Phaonia latipalpis SPECIES GROUP (DIPTERA, MUSCIDAE): REVIEW OF FAUNA OF RUSSIA AND SHORT NOTES ON FAUNAS OF ADJACENT TERRITORIES Текст научной статьи по специальности «Биологические науки»

Phaonia latipalpis SPECIES GROUP (DIPTERA, MUSCIDAE): REVIEW OF FAUNA OF RUSSIA AND SHORT NOTES ON FAUNAS OF ADJACENT TERRITORIES

N.E. Vikhrev, E.A. Erofeeva

Zoological Museum of Moscow University, Russia e-mail: nikita6510@yandex.ru

Received: 27.12.2022. Revised: 30.01.2023. Accepted: 13.02.2023.

The Phaonia latipalpis species group (Diptera, Muscidae) is reviewed. Some problems on taxonomy of the P. latipalpis group are clarified. Based on an examination of more than 400 specimens from several remote localities, the variability of P. zugmayeriae is studied, such an approach may turn out to be useful for other taxa of Phaoninae. The faunas of the P. latipalpis group of the West Palaearctic, Russian Far East and the Kuril Islands are considered separately, identification keys for each of these territories are offered. The faunas of the P. latipalpis group of China, Japan and the Nearctic region are briefly considered. One new synonym: P. apicata Johannsen, 1916 = P. solitaria Stein, 1920; syn. nov., and two new statuses: P. ommatina Zinovjev, 1981 to P. angustifrons ommatina Zinovjev, 1981 stat. nov.; P. apicata Johannsen, 1916 to P. apicalis apicata Johannsen, 1916; stat. nov. are proposed. Key words: Diptera, Muscidae, Phaonia latipalpis group, new records

https://dx.doi.org/10.24412/cl-31646-2686-7117-2023-32-159-180

Introduction

In present paper, we would try to clarify taxonomy of several Eurasian species of Phaonia Robineau-Desvoidy, 1830 with scutellum partly or entirely yellow. The considered species can be characterized as follows:

- Scutellum partly or entirely translucent or amber-like yellow (not opaque yellowish under whitish dusting as in P. trimaculata or P. errans);

- Abdomen entirely dark (not yellow as in species placed by Hennig in the P. pallida group which was recently reviewed by Vikhrev & Erofeeva (2018);

- Notopleuron entirely bare (except for P. asierrans and partly

P. zugmayeriae);

- Prealar seta two times longer than posterior notopleural; posterior notopleural seta 0.5-0.75x as long as anterior one (except for P. zugmayeriae and Nearctic P. bysia).

The following species of Phaonia known from Russia meet the above criteria: P. apicalis Stein, 1915 P. angustifrons Shinonaga & Kano, 1971 P. asierrans Zinovjev, 1981

P. latipalpis Schnabl, 1911 (= P. umbraticola Fonseca, 1957) P. ommatina Zinovjev, 1981 P. zugmayeriae Schnabl, 1888

The oldest valid species name among Palaearctic fauna is P. zugmayeriae, however, it is not easy to choose the name for this species-group. Hennig (1963) placed P. umbraticola and P. zugmayeriae in the P. zugmayeriae group. A set of Phaonia species known from Russia which was very similar to that given above was proposed by Zinovjev (1981a) as the P. umbraticola group. In the latest publications (mostly Chinese and Japanese ones), the name the P. latipalpis group is used, so we will name it in order to minimize the number of terms used.

In this paper, we do not focus on how our species set corresponds to the real and as yet unknown intraspecific phylogeny of Phaonia. Our main task is to delimit the described taxa more clearly and help with their identifications. In order to better address the task at hand, we will look at the regional faunas separately, so the article is divided into several chapters:

I-1. The Western and Central Palaearctic.

I-2 The Russian Far East (excluding the Kuril Islands).

I-3 The Kuril Islands.

II. Short notes on faunas of the P. latipalpis group of China, Japan and the Nearctic region.

Material and Methods

The specimens examined in this study are deposited in the following institutions:

MNHN - Muséum national d'Histoire naturelle, Paris, France;

ZIN - Zoological Institute, Saint Petersburg, Russia;

ZMUM - Zoological Museum of Moscow University, Russia.

Geographical coordinates are given in the Decimal Degrees format.

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).

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 name (taken from the English version of Wikipedia) and word "region" (abbreviated in the text - "reg.").

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

Results and Discussion I-1. The Western and Central Palaearctic

The fauna includes three species and one form of uncertain taxonomic status. The Western and Central Palaearctic, as reviewed here, cover all of Europe, the Caucasus and Siberia till the Yenisey River to the east. The distribution of

P. zugmayeriae is within the boundaries proposed here, while both P. apicalis and P. latipalpis have ranges extended to the Far East. For the last two species, we present here the examined material, including specimens from the Far East but excluding those from the Kuril Islands.

MATERIAL EXAMINED

Phaonia apicalis Stein, 1915 (Fig. 1, Fig. 2)

RUSSIA: Bashkortostan reg., 10 km SE of Beloretsk, 850 m, 53.89° N, 58.5° E, 15-19.06.2020, N. Vikhrev, 1 $ (ZMUM);

Irkutsk reg., Slyudyanka, 51.68° N, 103.69° E, 480 m, 27-29.06.2021, N. Vikhrev, 2 $$ (ZMUM);

Kamchatka reg., 60 km N Milkovo, (55.1° N, 159.1° E), 28-30.07.1978, A. Zinovjev, 2 $$; 47 km E Esso, (55.8° N, 159.4° E), 6.08.1978, A. Zinovjev, 1 $; Koryaki env., (53.25° N, 158.15° E), 21.07.1978, A. Zinovjev, 1 $ (all ZIN);

Khabarovsk reg., 15 km N Bikin, (46.9° N, 134.3° E), 1.06.1983, A. Zinovjev, 2SS, 1 $ (ZIN); Bolshekhekhtsirsky NP, (48.26° N, 134.77° E), 2.07.1982, A. Zinovjev, 2 $$ (ZIN); Solnechny env, 50.72° N, 136.67° E, 17-19.6.2022, N. Vikhrev, 1 & 2 $$ (ZMUM); Nizhnyaya Manoma, 49.33° N, 136.61° E, 22.06.2022, N. Vikhrev, 1 $ (ZMUM);

Khanty-Mansy reg., E Ural, Khulga R., 65.151° N, 62.110° E, 13-16.07.2018, K. Tomkovich, 1 $;

Krasnoyarsk reg., Krasnoyarsk env., E bank, Stolby, 55.963° N, 92.745° E, YPT, 18-19.06.2011, K. Tomkovich, 2 $$ (ZMUM);

Mordovia reg., Smolny National Park (16 km NE of Kemlyaj, 54.76° N, 45.47° E, 17.09.2019, G.Semishin, 1 $;

Moscow reg., Moskovskiy env., 55.58° N, 37.33° E, YPT, 22.06.2015, K. Tomkovich, 1 $; Ruza env., 55.66° N, 36.05° E, E. Erofeeva, 21-31.05.2016, 1 $; 11-20.06.2017, 1 $; 21-30.06.2017, 1 & 1 $; 21-31.07.2017, 2 $$; 1-11.08.2017, 1 $; 11-20.08.2017, 1 $; 21-30.06.2020, 1 S (all ZMUM);

Perm reg., Kungur, Uchleskhoz (= presently abolished Preduralie National Park, 57.36° N, 57.14° E), A. Zinovjev, 24.06.1979, 1 $ (ZIN);

Primorsky reg., Kamenushka env.: (43.622° N, 132.232° E), 6.06.1979, A. Zinovjev, 2 $$; Vladivostok, Sedanka, (43.2° N, 132.0° e), 3.06.1979, A. Zinovjev, 1 S; Anisimovka env., (43.17° N, 132.79° E), 17.06.1979, A. Zinovjev, 1 S (all ZIN);

Saint Petersburg reg., Krasnitsa env., (59.45° N, 30.35° E), 8.06.1980, D. Kasparyan, 1 $ (ZIN);

Tuva reg., Uyuk R., 52.07° N, 94.04° E, 800 m, 1-3.07.2017, N. Vikhrev, 1 $;

Yaroslavl reg., Berdicino, (57.45° N, 40.10° e), 28.05.1906, A. Yakovlev, 1 $

(ZIN).

DISTRIBUTION. Northern and Central Europe (Pont, 2013), Siberia, Far East, not recorded for the Caucasus. The records from Alaska (Huckett, 1965) and Japan (Kato, 1936) requires confirmation.

Fig. 1, Fig. 2, Fig. 3, Fig. 4. 1-2: male P. apicalis: 1 - lateral view, 2 - dorsal view; 3-4: male P. latipalpis: 3 - lateral view, 4 - dorsal view.

Phaonia latipalpis Schnabl, 1911 (Fig. 3, Fig. 4) P. umbraticola Fonseca, 1957

RUSSIA: Altai Rep. reg., Ust-Sema env., 51.6° N, 85.8° E, 21-26.06.2016, N. Vikhrev, 1 $ (ZMUM); Altai between Ada and Matur Rivers (= 50 km E of Tashtagol, 52.7° N, 88.6° E), 11.07.1897, Yu. Wagner, 1 S (ZIN);

Chelyabinsk reg., Taganay Mts., (55.221° N, 59.734° E), 18-24.07.2008, K. Tomkovich, 1 $ (ZMUM);

Khakasia reg., Shira env., Beljo salt-lake, 54.65° N, 90.18° E, 382m, YPT,

I-3.07.2011, K. Tomkovich, 1 $; Kubayka, 52.33° N, 89.82° E, 620 m, 10-13.07.2017, N. Vikhrev, 1 $ (all ZMUM);

Krasnoyarsk reg., 35 km WWS of Krasnoyarsk, Shumiha R., 19.07.1958, G. Bey-Bienko, 1 $ (ZIN); Krasnoyarsk env., E bank, Stolby, 55.963° N, 92.738° E, 209-260 m, YPT, 30-31.07.2009, K. Tomkovich, 1 $ (ZMUM);

Moscow reg., Ruza env., 55.66° N, 36.05° E, E. Erofeeva, 1-11.06.2019, 1 $;

II-20.06.2020, 1 $ (all ZMUM);

Primorsky reg., Andreevka env., 42.7° N, 131.1° E, 26.07-3.08.2018, N. Vikhrev, 1 S, 1 $ (ZMUM);

Saint Petersburg reg., Tolmachevo env., (58.85° N, 29.9° E), 1.08.1935, A. Stackelberg, 1 S (ZIN).

DISTRIBUTION. Similar to that of P. apicalis: Northern and Central Europe (Pont, 2013), Siberia, Far East, not recorded for the Caucasus.

Phaonia zugmayeriae Schnabl, 1888 (Fig. 5, Fig. 6, Fig. 17)

AZERBAIJAN: unknown locality: alpine meadow at 1800 m, 4.08.1962, Zagulyaev, Pastuhov, 1 $ (ZIN);

CZECHIA: Marienbad, (Marianske Lazne, 50.0° N, 12.7° E), Kowarz, 1 S

(ZIN);

Fig. 5, Fig. 6. P. zugmayeriae: 5 - male (Slovakia, Mala Fatra, 49° N, 19° E, 750 m, 22.07.2012, photo Mucha Fero, diptera.info); 6 - our records of the species in Europe and Siberia.

GEORGIA: Kazbegi, (42.66° N, 44.65° E), 1810 m, 07.1983, A. Pont, 1 S

(ZIN);

RUSSIA: Altai Rep. reg., Seminsky Pass., 51.05° N, 85.59° E, 1650 m, 27-30.06.2016, N. Vikhrev, 5 SS; Turala R., 50.99° N, 85.68° E, 1350 m, 8-12.07.2016, N. Vikhrev, 1 S (all ZMUM); Altai between Ada and Matur Rivers (= 50 km E of Tashtagol, 52.7° N, 88.6° E), Yu. Wagner, 3-12.07.1897, 2 $$ (ZIN);

Bashkortostan reg., 10 km SE of Beloretsk, 53.89° N, 58.5° E, 850 m, 15-19.06.2020, N. Vikhrev, 2 SS, 1 $ (ZMUM);

Khanty-Mansy reg., E Ural, 63.818° N, 59.562° E, 1-8.07.2010, K. Tomkovich, 1 S, 1 $ (ZMUM);

Komi reg., Eletskaya, 67.042° N, 64.22° E, 9.07.2019, N. Vikhrev, 1 S (ZMUM); Krasnodar reg.: Psekhako Mt., 43.697° N, 40.367° E, 2000 m, 14-18.06.2008, K. Tomkovich, 4 SS; Lagonaki campsite env., 44.09° N, 40.02° E, 1700 m, 26-28.06.2009, K. Tomkovich, 4 SS; Lagonaki, 44.009° N, 39.994° E, 1500-1900 m, N. Vikhrev, 27-30.06.2011, 7 SS; 5-9.06.2015, 1 $ (all ZMUM);

Krasnoyarsk reg.: W of Krasnoyarsk, Kryuchkovo, 56.11° N, 92.13° E, 14-23.07.2009, K. Tomkovich, 1 Krasnoyarsk env., E bank of Yenisey R., Stolby, 55.963° N, 92.745° E, 209-260 m, 18-19.06.2011, K. Tomkovich, 1 Ergaki NP, 52.84° N, 93.25° E, 1450 m, 27-29.06.2017, N. Vikhrev, 12 SS (all ZMUM);

Moscow reg.: Ruza env., 55.66° N, 36.05° E, 21-30.06.2015, N. Vikhrev, 1 S; E. Erofeeva, 1-10.07.2016, 1 11-20.06.2017, 1 S, 1 ?; 21-30.06.2017, 3 SS;

I-11.07.2017, 3 21-31.05.2018, 27 SS, 1 ?; 1-10.06.2018, 27 SS, 8

II-20.06.2018, 31 SS, 12 21-30.06.2018, 1 1-11.07.2018, 5 SS, 5 1-10.08.2018, 1 21-31.08.2018, 4 1-10.09.2018, 1 21-31.05.2019, 16 SS, 1 ?; 1-11.06.2019, 12 SS, 4 11-20.08.2019, 3 21-31.08.2019, 5

1-10.06.2020, 49 SS, 34 11-20.06.2020, 18 SS, 44 21-30.06.2020, 4 SS, 7 1-10.07.2020, 5 Dmitrov distr., Kostino env., (56.31° N, 37.75° E),

N. Vikhrev, 24.05-2.06.2010, 2 SS; 1.06.2011, 1 S; Volokolamsk env., Lama R., 55.99° N, 36.00° E, 29-30.05.2019, M. Yanbulat, 1 S (all ZMUM);

Perm reg., Kungur, Uchleskhoz (= Preduralie NP, presently abolished, 57.36° N, 57.14° E), 9.07.1979, A. Zinovjev, 1 S (ZIN);

Saint Petersburg reg., st. Mozhayskaya, Voronja gora, (59.70° N, 30.13° E), 26.06.1980, A. Zinovjev, 2 SS (ZIN);

Yamalo-Nenets reg.: Sobj env., 67.06° N, 65.46° E, 26-31.07.2011, K. Tomkovich,1 Kharp env., 66.81° N, 65.78° E, 10-13.07.2019, N. Vikhrev, 2 SS (all ZMUM);

SERBIA: Babin Zub, 43.375° N, 22.625° E, 1550 m, 1-8.07.2015, N. Vikhrev, A. Ozerov, M. Krivosheina, 30 SS, 8 $$ (all ZMUM);

UKRAINE: Zakarpatsky reg., Kozmeschik, 10 km N of Goverla Mt., (48.21° N, 24.48° E), 28.05.1978, A. Zinovjev, 1 S (ZIN);

DISTRIBUTION. P. zugmayeriae is recorded from the majority of Western European countries (Pont, 2013). Data from Eastern Europe and Siberia were quite scarce: Russia: Perm region (Zinovjev, 1981c) and Khanty-Mansi region (Neroika Mt. env., 64.6° N, 59.6° E, in Ural Mts. in the border between Europe and Asia) (Malozemov, 1992). Our records show that it is distributed eastward to the Yenisey River (93° E). In the North, the range of P. zugmayeriae extends beyond the Arctic Circle (66.8° N). The southern limit of distribution is at about 56° N, in the lowlands and is extended southwards to 43° N, in the Balkan and Caucasus Mountains.

P. zugmayeriae Schnabl, 1888form with widenedfrons (Fig. 7, Fig. 8, Fig. 9)

RUSSIA: Dagestan reg., Hala-Hol Mts, (41.97° N, 46.37° E), 7.08.1913, L.F. Mlokosiewicz, 1 S (ZIN);

Krasnodar reg., Lagonaki campsite env., 44.09° N, 40.02° E, 1700 m, 26-28.06.2009, K. Tomkovich, 2 S; (Guzeripl env.), 44.009° N, 39.994° E, 1700 m, N Vikhrev, 27-30.06.2011, 1 S (all ZMUM).

REMARKS. These four males could have been described as a new species based on several differences from the typical form which are summarized in the identification key. However, we decided to list these specimens as a form of P. zugmayeriae with the uncertain taxonomic status because we have only few specimens from different locality each. As follows from the discussion below, only the study of large series would allow to come to reasonable taxonomic conclusions. Male genitalia were examined and were found the same as those of P. zugmayeriae.

DISCUSSION

The most important revision of Western Palaearctic Phaonia has been done by Hennig (1963), who established synonymies, gave redescriptions, discussed intraspecific variability and offered the first comprehensive identification key. Hennig also discussed the intraspecific variability of the three species of Phaonia treated here, although he did not include this information in the key. Apart from the

Fig. 7, Fig. 8, Fig. 9. P. zugmayeriae Schnabl, 1888 form with widenedfrons, male from Guzeripl: 7 - lateral view, 8 - abdomen, posterior view; 9 - head.

Hennig's key, the only original and very useful key for Phaonia was published by d'Assis-Fonseca (1968) for the British fauna. After publication of the Palaearctic Catalogue (Pont, 1986), knowledge of the nomenclature of Phaonia almost reached its current state.

The latest key for Central European Phaonia (Gregor et al., 2002) duplicates the Hennig's key excluding the non-European species, along with its errors and omissions: Recommendations of Gregor et al. (2002) are as follows: couplet 120) or 19(5). strongprst ac absent; notopleuron bare. ... apicalis Stein

- 1 pair of strong prst ac present; notopleuron bare (while in the descriptive notes indicated that notopleuron bare in P. latipalpis but haired in P. zugmayeriae (Gregor et al., 2002). ... 17/14

couplet 170) or 14(5):

-f1 dark brown to black; postpronotal calli yellow. ...zugmayeriae Schnabl

- f1 yellow; postpronotal calli black. ... latipalpis Schnabl Recommendations of d'Assis-Fonseca (1968) are as follows (55): Phaonia: prst ac absent; 3 post dc; scutellum at least partly yellow:

32 (33) scutellum entirely yellow. ... latipalpis Schnabl (as umbraticola)

33 (32) scutellum darkened at base. ... apicalis Stein

- prst ac present; abdomen dark; scutellum at least partly yellow; at least posterior femora yellow; usually 3 post dc:

56 (57) coxae and femora entirely yellow; postpronotal calli dark. ... latipalpis Schnabl (as umbraticola)

57 (56) fore coxa dark, f1 at least partly dark; postpronotal calli yellow. ... zugmayeriae Schnabl

These recommendations do not allow many specimens to be identified. For example, what about the not rare specimens with prst ac, in which both f1 and postpronotal calluses are either yellow or both dark? Or what about P. apicalis with a pair of strong prst ac setae?

We aimed to collect and study as many specimens from different regions as possible, investigate the variability, and then compile a key, taking it into account.

VARIABILITY

P. apicalis. According to Hennig (1963: 800), this species normally has no strong prst ac and 2 + 3 dc, but specimens with a pair of prst ac and 2 + 4 dc on one side of thorax were mentioned. Among our material (38 specimens - 7 SS, 31 strong prst ac present in 5 SS and 1 usually 2 + 3 dc, rarely 2 + 4 dc (2 specimens) or 2 + 3/4 (3 specimens). Female palpi are more distinctly dilated in the European specimens, less distinctly so in specimens from the Far East. One female from Moscow region has p seta on t1.

P. latipalpis. According to Hennig (1963: 800) the species has a pair of strong prst ac and 2 + 3 dc, except for a male from Saint Petersburg region with 2 + 4 dc. According to d'Assis-Fonseca's (1968) key in this species strong prst ac may be either present or absent. Our material (12 specimens - 3 SS, 9 shows that there is usually dc 2 + 3, rarely 2 + 4 dc (1 specimen); prst ac absent in all females and present in all males. Therefore, both the presence or absence of prst ac setae and number of post dc setae are variable characters and can hardly serve as diagnostic ones. Female palps are usually dilated, sometimes with normal width.

Thus, P. apicalis differs from P. latipalpis only in darker colouration off1 and scutellum (yellow only on apical part) and also in dark spiracles. Also P. apicalis has more slender body (Fig. 1), while the body of P. latipalpis is more robust (Fig. 3), but it is difficult to use as a diagnostic character.

P. zugmayeriae. In contrast with previous species, here we examined a large series of 423 specimens in total (Table 1), so our judgment on variability is much more reliable.

We can see that in all localities prst ac is almost always present in both sexes; notopleuron around strong anterior seta normally with hairs in males, bare or hairy in females; scutellum darkened basally in males, entirely or almost yellow in females. Other variable characters differ depending on region. In most localities specimens have 2 + 4 dc but in Moscow region the majority of specimens have 2 + 3 dc.

Table 1. Variability of P. zugmayeria

\Character Amount of speciments post dc strong ac f1 yellow f1 dark f1 dark ntp hairy ntp bare scutellum, % of

Region \ 3 4 3/4 0 + 1 1 + 1 1 + 2 pc yellow pc yellow pc dark yellow

Balkan 30 S 2 26 2 0 23 7 0 27 3 30 0 40-70

Mts

8 2 0 7 1 0 6 2 0 8 0 3 5 80-90

Altai- 18 S 3 12 3 1 15 2 0 15 3 14 4 30-50

Sayan Mts

4 2 2 2 0 1 3 0 4 0 0 0 4 80-100

N Ural 4 S 1 3 0 0 3 1 0 4 0 3 1 50-70

lowlands

2 2 0 2 0 0 2 0 0 2 0 0 2 80-100

Caucasus 16 s 0 15 1 0 14 2 10 3 3 15 1 30-80

Mts

2 2 1 1 0 1 1 0 2 0 0 1 1 100

Moscow 198S 124 41 33 0 151 47 1 9 188 194 4 40-75

reg.

141 2 105 12 24 2 122 17 42 99 0 29 112 90-100

total 423 S, 2

Note: Balkan Mts - Serbia, Stara Planina;

Altai-Sayan Mts - Altai Republic and Krasnoyarsk reg., Russia, S Siberia;

N Ural lowlands - lowlands near northern Ural Mountains, it includes Khanty-Mansi, Komi and Yamalo-Nenets reg. of Russia; Caucasus - Krasnodar reg, Russia and Georgia; Moscow region - NW part of the region.

Abbreviations used in Table 1: pc - postpronotal calli; ntp - notopleuron.

As Table 1 shows, the colouration generally accepted as typical "fore femur mostly dark, postpronotal calli yellow" (Hennig, 1963; d'Assis-Fonseca, 1968; Gregor et al., 2002) is mostly observed in specimens from the Balkan and Altai-Sayan Mts and from lowlands of the Polar Urals. Our series from the Caucasus has yellow f1 even in most male specimens. In our largest series from Moscow region, 95% of males have both f1 and postpronotal calli dark (126 of 188 such males have f2 and f3 dark), 30% of females have both f1 and postpronotal calli yellow, while only 5% of males and 70% of females have "typical" colouration. Therefore, the characters considered in Table 1 may be described as follows: prst ac present; 3 or 4 post dc; postpronotal calli yellow in females, yellow or dark in males; f1 dark or yellow; scutellum mostly yellow in females, partly yellow in males; notopleuron around strong anterior seta with hairs in males, bare or hairy in females.

Differences in the colouration of Caucasian specimens are not surprising, the Caucasus is a well-known hot spot of biodiversity. However, the fact that the series from Moscow region which is the center of the Eastern European lowlands shows the most deviating characters is surprising, we have no explanation for this.

Regular collecting in Moscow region also allowed us to estimate the year-to-year variability of the same population. We found that some characters may significantly vary, for example, in females collected in 2018-2019: 65% had f1 yellow and only 35% f1 dark; 55% had bare notopleuron. In 2020, the ratio of colouration of fore femur became reversed: 30% f1 yellow and 70% f1 dark; number of females with bare notopleuron increased to 80%.

P. zugmayeriae, form with widened frons. The narrow frons of a typical P. zugmayeriae is shown in Fig. 5 and Fig. 17, the typical distinctly widened frons of this form in Fig. 8. A male from Hala-Hol has f2 also mostly dark. The male from Lagonaki campsite has abdominal tergites 1 + 2, 3 and 4 yellow laterally, tergite 3 is also yellow on anterodorsal surface (Fig. 9). Two other males have abdomen dark in dorsal view; in lateral view tergite 3 is translucent yellow (Fig. 7). All males have scutellum completely yellow.

MALE GENITALIA

Examination of genitalia often helps to identify at least males. Gregor et al. (2002) provided drawings of terminalia of most European species of Phaonia (Fig. 10, Fig. 11, Fig. 12). The drawings are similar to each other, but differ in details. The question is whether these minor differences are helpful for diagnostic use. After re-examination of genitalia of our material we came to a conclusion that the diagnostic value of structure of genitalia seems doubtful for species described here. Even a slightly different angle of view on cercal plate of P. zugmayeriae (Fig. 13, Fig. 14) easily renders its shape similar to that in any illustration given by Gregor et al. (2002).

Fig. 10, Fig. 11, Fig. 12, Fig. 13, Fig. 14. Cercal plates of P. latipalpis group: 10 - P. apicalis; 11 - P. latipalpis; 12 - P. zugmayeriae; (10-12 from Gregor et al., 2002); 13-14 - P. zugmayeriae.

To sum up, high variability of characters was revealed for P. zugmayeriae. Despite this, a reliable identification of this species is possible, based on the set of characters recommended in the key below. At least we did not have a single questionable specimen among the 423 examined.

Previously proposed identification keys did not allow the identification of a significant proportion of specimens.

In contrast, the differences between P. apicalis and P. latipalpis are much less convincing. They are only based on the colouring of f1, spiracles and scutellum, darker in P. apicalis, and more yellow in P. latipalpis.

EXCLUDED SPECIES

Two species of Phaonia were placed in identification keys together with species considered here and differed from them only in details of colouration. We have not seen neither the type material nor specimens which presumably belong to these species, the data is taken from literature.

Phaonia szlenyii Mihalyi, 1974. Included in the key of Gregor et al. (2002). The species has short prealar seta (Mihalyi, 1974), it should be related to P. rufipalpis Macquart, 1835.

Phaonia tersa Villeneuve, 1936. Included in Hennig's (1963) key as similar to P. zugmayeriae. Doubtful taxon is described from a single female specimen. The study of new material is required to clarify the situation.

IDENTIFICATION KEY for Phaonia latipalpis group (WEST and CENTRAL PALAEARCTIC FAUNA), SS, ??.

Almost 500 yellow-scutellum specimens of Phaonia examined may be distinguished as recommended by the key without any exception.

SS, ??

1. Katepimeron with 1-6 distinct hairs (S - 2-6, ^ - usually 1-2 hairs). Prealar seta as long as posterior notopleural, posterior notopleural seta as long as anterior. Male eyes haired densely and long. Wings are intensively yellow at base. Notopleuron haired at anterior part in S, bare or haired in Strong prst ac present. (Thoracic spiracles: anterior - yellow, posterior - dark). ... 2

- Katepimeron bare. Prealar seta twice longer than posterior notopleural, posterior notopleural seta 0.5-0.75x as long as anterior one. Male eyes haired sparsely short. Wings are with even light-yellow tint. Notopleuron bare. Strong prst ac present or absent. ... 3

2. S: Fronto-orbital plates touch, frontal vitta is absent (Fig. 17). Parafacials are narrower than width of postpedicel. Cheek is narrow, 1/6 as wide as height of eye. Abdomen is entirely dark. Scutellum is partly yellow (30-80%). ... zugmayeriae Schnabl

- S ($ unknown): Fronto-orbital plates are separated by frontal vitta which is wider than diameter of anterior ocellus (Fig. 9). Parafacials is wider than width of postpedicel. Cheek is wide, about 1/3 as wide as height of eye or twice wider than postpedicel. At least abdominal tergite 3 laterally is translucent yellow (Fig. 7, Fig. 8) or abdomen is with extensive yellow pattern (Fig. 8). Scutellum is completely yellow. ... zugmayeriae form with widened frons

3. Fore femora dark. Anterior and posterior spiracles are dark. Scutellum is yellow only on apical part. Body is slenderer (Fig. 1). Mid tibia usually has 2 p seta. ... apicalis Stein

- Fore femora yellow, sometimes with dark dorsal streak. Anterior and posterior spiracles are yellow. Scutellum is entirely yellow. Body is more robust (Fig. 3). Mid tibia usually has 3 p seta. ... latipalpis Schnabl

I-2. The Russian Far East (excluding the Kuril Islands)

In Hennig's (1963) revision of Palaearctic Phaonia, there is no information on the Far Eastern fauna, since the material from there was not available in European collections (including ZIN, Saint Petersburg), except for Kato's (1936) record of P. apicalis for Japan. In 1971, Shinonaga & Kano (1971) published a large review of

the Japanese Muscidae including Phaonia. This is a rare book, but the descriptions given there are repeated verbatim in a later monograph (Shinonaga, 2003), which is much easier to find, and we will refer to it in the text. According to Shinonaga (2003), the Japanese fauna of the P. latipalpis group includes five species, of which one species, P. angustifrons Shinonaga & Kano, 1971 is recorded from Russia.

The fauna of Phaonia of the Russian Far East was studied by Alexey Zinovjev (1980, 1981a, 1981b) who newly described two species from the P. latipalpis group: P. asierrans Zinovjev, 1981 (we included this species in P. latipalpis group, majority of other authors do not) and P. ommatina Zinovjev, 1981 from the Kuril Islands (see chapter I-3). Zinovjev's papers were published in Russian and therefore are not well known to non-Russian dipterists.

The study of fauna of the P. latipalpis group in China began in 1986. Our short notes on the Chinese, Japanese and Nearctic species of the P. latipalpis group are given in chapter II.

We decided to divide the Far Eastern fauna of the P. latipalpis group into subregional faunas and discuss them separately. This chapter covers the Russian Far East mainland (Kamchatka, Khabarovsk, Primorsky regions and large Sakhalin Island) which, in our opinion, may be described satisfactorily. The faunas of the P. latipalpis group from the Kuril Islands, Japan and China cannot be satisfactorily described. These territories are shortly reviewed in chapters I-3 and II below.

MATERIAL EXAMINED

Phaonia apicalis Stein, 1915 (Fig. 1, Fig. 2)

Material examined: see chapter I-1.

Phaonia angustifrons Shinonaga & Kano, 1971 (Fig. 15, Fig. 18)

RUSSIA: Primorsky reg., Andreevka env., 42.7° N, 131.1° E, 26.07-3.08.2018, N. Vikhrev, 4 SS, 2 22 (ZMUM); Anisimovka env., (43.17° N, 132.79° E), 20.06.1982, A. Zinovjev, 1 2; Gorno-Tayozhnoe, (43.70° N, 132.16° E), 22.08.1962, E. Narchuk , 1 2; Kamenushka env., 30 km SE Ussuriyska, (43.622° N, 132.232° E), 7.06.1979, A. Zinovjev, 1 S, 3 22; Partizansk env., (43.15° N, 133.11° e), 27.08-14.09.1978, A. Zinovjev, 2 22; Vladivostok: Lesnaya Zaimka, (43.26° N, 132.10° E), 26.06.1982, A. Zinovjev, 1 2; Okeanskaya, (43.26° N, 132.04° E), 30.09.1978, A. Zinovjev, 1 2; Sedanka, (43.2° N, 132.0° E), A. Zinovjev, 27.08.1978, 1 2, 1-14.06.1979, 2 SS, 6 22; 25.07.1979, 1 S (all ZIN); Volchanets env., 42.908° N, 132.726° E, 1-4.08.2019, E. Erofeeva, 2 SS, 1 2 (ZMUM).

Sakhalin reg., Sakhalin Isl.: Poluostrov Terpeniya, (49.13° N, 144.25° E), 27.07.1956, N. Violovich, 1 S; Yuzhno-Sakhalinsk env.: (46.96° N, 142.76° e), N. Violovich, 29.06.1953, 1 2, 16.07.1955, 2 SS,1 2, 8.06.1956, 1 S; Mt. Chekhova, (46.99° N, 142.83° E), 5.06.1972, M. Kozlov, 1 S (all ZIN).

DISTRIBUTION. Japan and Russian Primorsky and Sakhalin regions. Probably also China (Jilin prov. as P. jilinensis and Liaoning prov. as P. riparia) (Ma et al., 2002).

Fig. 15, Fig. 16. Far Eastern species of the P. latipalpis group: 15 - male P. angustifrons Shinonaga & Kano, 1971; 16 - Holotype P. asierrans Zinovjev, 1981.

Phaonia asierrans Zinovjev, 1981 (Fig. 16)

Type material. Holotype, RUSSIA, Primorsky reg., Khasansky distr., Barabashevka River flood plain, 43.19° N, 131.50° E, 2.08.1978, D. Kasparyan (ZIN).

Paratypes, 2 1 CHINA, Sichuan prov., Tachienlu (= Dartsedo = Kangding, 30.05° N, 101.96° E), G. Potanin, 6.06.1893 (ZIN).

Other material. RUSSIA: Primorsky reg.: Vladivostok, Sedanka, 43.2° N, 132.0° E, A. Zinovjev, 19.07.1979, 1 (ZIN); Andreevka env.,42.64° N, 131.13° E, N. Vikhrev, 3-8.08.2018, 1 1 $ (ZMUM); Sakhalin reg., Kunashir Isl., Yuzhno-Kurilsk env., 44.03° N, 145.86° E, A. Zinovjev, 3.07.1979, 1 & (ZIN).

DISTRIBUTION. Known from Russian Primorsky and Sakhalin regions and China.

Phaonia latipalpis Schnabl, 1911 (Fig. 3, Fig. 4)

Material examined: see chapter I-1.

DISCUSSION

P. apicalis. In chapter I, we provided diagnosis of this species, the Far Eastern specimens of P. apicalis fit the diagnosis.

P. angustifrons.

DESCRIPTION. Male. Head is dark. Eyes have rather sparse hairs; facets are moderately enlarged. Fronto-orbital plates touch; frontal setae is only on lower third

of frons. Aristal hairs are 1.5x longer than width of postpedicel. Palpi is dark. Thorax is dark, with a pair of black submedian vittae. Scutellum yellow at apex. Mesonotum is with 2 + 3 dc, 1 + 1 ac, pra slightly longer than anterior notopleural seta, the latter is 2x longer than posterior one. Notopleura, meron and katepimeron are bare. Spiracles are brown. Legs are with coxae, throchanters, femora and tarsi dark, only tibiae yellow (f1 yellow only at very apex, f2 and f3 dark in basal 4/5-5/6 in males, 2/3-3/4 in females). t1 without submedian setae, t2 with 2-3 p, t3 with 4 short av, 2 ad and 1 pd. Abdomen is grey dusted with indistinct dark median vitta. Sternite is bare. Female differs as follows: frons wide, without crossed interfrontal setae; prst ac absent.

REMARKS. Among the fore species of the P. latipalpis group described by Shinonaga & Kano, this one is the least problematic due to dark femora. We agree that diagnosis based mostly on leg colouration is not very convincing, but in the Far East mainland we have not seen any specimens with intermediate colour of legs.

P. asierrans.

The species was described in Russian, here are descriptive notes in English.

DESCRIPTION. Male. Head is dark. Eyes have rather sparse hairs; facets are moderately enlarged. Fronto-orbital plates touch in middle; upper frontal setae weak though extend to level of anterior ocellus. Aristal hairs are longer than width of postpedicel. Palpi is dark. Thorax is dark, somewhat bluish, with a pair of black submedian vittae. Scutellum is yellow at apex (the Holotype) or entirely yellow (S, 2 from Andreevka). Mesonotum is with 2 + 4 dc, 0 + 1 ac, pra slightly longer than posterior notopleural seta, anterior notopleural distinctly longer than posterior one. Notopleura with 1 hair near anterior seta and 4-5 around posterior one. Meron are bare, katepimeron have 4-5 hairs. Anterior spiracle is white, posterior one is brownish-yellow. Legs are yellow including posterior coxae (Zinovjev, 1981): coxae dark), tarsi are black, f1 is yellow (the Holotype) or darkened in basal half (S from Andreevka). t1 is without submedian setae, t2 with 2 p, t3 with 2 av, 2 ad and 1 pd. Abdomen is grey dusted with indistinct dark median vitta. Sternite 1 have several hairs. Female differs as follows. Frons is wide, without interfrontals. Notopleura has only 1 hair placed near posterior seta. Hairs are on katepimeron and sternite 1 is weaker.

REMARKS. P. asierrans was not included in the P. latipalpis group neither by Zinovjev (1981a,b) nor by Chinese authors (Ma et al., 2002). Zinovjev (1981b) supposed that it is related to P. errans Miegen, 1826. According to Ma et al. (2002), it is related to Oriental P. simultans Malloch, 1931. We do not share these opinions, since at least P. asierrans differs from other species of the P. latipalpis group less than P. zugmayeriae.

IDENTIFICATION KEY FOR Phaonia latipalpis group (Russian Far East),

SS, 22.

1. Katepimeron have 4-5 hairs. Notopleuron have hairs around posterior seta. Strong prst ac is absent. Frontal setae extend to level of anterior ocellus. Sternite 1 setulose. Anterior spiracle is whitish. ...asierrans Zinovjev

- Katepimeron is bare. Notopleuron is bare. Strong prst ac usually present, rarely is absent. Frontal setae are present only on lower half of frons. Sternite 1 is bare. Anterior spiracle is yellow or dark. .. .2

2. Fore femur is yellow, at most with dark dorsal streak. Anterior and posterior spiracles are yellow. Scutellum is entirely yellow. ... latipalpis Schnabl

- At least fore femur is mostly dark. Anterior and posterior spiracles are dark. Scutellum is yellow only at apex. ... 3

3. f2, f3 are yellow. Aristal hairs are 0.5-0.8x as long as width of postpedicel. Strong prst ac usually present in S and is absent in 2. S: Facets of the eyes not enlarged. Mid tibia usually has 2p seta. ... apicalis Stein

- f2, f3 are dark. Aristal hairs are 1.3x as long as width of postpedicel. Strong prst ac present in S, is absent in 2. S: Facets of the eyes are enlarged, 1.2-1.6 times larger than the facets in the lower part of the eyes. Mid tibia usually has 3p seta. ... angustifrons Shinonaga & Kano

I-3. The Kuril Islands

MATERIAL EXAMINED

Phaonia angustifrons ommatina Zinovjev, 1981 stat. nov. (Fig. 19, Fig. 20) Phaonia ommatina Zinovjev, 1981

Fig. 17, Fig. 18, Fig. 19, Fig. 20. Heads of Phaonia, SS: 17 - P. zugmayeriae; 18 -P. angustifrons from Far East mainland; 19 - P. angustifrons ommatina, with widened frons 20 -P. angustifrons ommatina, paratype.

Type material. Holotype, S: RUSSIA, Sakhalin reg., Kunashir Island, Yuzhno-Kurilsk env., volcano Mendeleeva, (44.0° N, 145.8° E), 9.07.1979, A. Zinovjev, (ZIN). Paratypes: the same lebel as the holotype, 4-9.07.1979, 16 SS, 1 2 (ZIN and ZMUM); Iturup Island, (45.20° N, 147.84° E), 25.06.1968, V. Rikhter, 1 S (ZIN).

Other material: RUSSIA: Sakhalin reg.: Iturup Isl., Rybaki, 5 km SW of Kurilsk, (45.20° N, 147.84° E), 22-23.06.1968, V. Rikhter ,1 S, 1 2 (ZIN); Kunashir Isl.: Kurilsky NR, caldera of the Golovnin volcano, 43.841° N, 145.509° E,

25-29.08.2009, YPT, I. Melnik, 1 $, K. Makarov, A. Zaitsev, 1 $ (all ZIN), 3-5.07.2014, I. Gomyranov, 3 SS, 5.07.2014, T. Galinskaya, 1 & (all ZMUM); Filatovsky cordon, 44.11° N, 146.01° E, 18-19.07.2014, I. Gomyranov, 1 S (all ZMUM); Tretyakovo village, 43.59° N, 145.38° E, 13-22.09.2009, YPT, I. Melnik, 3 $ (ZIN), 8-15.07.2014, I. Gomyranov, 8 SS (ZMUM); Simushir Isl., volcano Milna env., (46.82° N, 151.78° E), 18.07.1958, N. Violovich, 1 S (ZIN).

DISCUSSION. Zinovjev (1981b: 625) gave the following diagnosis of P. ommatina: "Similar to P. angustifrons but the colouring of femora; scutellum and wings are lighter. Male frons is on average narrower than in P. angustifrons, cerci have a little bit different shape. Abdomen is less dusted than in P. angustifrons, with less distinct median vitta. Male eyes are with upper facets more strongly enlarged (1.8-2.1 times larger than lower facets, while in P. angustifrons 1.2-1.6 times) (this character is unique for the P. latipalpis group)."

We reexamined Zinovjev's type series and a series of recently collected specimens and came to the following conclusions.

a. We have not found any differences in wing colour, shape of cerci and dusting of abdomen.

b. Colour of femora is lighter than that of P. angustifrons, f2 and f3 are dark only at basal 1/2 to 1/5, sometimes f3 is entirely yellow.

c. Colour of scutellum is more yellow than that of P. angustifrons, (varies from 50% of surface to almost entirely (90%) yellow).

d. Most specimens of the type series of P. ommatina have closely approximated eyes, fronto-orbital plates are almost invisible as in Figure 20. Unfortunately, the character is also variable, several freshly collected males in ZMUM have fronto-orbital plates divided by dark frontal vitta (Fig. 19) i.e., eyes are separated more widely than in P. angustifrons.

e. The size of upper facets of male eyes is also variable: they are enlarged strongly or moderately. Zinovjev did not include male specimens from the Kuril Islands with moderately enlarged facets in the type series of P. ommatina but identified them as P. angustifrons.

Therefore, P. ommatina is undoubtedly similar to P. angustifrons. There is a curious peculiarity of the distributions of these taxa. P. ommatina is recorded only from Kunashir and Shikotan Islands. Meanwhile, to south of the Kuril Islands, in Japan, the typical P. angustifrons occurs, and to north of the Kuril Islands, in Sakhalin Island, the typical P. angustifrons is again recorded. We suggest the following explanation to these observations. Japan, Sakhalin, and the Kuril Islands had divided from the continent and each other when the sea level had risen, that is, ca. 15.000 years ago. Since then, large populations haven't changed much, whereas small islands' populations have changed under influence of genetic drift and show wide ranges of intra-population variability.

That is why we prefer to identify P. ommatina as an insular subspecies of P. angustifrons - P. angustifrons ommatina Zinovjev, 1981 stat. nov.

We believe that Zinovjev's diagnosis of P. ommatina have to be corrected for the following reasons. (1) It requires changing of generally accepted diagnosis of P. angustifrons, a much widely distributed and earlier described species. (2) Using strongly enlarged eye facets as the main diagnostic character leaves unresolved the question of how to distinguish females of P. ommatina. We offer the following diagnosis of the Kuril subspecies:

- S $: ,f2 and f3 are only basally dark, sometimes f3 is entirely yellow. Scutellum is yellow in apical 50 to 90% of surface. Anterior and posterior spiracles are yellow or brownish-yellow. S: Facets of the eyes are usually greatly enlarged. ... angustifrons ommatina

- S $: f2 andf3 are dark, yellow only at apex. Scutellum is yellow only at very apex.

Anterior and posterior spiracles are dark. S: Facets of the eyes are moderately

enlarged. . angustifrons angustifrons

P. latipalpis Schnabl, 1911 Kuril form

RUSSIA: Sakhalin reg., Kunashir Isl.: Kurilsky NR, Andreevsky cordon, 43.54° N, 145.37° E, 6-8.07.2014, I. Gomyranov, 3 SS; Filatovsky cordon, 44.11° N, 146.01° E, 18-19.07.2014, I. Gomyranov, 1 S (all ZMUM); Mendeleevo env.: road to Sernovodsk, (43.95° N, 145.68° E), 9.06.1968, V. Rikhter, 1 S (ZIN), 6.10.1968, K. Gorodkov, 1 $ (ZIN), Tretyakovo env., 43.59° N, 145.38° E, 13-22.09.2009, YPT, I. Melnik, 1 $ (ZIN); Yuzhno-Kurilsk env., (44.0° N, 145.8° E), 25.09.1967, V. Sychevskaya, 1 $ (ZMUM); 2-7.07.1979, A. Zinovjev, 5 SS, 2 $$ (ZIN); Shikotan Isl.: Malokurilsk env., 43.84° N, 146.91° E, 22.09.1968, K. Gorodkov, 1 $ (ZIN); Tserkovnaya bay, 43.75° N, 146.70° E, 10-14.06.2012, Yu. Sundukov, 2 $$ (ZMUM).

REMARKS. It is remarkable that the Kuril specimens of P. latipalpis also somewhat differ from those from the Far East mainland: fore femur has colouration from entirely yellow to mostly darkened (as in Japanese specimens shown in Fig. 21, Fig. 22). We do not share the opinion of several Japanese and Chinese authors that these colour variations should be regarded as separate species. We interpret such deviation as some insular effect and consider all Kuril specimens with yellow f2 and f3 as Kuril form as P. latipalpis.

IDENTIFICATION KEY FOR Phaonia latipalpis group (Kuril Islands), SS, $$.

- f2 and f3 entirely yellow. Scutellum entirely yellow. S: Facets of eyes not enlarged. . latipalpis Schnabl

-f2 andf3 are dark basally. Scutellum is yellow in apical 50 to 90% of surface. S: Facets of eyes are enlarged. ... angustifrons ommatina Zinovjev

II. Short notes on faunas of the P. latipalpis group of China, Japan and Nearctic region

China

The study of fauna of the Phaonia latipalpis group in China began in 1986 with the description of P. subommatina. Nowadays the Chinese fauna of the

P. latipalpis group includes 17 species of which 15 were newly described from China: Xue & Chao (1998); Ma et al. (2002,); Xue et al. (2006); Wu et al. (2015). Most articles are written in Chinese, only a small part of the data is available in English. The majority of species differ from each other by details of colouration of scutellum, femora etc., therefore, the validity of them seem questionable. We found it impossible to make any assumptions about the Chinese fauna until a revision of the previously described and recorded species will be published.

Japan

According to Shinonaga (2003), Japanese fauna of the P. latipalpis group includes 5 species: P. angustifrons Shinonaga & Kano, 1971; P. dorsolineata Shinonaga & Kano, 1971; P. hydrocharis Shinonaga & Kano, 1971; P. japonica Shinonaga & Kano, 1971 and P. latipalpis Schnabl, 1911. Recorded by Kato (1936) P. apicalis was excluded from Japanese fauna by Shinonaga (2003). P. angustifrons was considered in chapter I-2.

Through scientific exchange, some Japanese specimens were deposited in European museums, and we studied the specimens kept at MNHN and ZIN: they are paratypes or have identification labels of Shinonaga himself.

MATERIAL EXAMINED

P. dorsolineata Shinonaga & Kano, 1971 (Fig. 21)

JAPAN, Kyushu Isl., Kagoshima, Yakushima: Kusugawa-hodo, 18.05.1972, S. Shinonaga, 2 SS (MNHN and ZIN); Kosugidani, 16.05.1972, S. Shinonaga, 1 2 (MNHN).

P. hydrocharis Shinonaga & Kano, 1971

Paratype 1 S: JAPAN, Honshu, Nagano prefecture, Karuizawa, (36.4° N, 138.6° E), 27.07.1970, R. Kano (MNHN).

P. japonica Shinonaga & Kano, 1971 (Fig. 22)

Paratypes: JAPAN, Honshu, Fukushima prefecture, Inawashiro (37.55° N, 140.10° E), 1 S, 1 2 (MNHN); 1 S (ZIN).

REMARKS. Examined specimens of P. japonica have scutellum entirely or almost yellow; darkening on f1 as broad basal ring, katepimeron has 1-2 setulae.

Shinonaga's (2003) identification key is based mostly on details of colouration off and scutellum, the key has obvious drawbacks.

a. It is not indicated that the key is for male only, females which have no prst ac cannot be identified.

b. Sometimes the key contradicts descriptions.

c. Some significant characters were overlooked. For example, all Japanese specimens of P. japonica have setulae on katepimeron but this character was not mentioned in Shinonaga & Kano (1971) or Shinonaga (2003).

Our attempts to identify Shinonaga's specimens by Shinonaga's (2003) key were not successful. Zinovjev (1980: 913) came to the same conclusions and excluded species of the P. latipalpis group from his review of Far Eastern Phaonia.

Nearctic

Fig. 21, Fig. 22. Japanese specimens in MNHN collection: 21 - specimen identified by S. Shinonaga as P. dorsolineata; 22 - paratype of P. japonica.

MATERIAL EXAMINED

Phaonia apicalis apicalis Stein, 1915

Phaonia apicalis Stein, 1915: Huckett (1965)

No Nearctic material seen. Palaearctic material see Part I-1.

DISTRIBUTION. Holarctic. In Nearctic Alaska and N Canada (Huckett, 1965).

Phaonia apicalis apicata Johannsen, 1916

P. apicata Johannsen, 1916

P. apicata Johannsen, 1916: Malloch (1919)

P. solitaria Stein, 1920; syn. nov.

P. apicalis apicata Johannsen, 1916; stat. nov.

CANADA: Ontario, Ottawa, Nepean, 45.32° N, 75.72°W, 16.06-2.07.2016, J. O'Hara, 5 $$ (1 $ with 1p on t1, all withoutprst ac, all 2 + 3dc) (ZMUM, ISEA).

USA: NC, Highlands, (35.06° N, 83.20°W), 1150 m, 25.05.1957, J.R. Vockeroth, 1 S; 29.08.1957, J. C. Chillcott, 1 S (with prst ac) (ZIN); RI, Coventry Co, 41.69° N, 71.55°W, 8-14.05.2017, A. Medvedev, 1 S (without prst ac) (ZMUM).

DISTRIBUTION. Canada and USA.

Phaonia bysia Walker, 1849

USA, NC, Highlands, (35.06° N, 83.20°W), 1150 m, 4.06.1957, J.R. Vockeroth, 1 TN, Great Smoky Mountains National Park, 1550 m, 23.08.1957, J.C. Chillcott, 1 S, with det. label by Vockeroth (all ZIN).

DISTRIBUTION. Canada and USA.

REMARKS. According to Malloch (1923), humeri (postpronotal calli) is at least partly yellow. In our specimens, postpronotal calli are entirely dark.

DISCUSSION on the Nearctic fauna.

Unfortunately, there is still no major revisional study on the Nearctic Muscidae like the Hennig's study on the Palearctic fauna. The Nearctic fauna of Phaonia was reviewed by Malloch (1923); the key for Phaonia of Northern Canada, Alaska and Greenland was published by Huckett (1965). We restrict our review to three species we were able to examine.

P. a. apicalis. Huckett (1965) listed the species for the USA, Alaska and Canada: Northwest Territories, Manitoba and Quebec. Given the fact that some our specimens are from Kamchatka, its presence in North America is not surprising. Huckett indicated that antenna was entirely dark, so his North American material represents the Palaearctic subspecies of P. a. apicalis.

P. a. apicata. According to the original description of P. apicata, a pair ofprst ac is either present or absent in males (Johannsen, 1916): "Male ... one pair of inner dorso-centrals in front of the transverse suture present, though but small in some specimens and absent in one ..."; in females prst ac is absent. Material we examined fits the description: one our male has prst ac present, another one does not, in all 5 females prst ac is absent. One of our 5 22 has p seta on t1. Thus, the variability of P. apicata is quite similar to that of P. apicalis, there are only some differences in colouration among these taxa. We propose to identify P. apicata as P. apicalis apicata Johannsen, 1916; stat. nov., a Nearctic subspecies of nominotypical P. a. apicalis.

SYNONYMS. According to Malloch (1923), the only difference between P. apicata and P. solitaria is that the latter has prst ac. However, according to above discussion, P. apicata may or may not have prst ac. Therefore, P. a. apicata Johannsen, 1916 = P. solitaria Stein, 1920 syn. nov.

EXCLUDED SPECIES. At least two more species of Nearctic Phaonia should belong to the P. latipalpis group. We know them only from Malloch (1923) publication.

Phaonia curvinervis Malloch, 1919. Margins of postpronotal calli, posterolateral margins of mesonotum, scutellum and margins of pleural sclerites are yellowish. No prst ac; dc 2 + 3; pra as long as in P. apicalis; notopleuron and meron are bare. t1 is with 2 ad and 2 p. Vein R4 + 5 slightly curved forward apically.

Phaonia winnemanae Malloch, 1919. Similar to P. a. apicata, but postpronotal calli, posterolateral margins of mesonotum and tarsi is yellow.

IDENTIFICATION KEY FOR Nearctic Phaonia latipalpis group

1. pra is about 0.5x as long as posterior notopleural seta; the latter is slightly shorter than anterior one. Cheek is distinctly narrower than postpedicel. ... bysia Walker

- pra is about 2.5x longer than posterior notopleural seta and 1.5x longer than anterior one. Cheek is as wide or wider than postpedicel. ...2

2. Postpedicel at base and pedicel are yellowish. f1 is yellow or slightly darkened; trochanters are yellow. The USA and Canada. ... a. apicata Johannsen

Antenna is dark. f1 is mostly dark; trochanters are brown. Palaearctic and North of America. ... a. apicalis Stein

Acknowledgements

We thank Olga Ovchinnikova and Galina Suleymanova (ZIN, Saint Petersburg); Christophe Daugeron and Emmanuel Delfosse (MNHN, Paris) for the possibility to examine their material. We thank Dmitry Gavryushin (Moscow) for his corrections and advice. We are especially grateful to all collectors for the rich material listed above.

References

d'Assis-Fonseca E.C.M. 1968. Diptera Cyclorrhapha Calyptrata, Muscidae // Handbook for the Identification of British Insects. London: Royal Entomological Society. Vol. X. Part 4(b). P. 22-27.

Gregor F., Rozkosny R., Bartak M., Vanhara J. 2002. The Muscidae (Diptera) of Central Europe. Brno: Mazaryk University. 280 p.

Hennig W. 1963. Muscidae (Part, Lieferung 233, 234 and 241) // E. Lindner (ed.). Die Fliegen der Palaearktischen Region. Stuttgart: Schweizerbart. 63b. P. 772-899. [In German]

Huckett H.C. 1965. The Muscidae of Northern Canada, Alaska and Greenland (Diptera) // Memoirs of the Entomological Society of Canada. Suppl. S42. 369 p.

Johannsen O. A. 1916. New Eastern Anthomyiidae (Diptera) // Transactions of the American Entomological Society (1890-). Vol. 42(4). P. 385-398.

Kato S. 1936. A New and Four Unrecorded Species of Phaoniinae (Dipt., Muscidae) from Japan // Insecta matsumurana. Vol. 11. P. 24-27.

Ma Z., Xue W., Feng Y. 2002. Fauna Sinica Insecta. Vol. 26. Diptera: Muscidae II, Phaoniinae I. Beijing: Science Press. 421 p. [ In Chinese].

Malloch J. R. 1923. Flies of the Anthomyiid genus Phaonia Robineau-Desvoidy and related genera, known to occur in North America // Transactions of the American Entomological Society (1890-). Vol. 48(3). P. 227-282.

Malozemov A.Yu. 1992. On the fauna and ecology of true flies (Diptera, Muscidae) of the eastern macroslope of the Pripolar Ural Mountains // N.V. Nikolaeva (ed.). Insects in natural and anthropogenic biogeocenozes of the Urals. Yekaterinburg: Nauka. P. 91-93. [In Russian]

Mihalyi F. 1974. Description of Phaonia szelenyii sp. nov. found in Hungary (Diptera: Muscidae) // Folia entomologica Hungarica. Vol. 27. P. 119-120.

Pont A.C. 1986. Family Muscidae // A. Soos, L. Papp (eds). Catalogue of Palaearctic Diptera 11. Budapest: Akademia Kiado. P. 57-215.

Pont A.C. 2013. Fauna Europaea: Fanniidae & Muscidae // Beuk P., Pape T.P. 2013. Fauna Europaea: Diptera: Brachycera. Fauna Europaea version_2018.12. https://fauna-eu.org.

Schnabl J. 1888. Contributions a la faune dipterologique. Additions aux descriptions precendentes des Aricia et descriptions des especes nouvelles // Horae societatis entomologicae rossicae. Vol. 22. P. 378-486. [In French]

Shinonaga S. 2003. Monograph of the Muscidae of Japan. Tokyo: Tokai University Press.

347 p.

Shinonaga S., Kano R. 1971. Fauna Japonica. Vol. I. Muscidae (Insecta: Diptera). Academic Press of Japan. 242 p.

Vikhrev N., Erofeeva E. 2018. Review of the Phaonia pallida group (Diptera, Muscidae) // Russian Entomological Journal. Vol. 27(3). P. 315-322. https://dx.doi.org/10.15298/rusentj.27.3.14.

Wu C-G., Dong J-M., Wei L-M. 2015. Two new species of Phaonia latipalpis group (Diptera: Muscidae, Phaonia) from China // Chinese Journal of Vector Biology and Control. Vol. 26(4). P. 404-407. https://dx.doi.org/10.11853/j.issn.1003.4692.2015.04.019.

Xue W-Q., Chao C-M. 1998. Flies of China. Vol. 1. Shenyang: Liaoning Science and Technology Press. 1365 p. [In Chinese]

Xue W-Q., Zhang C., Zhu Yu. 2006. Three new species of Phaonia Robineau-Desvoidy (Diptera: Muscidae) from China // Oriental Insects. Vol. 40(1). P. 127-134. https://dx.doi.org/10.1080/00305316.2006.10417464

Zinovjev A.G. 1980. Phaoniinae (Diptera, Muscidae) of the Far East // Entomologicheskoe Obozrenie. Vol. 59(4). P. 904-913. [In Russian]

Zinovjev A.G. 1981a. On the classification of the Palearctic flies of the genus Phaonia R.-D. (Diptera, Muscidae) // Entomologicheskoe Obozrenie. Vol. 60(3). P. 686-697. [In Russian]

Zinovjev A.G. 1981b. Two new species of the genus Phaonia (Diptera, Muscidae) from the Soviet Far East // Zoological Journal. Vol. 60(4). P. 623-625. [In Russian]

Zinovjev A.G. 1981c. To the fauna of robberflies (Diptera, Asilidae) and true flies of the genus Phaonia (Diptera, Muscidae) of the Central Urals // L.L. Savenkova (ed.). Fauna and ecology of insects. An interdisciplinary collection of scientific works. Perm: Perm State Institute. P. 119-126. [In Russian]

ГРУППА ВИДОВ PHAONIA LATIPALPIS (DIPTERA, MUSCIDAE): ОБЗОР РОССИЙСКОЙ ФАУНЫ И ЗАМЕТКИ ПО ФАУНАМ СОПРЕДЕЛЬНЫХ

ТЕРРИТОРИЙ

Н.Е. Вихрев, Е.А. Ерофеева

Зоологический музей МГУ им. М.В. Ломоносова, Россия e-mail: nikita6510@yandex.ru

Рассмотрена группа видов Phaonia latipalpis (Diptera, Muscidae). Были прояснены некоторые проблемы таксономии группы P. latipalpis. Так, на основе изучения более 400 образцов из нескольких отдаленных местностей была исследована изменчивость P. zugmayeriae. Такой подход может оказаться полезным для других таксонов Phaoninae. Фауна группы P. latipalpis Западной Палеарктики, Дальнего Востока России и Курильских островов рассматривалась отдельно, предложены идентификационные ключи для каждой из этих территорий. Кратко рассмотрена фауна группы P. latipalpis Китая, Японии и Неарктического региона. Предложен один новый синоним: P. apicata Johannsen, 1916 = P. solitaria Stein, 1920, syn. nov. Предложено понизить с видового до подвидового два таксономических статуса: P. ommatina Zinovjev, 1981 = P. angustifrons ommatina Zinovjev, 1981 stat. nov.; P. apicata Johannsen, 1916 = P. apicalis apicata Johannsen, 1916 stat. nov.

Ключевые слова: Diptera, Muscidae, группа Phaonia latipalpis, новые находки