing, although its identification is confirmed by the examination of genitalia.

Other important diagnostic characters like bare vs hairy prosternum or densely vs sparsely setulose anepimeron are sometimes difficult to apply because of wide variability in the number of setulae.

2. Genitalic characters. We have not found significant intraspecific variability of the structure of male genitalia, so the identification of the five above considered species based on the examination of a male postabdomen seems to us unmistakable. Rozkosny's drawings (shown here in Fig. 3) of the postabdomen of three Fennoscandian species (I. albiseta, I. lineata, I. rossica) are easily recognizable and correspond to our photographs (Rozkosny 1987: 222-224). Verbeke's drawings of the postabdomen (shown here in Fig. 1) are less understandable but still useful (Verbeke 1964: 9-16). Verbeke gave two drawings of the surstyli of I. rossica (Fig. 1e), which he indicated as a typical and an atypical form. We have quite a different interpretation of that: Fig. 21 shows that the same surstylus of I. rossica may look 'typical' or 'atypical' depending on the angle of view.

The surstylus of I. trifaria shows even more significant differences depending on the angle of view, as shown in Fig. 23. Once again, it is important to pay attention of colleagues that 3-dimensional sclerites may look very different in 2-dimensional projections (Vikhrev, Yanbulat 2019; Vikhrev 2022; 2023). In the present paper we tried to provide several projections for each illustrated postabdomen.

The shape of sternite 4 is also useful: our images (Figs 15-18) are similar to those from Verbeke (shown here on Fig. 2), except for sternite 4 of I. lineata which is actually much more elongated than in Verbeke's drawing (Verbeke 1964: 17-20). Sternites 4 of I. albi-seta and I. trifaria + I. turcestanica are usually well recognizable in intact specimens.

Key to common Palaearctic species of II-ione,

1. Vein M1+2 with a round spot in apical section (Fig. 4). Body length 6.5-9.0 mm. Presternum bare. surstyli in basal half wide and covered with spinulose setulae, in apical half surstyli narrow and hooked at very apex (Figs 1a, 3a, 19); sternite 4 at posterior margin with a pair of hill-like protrusions (Figs 2a, 15). 9: f3 ventrally without rows of spinulose pv and av setae in apical part, usually with 1-3 irregularly placed pv setae near middle. (Arista with short, dense, white hairs, total width of hairing at most as wide as basal width of arista (Fig. 12). Vein M1+2 with 1(0-2) dark spot(s) in section between crossveins. Inner margin of hind coxa with about 4 setulae, of which one markedly longer and stronger than the others. Anepisternum and propleuron in upper third with subglossy brown stripe. Prescutellar ac and subalar setae strong. Tergite 5 with 1-3 pairs of strong latero-

marginal setae.).........albiseta Scopoli

— Vein M1+2 without dark spot in apical section. Body length 5.0-7.5 mm. termi-nalia not as described above. Prosternum hairy or bare. $: f3 in apical third to half with two rows of av and pv spinulose setae or f3 without any ventral setae (I. lineata) ....................................... 2

2. Prosternum densely haired, with 4-12 pairs of hairs (Fig. 6). Anepisternum and propleuron in upper third with subglossy brown stripe (Fig. 7). Pleura with fine, sparse setulae (Fig. 9), usually 5-15 anepis-ternal setulae; anepimeron with 0-4 setu-lae. Arista with short, dense, white hairs, total width of hairing at most as wide as basal width of arista (Fig. 14). Inner posterior margin of hind coxa with minute setulae. sternite 4 posteriorly without rabbit-ears-like processes...............3

— Prosternum usually bare, in about one third specimens with 1-2 pairs of hairs. Upper part of anepisternum and propleu-ron not glossy, at most slightly darker than lower part (Fig. 8). Pleura densely covered with longer and stronger setulae (Fig. 10): 20-35 anepisternal setulae, 5-14 setulae on anepimeron. Arista with longer, sparse, mostly or partly brownish hairs, total width of hairing wider than basal width of arista (Fig. 13). Inner posterior margin of hind coxa with longer setulae, one of which is stronger and longer than the others. ster-nite 4 posteriorly with a pair of processes which look as hairy rabbit ears (Figs 2b, 16) ....................................... 4

3. Prescutellar ac setae distinct. Subalar setae long and strong. Prosternum haired on entire surface (Fig. 6). Wing usually with brownish longitudinal stripes (as in Fig. 7).

cerci swollen and exposed, usually distinct on intact specimen as in Fig. 7; sur-styli in the form of knives, deeply hidden under cerci (Figs 3b, 20); sternite 4 elongated (Fig. 17). $: v surface off3 without setae ...........................lineata Fallen

— Prescutellar ac setae absent. Subalar setae minute. Lower 1/3 of prosternum bare. Wing usually hyaline. cerci not enlarged; surstyli in basal half swollen, in apical half narrow and curved (Figs 3c, 21); sternite 4 not elongated (Figs 2d, 18). $: f3 in apical third with 2-5 av and 2-5 pv short spinulose setae.................rossica Mayer

4. Distributed in SW Palaearctic from Spain to Greece and in Maghreb in N Africa. Aristal hairs mainly white, dark at most

apical 2/5 (Fig. 5). Occiput with broad dark vitta. $: surstyli with wide basal half and

abruptly narrowed in apical half; this apical half directed ventrally, it looks straight or like bilobed horn depending on angle of view (Figs 1c, 5, 22, 23) — trifaria Loew — Distributed more easterly; in Asia from Turkey to E Kyrgyzstan (77°E); in Europe reported from the Balkans. Aristal hairs mainly black coloured, white hairs visible only at base (Figs 8, 13). Brown vitta on occiput blur if present. $: surstyli mostly evenly wide, only at very apex gradually narrowed (Figs 1d, 24, 29).................turcestanica Hendel

Part 2. Notes on the four rare species of Ilione

Three uncommon sub-Mediterranean species of Ilione and one newly described species from North India are considered here.

Ilione unipunctata Macquart, 1849 Figs 1h, 25

Ilione plumosula Becker, 1907 Material examined: SPAIN, Almeria Prov., Fuente de Cela, Tijola [37.37°N, 2.45° W], 13.03.1964, L. Knutson, 1$, identification label by J. Verbeke, 1966: Knutsonia unipunctata Macquart (ZIN).

Descriptive notes (based on a single examined $ specimen). Body length: 6.5 mm, wing: 5.5 mm. Head. Spots at bases of orbital setae and orbito-antennal spot distinct. Occiput with brown median stripe surrounded by whitish stripes. Antenna of a short type. Aristal hairing 2-2.5 times longer than width of aristal base; hairs brown, only in basal quarter white. Thorax. Prosternum with 2 pairs of hairs: on presternum and on upper margin of basisternum. Scutum with a pair of indistinct greyish submedian vit-tae. Anepisternum in upper third with brownish stripe (more distinct than in I. trifaria, less than in I. rossica). Prescutellar acrostichals strong, subalar strong. Anepisternum with 10/15 (left/ right) setulae, anepimeron with 4/5 setulae; setulae of 'strong type' (as in Fig. 10). Wing with costal spinules distinctly longer and stronger than in other Ilione (Fig. 25). Crossveins darkened, vein M1+2 with one spot between cross-veins. Dark longitudinal stripes in cells r4+5 and m12 moderately distinct. Legs. Inner posterior margin of hind coxa with 5-6 long setulae, one of them stronger than others. Chaetotaxy of f3 similar to that in I. albiseta: 10-11 pv in apical 2/3 and 8-9 av in apical half. Abdomen brownish-grey with 5 dark brown vittae: dorsal, lateral and ventro-lateral. Tergite 5 with 3-4

Fig. 25. I. unipunctata, male: general view; postabdomen; sternite 4 Рис. 25. I. unipunctata, самец: общий вид; постабдомен; стернит 4

pairs of strong latero-marginal setae. Sternite 4 similar to that of I. albiseta (Fig. 25). Surstyli characteristic, large and bilobed, one lobe wide and obtuse, the other lobe narrow, twisted, and with pointed apex (Fig. 25). The shape of sur-styli matches Verbeke's drawing reproduced in Fig. 1h here (Verbeke 1964).

Distribution. Reliably known from S Europe from Spain to Italy and Maghreb from Morocco to Tunisia.

Discussion. The general habitus of the examined specimen is quite similar to I. albiseta, but without a dark spot in the apical third of vein M1+2 The examined male also has a characteristic shape of the surstyli and several non-genitalic diagnostic characters: long costal spines on wing; aristal hairs long and sparse, mostly brown, only in basal quarter white; prosternum with two pairs of setulae. We are sure that it is a valid species and did not include it in the key because neither female nor variability are known to us.

Ilione corcyrensis Verbeke, 1964 Figs 1g, 11, 26

Material examined: GREECE, Corfu, Gardiki [39.476°N, 19.885°E], L. V. Knutson, 29.04.1963, paratype, J. Verbeke det., 1966, Knutsonia corcyrensis, 1$ (ZIN).

Descriptive notes (based on a single examined $ paratype). Body length: 7.5 mm, wing:

7 mm. Head. Spots at bases of orbital setae and orbito-antennal spot distinct. Occiput with a brown median stripe surrounded by whitish stripes. Antenna of intermediate length. Aristal hairing about 3 times longer than width of aristal base; hairs entirely white. Thorax. Prosternum bare. Scutum with a pair of indistinct greyish submedian vittae. Anepisternum and propleuron in upper third with subglossy brown stripe. Pres-cutellar acrostichals strong, subalar strong. Ane-pisternum with 22/24 (left/right) setulae (setulae on posterior margin rather strong, on upper margin weak), anepimeron with 3/4 setulae. Wing with costal spinules slightly longer than in other Ilione. Crossveins darkened, vein Mx+2 with one spot between crossveins. Dark longitudinal medial stripes in cells r4+5 and mx+2 indistinct. Legs. Inner posterior margin of hind coxa with 8 long setulae. Ventral surface off3 without strong setae. Abdomen yellow-brown with dark brown dorsal vitta. Tergite 5 with distinct marginal setae.

Distribution. So far known only from the island of Corfu.

Discussion. The general habitus of the examined specimen is quite similar to I. albiseta, without a dark spot in apical third of vein Mx+2 The examined female also has remarkably long and entirely white aristal hairs. Male surstyli are known only from the original description of Verbeke's

Figs 26-27. Ilione, general view: 26 — female paratype corcyrensis; 27 — male holotype I. bindata sp. nov.

Рис. 26-27. Ilione, общий вид: 26 — самка corcyrensis, паратип; 27 — самец bindata sp. nov., голотип

drawing (see Fig. 1g here), they look rather similar to that of I. albiseta (Verbeke 1964). To obtain a reasonable conclusion on validity, it is necessary to study more material, at least a male specimen.

Ilione truquii Rondani, 1863

Fig. 1f

This species was characterised by Verbeke mainly by negative characters: wing without dark spot in apical third of vein M1+2 (not I. albiseta) and without long costal spines (not I. unipunctata); aristal hairs not entirely white (not I. corcyrensis) and not short and dense as in I. albiseta (Verbeke 1964).

Distribution. So far known only from Syria.

Discussion. I. truquii was described from a female holotype from Syria. Verbeke has seen several more specimens again from Syria and his drawing of genitalia is based on the examination of these additional topotypes. Verbeke gave no verbal explanation as to which differences of the male genitalia are diagnostic (Verbeke 1964). Looking on Verbeke's drawing (Figs 1f, 1g, 1h) we can see that the surstyli of I. truquii are not similar to distinctive surstyli of I. unipunctata, but quite similar to those of I. corcyrensis. At the present political situation, it is unlikely that new Syrian material will be available. So far, we regard I. truquii as a doubtful species.

Ilione bindata sp. nov.

Figs 27, 28

https://zoobank.org/References/59BFC186-A536-4037-9CFA-47BD310C5A2D

Holotype: INDIA, Uttarakhand state, Ut-tarkashi, 30.735°N, 78.458°E, 1200 m, walnut-

oak forest, 01-02.05.2012, K. Tomkovich, 1$, abdomen dissected, stored in ZMUM.

Description. Body length: 5.5 mm, wing: 5.5 mm. Head. Spots at bases of anterior orbital setae and orbito-antennal spot large, black, very distinct. Occiput with brown median stripe surrounded by whitish stripes. An-tennal postpedicel darkened. Aristal hairing 2 times longer than width of aristal base; hairs white. Face with drop-like black spot (Fig. 27). Thorax. Prosternum with 2/3 setulae. Disc of scutum brownish, with a pair of greyish sub-median vittae. Brownish stripe on upper third of anepisternum hardly distinct. Thoracic chaetotaxy: 0+2 dc; prescutular acrostichals strong; 1 pospronotal seta; 2 notopleural setae; 1 presutural seta; 1 subalar seta; 2 postalar setae; 2 weak subalar setae. Anepisternum with 29 setulae, anepimeron with 9 setulae; meron with 3 minute setulae near posterior margin. Wing with costal margin darkened; vein M1+2 with darkened short 3/4 stump veins in section between crossveins (Fig. 27). Legs. Inner posterior margin of hind coxa with 8-10 short setulae. f3 with 7-8 pv in apical 1/2 and 5 av in apical 1/3. Abdomen brownish-grey, with distinct dark brown wide dorsal vitta and a pair of less distinct narrow lateral vittae. Tergite 5 without strong latero-marginal setae. Sternite 4 as in I. turcestanica. Surstyli similar to those in I. turcestanica, but narrower and medially curved in lateral view (Fig. 28).

Etymology. The species name is a Latinised adjective derived from 'bindi' — a coloured spot on a forehead often seen in India as a sign

Figs 28-29. Ilione, male postabdomen: 28 — bindata sp. nov., ventral and lateral; 29 — turcestanica, lateral

Рис. 28-29. Ilione, постабдомен самцов: 28 — bindata sp. nov., снизу и сбоку; 29 — turcestanica, сбоку

of belonging to the Brahmin varna. The characteristic spot on the face of the new species from India strongly resembles the bindi. In Latin 'bindata' means 'with a bindi.

Diagnosis. Ilione bindata sp. nov. is closely related to I. turcestanica, these species differ as follows

— Face with drop-like black spot (Fig. 27). Aristal hairs entirely white. Surstyli in lateral view curved at middle and narrower (Fig. 28). Costal margin of wing darkened (Fig. 27).......bindata sp. nov.

— Face without black spot. Aristal hairs mainly dark coloured, white hairs visible only at base (Fig. 13). Surstyli in lateral view not curved and wider (Figs 24, 29). Costal margin of wing not darkened . . . .

.................... turcestanica Hendel

Acknowledgements We thank Olga Ovchinnikova and Galina Suleymanova (Saint Petersburg) for the opportunity to examine the important material in ZIN. We thank Oleg Kosterin (Novosibirsk) for critically reading the manuscript.

References

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For citation: Vikhrev, N. E., Yanbulat, M. O. (2024) Review of the genus Ilione (Diptera, Sciomyzidae). Amurian Zoological Journal, vol. XVI, no. 2, pp. 310-325. https://www.doi.org/10.33910/2686-9519-2024-16-2-310-325

Received 20 February 2024; reviewed 20 March 2024; accepted 11 April 2024.

Для цитирования: Вихрев, Н. Е., Янбулат, М. О. (2024) Обзор рода Ilione (Diptera, Sciomyzidae). Амурский зоологический журнал, т. XVI, № 2, с. 310-325. https://www.doi.org/10.33910/2686-9519-2024-16-2-310-325 Получена 20 февраля 2024; прошла рецензирование 20 марта 2024; принята 11 апреля 2024.