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http://journals.tubitak.gov.tr/zoology/

Turkish Journal of Zoology Turk J Zool(2017) 41: 43-59© TÜBİTAKdoi:10.3906/zoo-1512-60

A review of subtribe Phrosinellina Verves, 1989, with descriptionof Phrosinella (Asiometopia) kocaki sp. nov. from the Middle East

(Diptera: Sarcophagidae: Miltogramminae: Metopiaini)

Yury VERVES1,*, Liudmyla KHROKALO2

1Institute for Evolutionary Ecology, National Academy of Sciences of Ukraine, Kyiv, Ukraine2National Technical University of Ukraine “Kyiv Polytechnic Institute”, Kyiv, Ukraine

* Correspondence: fly_@voliacable.com

1. IntroductionSubtribe Phrosinellina Verves, 1989 includes 31 species from three genera: Gymnoprosopa Townsend, 1892 (6 species); Gymnopsidia Shewell, 1987 (1 species); and Phrosinella Robineau-Desvoidy, 1863 (24 species). All species are very xerophilous and psammophilous and mainly distributed in arid and subarid zones of the Holarctic and Oriental regions. Larvae are known as kleptoparasites (inquilines) in ground nests mainly of different sphecoid and occasionally of pompiloid wasps and bees (Allen, 1926; Charykuliev, 1965; Downes, 1965; Evans, 1966a; Krombein, 1967; Rohdendorf, 1967, 1971a, 1971b; Myartzeva, 1972a; Verves, 1976; Spofford et al., 1989; Verves, 1989; Spofford and Kurczewski, 1990, 1992; Povolný and Verves, 1997; Verves and Khrokalo 2006; Zerova et al., 2006; Evans and O’Neil, 2007; Pickering, 2011; Pulawski, 2015).

2. Materials and methodsThis article is a result of study of the Natural History Museum (London) collection of miltogrammine flies and original analysis of literature data.

All photographs were prepared using a stereomicroscope (Leica M205C, Leica Microsystems, Wetzlar, Germany) with a Canon EOS 5D Mark II body camera (Canon Inc., Tokyo, Japan). As a mount was essentially not flat, a series of photographs were made at different focal depths.

They were compiled into one sharp image, in accordance with the criterion of maximal power in the spatial high-frequency domain, with the software Helicon Focus Pro 5.3.14 X64 (Helicon Soft Ltd., Kharkiv, Ukraine).

Authors of faunistic data not registered in catalogues (Downes, 1965; Verves, 1986; Pape, 1996) are given in footnotes.

Acronyms: CNC: Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada; BMNH: Natural History Museum, London, United Kingdom; IEE: Institute for Evolutionary Ecology, National Academy of Sciences, Kyiv, Ukraine; MZLU: Museum of Zoology, Lund University, Lund, Sweden; MNHN: Muséum National d’Histoire Naturelle, Paris, France; MZUF: Muzeo Zoologico de “La Specola”, Florence, Italy; SEM: Snow Entomology Collection, University of Kansas, Lawrence, Kansas, USA; SMNS: Staatliches Museum für Naturkunde, Stuttgart, Germany; TAU: Tel Aviv University, Israel; USNM: Department of Entomology, Smithsonian Institution, National Museum of Natural History, Washington, DC, USA; ZIN: Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russian Federation.

Abbreviations of morphological features: acr - acrostichal seta; ad - anterodorsal seta; ap - apical seta; d - discal seta; dc - dorsocentral seta; dm-cu - discal medial-

Abstract: Phrosinella kocaki Verves & Khrokalo sp. nov. from Israel is described as being new to science. A similar species, Phrosinella kozlovi (Rohdendorf, 1925), is recorded for the first time for Turkey and redescribed. Faunistic and ecological data on 31 species from 3 genera of subtribe Phrosinellina are given: Gymnoprosopa Townsend, 1892 (6 species); Gymnopsidia Shewell, 1987 (1 species); and Phrosinella Robineau-Desvoidy, 1863 (24 species). An original key to the genera and subgenera is presented.

Key words: Phrosinella kocaki, Middle East, new species, Sarcophagidae, Phrosinellina, genera, key, species composition, geographical distribution, habits

Received: 22.12.2015 Accepted/Published Online: 28.05.2016 Final Version: 25.01.2017

Research Article

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cubital crossvein; fr - frontal seta; ia - intraalar seta; kepst - katepisternal seta; M - medial vein; npl - notopleural seta; oc - ocellar seta; orb - orbital seta; pd - postdorsal seta; postorb - postorbital seta; pprn - postpronotal seta; R1 - first longitudinal vein; R4+5 - third longitudinal vein; r5 - first posterior cell; t2 - midtibia; vte - outer vertical seta; vti - inner vertical seta.

3. Results and discussionSubtribe Phrosinellina Verves, 1989Verves, 1989: 119; Povolný and Verves, 1997: 107; Nandi, 2002: 122; Verves and Khrokalo, 2006: 101.

Bright colored flies of small or medium size (3.0–10.0 mm in length). Eyes large, bare; frons as wide as or more widened than eye, parafacials very broad, genae narrow or midheight; flagellomere 3–7× as long as pedicel; arista bare or microchaetose, widened in the proximal 0.6–0.8. Lower margin of head more or less shortened; prescutellar acr absent or unclear; wing cell r5 narrowly open or closed; abdomen usually with distinct dorsal black or brown spots or bands at hind surfaces of tergites, or sometimes with checkering pattern; sexual dimorphism inconsiderable (male fore tarsi often with bristles or brushes of specialized setae).

Description of Phrosinella (Asiometopia) kocaki Verves & Khrokalo, sp. nov. (Figures 1–6)

Male: Body color: Bright colored. Head almost entirely densely silvery gray pollinated, frontal vitta black, finely dusted, scapus and pedicel brown, flagellomere and arista black, palpi yellow. Thorax densely light gray dusted, mesonotum with broad dark median longitudinal stripe before suture only. Legs black, fore tarsi brown. Wings hyaline, basicosta and epaulette yellow. Abdomen yellowish gray pollinated, in basal half yellowish brown laterally, with black dorsal drawing. 1+2nd tergite with narrow indistinct dark band or badly developed median spot and paired lateral stripes in hind 0.3; each of 3rd and 4th tergites in hind 0.4–0.5 with square or rounded three spots separated from one another by light dusting of narrow longitudinal bands; 5th tergite with similar drawing in hind 1/2–2/3. Genitalia shining black.

Body length: 6.5–8.5 mm.Head: Frons at vertex 0.43–0.45×, at level of antennal

base 0.47–0.50× of head width. Frontal vitta 1.2–1.5× widened backwards, at level of fore orb 0.8–0.9× as wide as one of parafrontalia. Flagellomere 3.7–4.5× as long as pedicel, arista widened in basal 0.8–0.9, bare. Parafacialia at level of antennal base 0.25–0.31×, genae 0.13–0.20× of eye height. Palpi elongate, at apex widened. Two regular rows of postorb; vte strong, 0.5× as long as vti; oc midlong; orb 1+2, strong; fr 9–14, beside them 4–5 fore pairs crossed, hind pairs reclinate; hind part of parafrontalia with several black erect long hairs; fore part of parafrontalia and

parafacialia with microscopic yellow setae, almost bare; facial ridge above angular bristles with 2–3 black short setae; oral bristles numerous, their fore 2–3 pairs strong, others hair-like; genae and occiput with numerous yellow erect midlong setae (Figures 1–3).

Thorax: Covered with densely black erect midlong hairs; acr 0+1, fine; dc 2–3+3, only prescutellar pair strong; ia 0–1+2–3, very fine; pprn 1–2; kepst 1+2, mid long; npl 2–3. Scutellum with 3 pairs of strong marginals (ap crossed), d less.

Legs: Each of 1st and 2nd tarsomeres of fore tarsi with elongate curved ad and pd, which reached to the end of 4th tarsomere, 3rd tarsomere with more short similar setae, 4th and 5th tarsomeres without long setae; t2 with one ad (Figure 4).

Wings: Costal spine small, badly developed, r5 narrowly open, R1 at upper side with 1–3 black hairs in basal half, R4+5 dorsally with a row of setae in basal 0.5–0.7 of first section, ventrally with 2–3 basal hairs, M right-angled, dm-cu slightly s-like curved, almost straight, the ratio of 3rd and 5th costal sections is 1:1.5–2.3.

Abdomen: 1+2nd tergite without mediomarginals, 3rd tergite with a pair of erect long mediomarginals (Figures 5 and 6).

Female: Related to male; fore tibia without specialized setae; median spot placed in hind 0.2–0.3 of length of 3rd abdominal tergite, and lateral stripes of this tergite narrow, partly reduced.

Etymology: The specific name is given in honor of one of the most known Turkish entomologists, Prof Emeritus Dr Ahmet Ömer Koçak, Van, Turkey.

Type material. Holotype (male): Israel: Sedom, 26.vi.1976, A. Freidberg leg. (TAU). Paratypes (males and females): Israel: Zin Wilderness, Nahal Zin at En Akrabim, cane-covered sandy wadi el. – 61 m, 30°53′38″N, 35°09′39″E [GPS], 12, 15, 23.iii. et 4.iv.1995, 3 ♂, 2 ♀ (Irwin) (TAU & IEE); Negev, Ein Avdat National Park, Ein Aqev spring, 25.v.2006, 1 ♀ (K. Szpila) (IEE); ‘Enot Samar, 22.04.1998, 1 ♂ (A. Freidberg) (TAU).

Comparison: This species is related to P. (A.) kozlovi (Rohdendorf, 1925) (Figures 7–11) by habitus, drawings of 3rd and 4th abdominal tergites and badly developed costal spine, but differs by ciliation of ♂ fore tarsi: bristles of 1st and 2nd tarsomeres very long, but 4th tarsomere without similar setae, and by three distinctly separated hind black spots at 5th abdominal tergite in both sexes.

Redescription of Phrosinella (Asiometopia) kozlovi (Rohdendorf, 1925)

(Figures 7–11).Male: Body color: Brightly painted. Head densely

silvery gray pollinated, frontal vitta matt black, with fine brownish tincture and white pruinescence; ocellar triangle and antennae entirely black, palpi brownish yellow. Thorax

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Figures 1–6. Phrosinella (Asiometopia) kocaki Verves & Khrokalo, sp. nov. Holotype ♂: 1- head anteriorly; 2- head laterally; 3- head dorsally; 4- fore left tarsus dorsally; 5- abdomen dorsally; 6- total view dorsally.Figures 7–11. Phrosinella (Asiometopia) kozlovi (Rohd.). ♂, Turkey: Iğdır, Ararat [recent: Mountain of Ağrı] below Serdarbulak, 5000′, 4.ix.1960: 7- head anteriorly; 8- head laterally; 9- fore tarsus dorsally; 10- abdomen dorsally; ♂, Mongolia: Kobdo aimak, Bulgan river, 25 km NNW of Bulgan habitation, 2.vii.1980: 11- abdomen dorsally.

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densely light gray pollinate, mesonotum with three dark median longitudinal stripes in anterior part, which reach middle level of postsutural arena; fore spiracles gray, hind ones yellowish-white. Wings hyaline, basicosta and epaulette yellow, legs brownish black. Abdomen silver gray pollinated, in basal half yellowish brown laterally, with black dorsal drawing. 1+2nd tergite with badly developed median spot and paired lateral stripes in hind 0.6, each of 3rd and 4th tergites in hind 0.4–0.5 with square median spot and separated from it by narrow areas of light dusting paired lateral stripes, 5th tergite with shining black marginal band in hind 0.6, and silver gray dusting in fore 0.4. Genitalia shining black.

Body length: 6.0–11.0 mm.Head: Frons at vertex 0.46–0.50×, and at level of

antennal base 0.48–0.51× of head width. Frontal vitta almost parallel-sided, at level of fore proclinate orb 0.7–0.8× wide as one of parafrontalia. Flagellomere 4.0–5.0× as long as pedicel, arista widened in basal 0.7–0.8, almost bare, microchaetose. Parafacialia at level of antennal base 0.40–0.44×, genae 0.22–0.26× of eye height. Palpi midlong, at apex widened. Two regular rows of postorb in upper part of head; vte strong, only slightly shortly as vti; ocellar triangle with a pair of very strong lateroproclinate oc, and covered with numerous erected hairs; orb 1+2, strong; fr 10–16, crossed and strong; parafrontalia entirely covered with sparse black hairs, more long in hind part of frons; parafacialia in fore part with numerous fine black setae, bare near eyes; facial ridge above long and erected angular bristles with 2 black short setae; 6–8 pairs of oral bristles, their fore 2–3 pairs strong, others hair-like; genae and occiput with numerous pale erect midlong setae (Figures 7 and 8).

Thorax: Covered with numerous erect midlong black hairs; acr 0+1; dc 2+3-4; ia 1+2-3, very fine; pprn 2; kepst 1+1; npl 2. Scutellum with 3 pairs of long and strong marginals (ap crossed), d less.

Legs: Each of 1st–4th tarsomeres of fore tarsi with a pair of very long curved apical ad and pd: each of them is accompanied by strong apical spine-like ad or filiform pd accordingly; t2 with one ad (Figure 9).

Wings: Costal spine small, badly developed, r5 closed at edge of wing or narrowly open, R1 at upper side with 3–4 black hairs in basal half, R4+5 dorsally with a row of setae in basal 0.6–08 of first section, ventrally with 2–3 basal hairs, M obtuse angled, dm-cu straight, the ratio of 3rd and 5th costal sections is 1:1.9–2.2.

Abdomen: Each of 1+2nd–5th tergites with erected pair of medial marginal bristles. Genitalia small (Figures 10 and 11).

Female: Related to male, but fore tarsi without specialized chaetom.

Material examined: Turkey: Iğdır, “Ararat” [recent: Mountain of Ağrı] below Serdarbulak, 4.ix.1960, 5000′,

1 ♂ (Guichard & Harvey) (BMNH); Mongolia: Kobdo aimak, Bulgan river, 25 km NNW of Bulgan habitation, meadow, 2.vii.1980 (I. Kerzner), 1 ♂ (ZIN).

Key to genera and subgenera1. orb 1+3–4. Genae covered by long upcurved setae

(Figure 12)……….........…….. g. Gymnopsidia Shewell- orb 1+2. Genae covered by fine inconspicuous hairs... 22. fr not reaching below antennal base, almost not

divergent (Figure 13).............. g. Gymnoprosopa Town.- fr reaching to level of middle of pedicel, distinctly

divergent (Figure 14) ….........…… g. Phrosinella R.-D.a. Abdomen with checkering pattern; 2nd tarsomere of

male fore tarsi with a tuft of long hairs (Figure 16) ……………....................…………sg. Euhilarella Town.

- Abdomen with distinct dark bands or spots at tergites (Figure 15); male fore tarsi with another chaetom ….. b

b. Frontal stripe as wide or wider than one of parafrontalia; R1 bare dorsal; male fore tarsi without long hairs …................................................................. sg. Phrosinella s. str.

- Frontal stripe distinctly narrower than one of parafrontalia (Figures 3 and 17); R1 often haired dorsall............................................................................… c

c. r4+5 hairy from base to the middle of its 2nd section dorsally (Figure 18); 5th abdominal tergite completely shining black (Fig. 19); male fore tarsi without long hairs …..................................…… sg. Caspiomyia Rohd.

- r4+5 hairy only in its 1st section; 5th abdominal tergite more or less lightly dusted anteriorly (Figures 5 and 10); male fore tarsi with paired elongate apical setae at several tarsomeres (Figures 4 and 9)…...........................................................................……sg. Asiometopia Rohd.Annotated list of genera and species of PhrosinellinaGenus Gymnoprosopa Townsend, 1892Gymnoprosopa Townsend, 1892: 108. Type species:

Gymnoprosopa polita Townsend, 1892; by original designation.

Gymnoprosopa: Allen, 1926: 95 [revision]; Downes, 1965: 936 [catalogue]; Krombein and Kurczewski, 1963: 139 [habits]; Shewell, 1987: 1182 [in key]; Spofford et al., 1989: 255 [habits]; Pape, 1996: 89 [catalogue].

Bright colored small flies (3.5–6.0 mm in length). Frontal vitta matt yellow to reddish, as wide or widened as one as parafrontalia; fr not reaching below antennal base, almost not divergent; orb 1+2; genae covered by inconspicuous black hairs, without pale setae; lower margin of head almost not shortened; lunule bare. r5 open; abdominal tergites with distinct black bands at hind surfaces of tergites; sexual dimorphism practically absent. Six species are distributed in the Nearctic region.

Habits: The larvae of Gymnoprosopa sp. were developed at paralyzed nymphs and adults of Tridactylus sp. (Orthoptera) in nests of sphecoid wasp Tachytes mergus (Krombein and Kurczewski, 1963; Spofford et al., 1989).

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Figure 12. Gymnopsidia inflaticornis (Allen). ♀: head laterally (after Shewell, 1987).Figure 13. Gymnoprosopa polita Townsend. ♀: head laterally (after Shewell, 1987).Figures 14–15. Phrosinella (s. str.) nasuta (Meigen), 14, ♂, Ukraine, Kyiv region, Rokytne District, Busheve village, N 49039′, E 30035′, open cast, 27.07.2012, head laterally (original photo); 15, ♂, total view dorsally (after Venturi, 1947).Figure 16. Phrosinella (Euhilarella) sannio (Zetterstedt, 1838). ♂: fore left tarsus laterally (after Verves, 1990).Figures 17–19. Phrosinella (Caspiomyia) persa (Rohdendorf, 1935) (orig.), ♂: 17- head dorsolaterally; 18- right wing dorsally; 19- abdomen dorsally.

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Gymnoprosopa argentifrons Townsend, 1892Gymnoprosopa argentifrons Townsend, 1892: 109 [♂♀]Gymnoprosopa argentifrons: Allen, 1926: 98

[redescriptions of ♂♀, faunistic]; Downes, 1965: 936 [catalogue]; Pape, 1996: 89 [catalogue].

Holotype (male): USA, southern Florida. Deposited in SEM.

Distribution: Nearctic: USA (Florida, Indiana, Iowa, Louisiana, Maryland, Minnesota, Mississippi, North Carolina1, Texas, Virginia).

Gymnoprosopa filipalpus Allen, 1926Gymnoprosopa filipalpus Allen, 1926: 100 [♂♀].Gymnoprosopa filipalpus: Downes, 1965: 936

[catalogue]; Pape, 1996: 89 [catalogue].Gymnoprosopa filipalpis: Rohlfien and Ewald, 1974:

128, incorrect subsequent spelling of filipalpus [type revised].

Holotype (male): USA, Mississippi, McHenry. Deposited in USNM.

Distribution: Nearctic: USA (Alabama, Connecticut2, Florida, Georgia, Mississippi, North Carolina3, Texas, Virginia).

Habits: Adult flies feed on flowers of Ceanothus americanus (Allen, 1926).

Gymnoprosopa latifasciata Reinhard, 1945Gymnoprosopa latifasciata Reinhard, 1945: 72 [♂♀]Gymnoprosopa latifasciata: Downes, 1965: 936

[catalogue]; Pape, 1996: 89 [catalogue].Holotype (male): USA, Texas, Brazos Co., College

Station. Deposited in CNC.Distribution: Nearctic: USA (Texas).Gymnoprosopa milanoensis Reinhard, 1945Gymnoprosopa milanoensis Reinhard, 1945: 72 [♂♀]Gymnoprosopa milanoensis: Downes, 1965: 936

[catalogue]; Pape, 1996: 89 [catalogue].Holotype (male): USA, Texas, Milano. Deposited in

CNC.Distribution: Nearctic: USA (Texas).Gymnoprosopa pallida Allen, 1926Gymnoprosopa pallida Allen, 1926: 101 [♂]Gymnoprosopa: Downes, 1965: 936 [catalogue]; Pape,

1996: 89 [catalogue].Holotype (male): USA, Idaho, Horseshoe Bend.

Deposited in USNM.Distribution: Nearctic: USA (Idaho).Gymnoprosopa polita Townsend, 1892 (Figure 13)Gymnoprosopa polita Townsend, 1892: 109 [♀]Gymnoprosopa polita: Allen, 1926: 96 [description of

♂, redescription of ♀; faunistic; type revised]; Krombein,

1961: 80 [habits]; Downes, 1965: 936 [catalogue]; Pape, 1996: 89 [catalogue]; Shewell, 1987: 1180, figure 90 [drawing of head].

Hilarella polita: Coquillett, 1897: 128 [redescription of ♀]; Aldrich, 1905: 477 [catalogue].

Holotype (female): USA, south part of Florida. Deposited in SEM.

Synonym: Gymnoprosopa clavifrons Townsend, 1892: 109 [♂].Gymnoprosopa clavifrons: Coquillett, 1897: 127

[redescription of ♂]. Holotype (male): USA, Illinois, Carlinville. Deposited

in SEM.Distribution: Nearctic: Canada (Ontario4), USA

(Arizona5, Connecticut, Florida, Illinois, Kansas, Minnesota, Mississippi, New Jersey, New Mexico, New York, North Carolina6, Ohio, Virginia).

Habits: Adults feed on flowers of Euphorbia albomarginata (Krombein, 1961) and from the honeydew at tulip tree; in mountains flies are distributed up to 5000 feet a.s.l. (Allen, 1926).

Genus Gymnopsidia Shewell, 1987Gymnopsidia Shewell, 1987: 1182. Type species:

Gymnoprosopa inflaticornis Allen, 1926; by original designation.

Gymnopsidia: Pape, 1996: 89 [catalogue].Bright colored small flies (4.0–5.0 mm in length).

Frontal vitta matt yellow, parallel-sided, 3× widened as one as parafrontalia; flagellomere 3.5× as long as pedicel; fr not reaching below antennal base, almost not divergent; orb 1+3–4; genae covered by long upcurved bristle-like setae; lower margin of head almost not shortened; lunule bare. r5 open; abdominal tergites with distinct black bands at hind surfaces of tergites; sexual dimorphism practically absent. One Nearctic species.

Gymnopsidia inflaticornis (Allen, 1926) (Figure 12)

Gymnoprosopa inflaticornis Allen, 1926: 102 [♀]Gymnoprosopa inflaticornis: Downes, 1965 [catalogue]Gymnopsidia inflaticornis: Shewell, 1987: 1180, figure

91 [drawing of head], 1182 [in key]; Pape, 1996: 89 [catalogue].

Holotype (female): USA, New Mexico, Pecos. Deposited in USNM.

Distribution: Nearctic: USA (New Mexico).Genus Phrosinella Robineau-Desvoidy, 1863Phrosinella Robineau-Desvoidy, 1863: 82. Type species:

Phrosinella argyrina Robineau-Desvoidy, 1863 [= Tachina nasuta Meigen, 1824], by monotypy.

1 after Brimley, 19382 after Britton, 19383 after Brimley, 1938

4 after Criddle, 19285 after Krombein, 19616 after Brimley, 1938

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Phrosinella: Rohdendorf, 1935: 110 [review]; Séguy, 1941: 304 [review]; Venturi, 1947: 126 [review]; 1960: 82 [review]; Rohdendorf, 1970: 643 [key to species]; 1971a: 446 [revision]; 1971b: 132 [review]; Mihályi, 1979: 54 [in key], 85 [review]; Verves, 1986: 102 [catalogue]; 1990: 523 [in key], 552 [key to species]; Pape, 1996: 29, 122 [catalogue]; Povolný and Verves, 1997: 60 [in key], 107 [review]; Nandi, 2002: 118 [in key], 122 [review]; Verves and Khrokalo, 2006: 75 [in key], 101 [key to species]; Zerova et al., 2006: 91 [in key], 109 [key to species].

Pedimyia Rohdendorf, 1925: 81. Type species: Pedimyia fedtshenkoi Rohdendorf, 1925, by original designation.

Phrosina: Robineau-Desvoidy, 1863: 101, incorrect original spelling of Phrosinella Robineau-Desvoidy, 1863 [potential junior homonym of Phrosina Risso, 1836, Crustacea].

Bright colored small or midsized flies (3.0–11.0 mm in length). Frontal stripe usually blackish; parafacialia usually with short black hairs, rarely almost bare; fr reaching level of middle of pedicel, distinctly divergent; lunule setose; genae covered by fine inconspicuous pale setae; lower margin of head distinctly shortened; orb 1+2. r5 open or closed at wing margin; male fore tarsi often with specialized setae, rarely without them. Twenty-four species in arid zones of Holarctic and north part of Oriental (Jammu & Kashmir) regions.

Habits: The larvae of “Phrosinella sp.” have been registered in nests of two species of marine turtles: leatherback turtle Dermochelys coriacea and olive ridley turtle Lepidochelys olivacea in Mexico (Andade et al., 1992). These data most probably are the result of erroneous determination. Really common predators of eggs of marine turtles on Mexican seashores are the larvae of the sarcophagid species Eumacronychia sternalis (Lopes, 1982; Lòpez Barbosa, 1989).

Subgenus Asiometopia Rohdendorf, 1935Asiometopia Rohdendorf 1935: 126 (as genus). Type

species: Pedimyia kozlovi Rohdendorf, 1925; by original designation.

Asiometopia: Rohdendorf, 1935: 126 [key to species], 1971b: 129 [revision]; Chao and Zhang, 1998: 1528 [in key], 1532 [redescription and faunistic]; Verves, 1986: 88 [catalogue].

Phrosinella (Asiometopia): Verves, 1990: 553 [in key]; Verves and Khrokalo, 2006: 102 [in key].

Bright colored small or midsized flies (4.5–10.0 mm in length). Frontal vitta distinctly narrower than one of parafrontalia, blackish, more or less lightly dusted; lower margin of head very short; abdomen with dark bands or spots in hind parts of tergites; each of 1st–4th tarsomeres of ♂ fore tarsi with a pair of very long curved apical ad and pd; R1 dorsally bare or with several black hairs in basal half; 5th tergite densely lightly dusted in fore 0.3–0.6.

Seven species are distributed in arid zones of the Palearctic region.

Habits: The larvae of Asiometopia sp. were developed in dead insects in nests of such sphecid wasps as Liris opalipennis (prey - nymphs of crickets), Palarus aurantiacus, and Philanthus triangulum (prey - different Apoidea and Sphecoidea) (Myartzeva, 1972a).

Phrosinella (Asiometopia) beludzha (Rohdendorf, 1961)

Asiometopia ujugura beludzha Rohdendorf, 1961: 9 [♂♀]

Asiometopia beludzha: Koçak and Kemal, 2015: 343 [faunistic]; Verves, 1986: 88 [catalogue].

Phrosinella beludzha: Pape, 1996: 123 [catalogue].Holotype (male): Iran, Khuristan, Jarrahi 18 km NE

Shadegan. Deposited in SMNS.Distribution: Palearctic: Iran (Khuristan).Phrosinella (Asiometopia) karakalpaka Rohdendorf,

1935 Asiometopia karakalpaka Rohdendorf, 1935: 127;

1971b: 129 [♂♀] Asiometopia karakalpaka: Rohdendorf and Verves,

1980: 499 [faunistic]; Verves, 1986: 88 [catalogue]. Phrosinella (Asiometopia) karakalpaka: Verves, 1990:

553 [in key]. Phrosinella karakalpaka: Pape, 1996: 124 [catalogue].Holotype (male): Uzbekistan, Khorezm District,

Ravat. Deposited in ZIN.Distribution: Palearctic: Mongolia; Turkmenia;

Uzbekistan.Habits: Psammophilous species.Phrosinella (Asiometopia) kocaki Verves & Khrokalo,

sp. nov. (Figures 1–6).Phrosinella (Asiometopia) kocaki Verves & Khrokalo,

2015 [♂♀]Holotype (male): Israel, Sedom. Deposited in TAU.Distribution: Palearctic: Israel. Habits: Psammophilous species.Phrosinella (Asiometopia) kozlovi (Rohdendorf, 1925)

(Figures 7–11) Pedimyia kozlovi Rohdendorf, 1925: 83 [♀].Phrosinella (Asiometopia) kozlovi: Verves and

Khrokalo, 2006: 102 [in key].Phrosinella kozlovi: Fan and Pape, 1996: 241, [faunistic];

Pape, 1996: 124 [catalogue].Asiometopia kozlovi: Rohdendorf, 1935: 127 [in key];

1971b: 129 [♀; redescription, faunistic]; Rohdendorf and Verves, 1980: 498 [description of ♂, faunistic]; Verves, 1986: 88 [catalogue]; 1990: 553 [in key]; Chao and Zhang, 1998: 1532 [♂♀ in key, faunistic].

Holotype (female): China, Gansu, Gashun Gobi, Shibendu-Shigusa N Dunhuang [“nördlich von Sadshzhou”]. Holotype in ZIN.

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Synonym: Asiometopia burjata Rohdendorf, 1935: 128 [♂]; 1971:

129 [♀ & drawing of ♂ fore tarsus]. Holotype (male): Russia, Buryatia, Chikoy River,

Dureny. Deposited in ZIN.Distribution: Palearctic: Turkey (firstly recorded),

China (Gansu), Mongolia, Russia (Buryatia).Habits: Psammophilous species.Phrosinella (Asiometopia) tadzhika (Rohdendorf,

1935)Asiometopia tadzhika Rohdendorf, 1935: 127 [in key];

1971b: 130 [♂♀]. Asiometopia tadzhika: Charykuliev, 1965: 16 [habits];

Verves, 1986: 88 [catalogue]; Chao and Zhang, 1998: 1532 [♂♀ in key, faunistic].

Phrosinella tadzhika: Fan and Pape, 1996: 241, [faunistic]; Pape, 1996: 125 [catalogue]; Kara and Pape, 2002: 292 [faunistic]; Koçak and Kemal, 2013: 137; 2015: 347 [faunistic].

Holotype (male): Uzbekistan, Bukhara Region, between Jargak & Sarat Lyalyk. Deposited in ZIN.

Distribution: Palearctic: China (Neimenggu, Xinjiang); Tajikistan; Turkmenia; Turkey (Konya, Tokat); Uzbekistan.

Habits: Psammophilous species; developmental time in burrows of sphecid wasp Bembicinus asiaticus from neonatal 1st stage larva to eclosion of adult flies in June–July occurs during 13–18 days. ♀ bears the larvae at victim of wasp (Cicadellidae) when host carries paralyzed leafhopper into burrow (Charykuliev, 1965).

Phrosinella (Asiometopia) ujugura (Rohdendorf, 1935)

Asiometopia ujugura Rohdendorf, 1935: 127 (in key); 1971b: 130 [♂♀].

Asiometopia ujugura: Charykuliev, 1965: 16; [habits & faunistic]; Myartzeva, 1972b: 79 [habits]; Verves, 1986: 88 [catalogue]; Chao and Zhang, 1998: 1532 [♂♀ in key, faunistic].

Phrosinella ujugura: Fan and Pape, 1996: 241 [faunistic]; Pape, 1996: 125 [catalogue]; Kara and Pape, 2002: 292 [faunistic]; Koçak and Kemal, 2015: 343 [faunistic].

Holotype (male): Uzbekistan, near Khiva. Holotype in ZIN.

Distribution: Palearctic: China (Xinjiang), Tajikistan, Turkmenia, Uzbekistan.

Habits: Larvae were developed in burrow nests of sphecid wasps Tachytes cornigera on paralyzed caterpillars of snout moths  Tegostoma baphialis (Charykuliev, 1965) and Bembix olivacea on freshly killed flies (Myartzeva, 1972b).

Phrosinella (Asiometopia) uzbeka (Rohdendorf, 1935)Asiometopia uzbeka Rohdendorf, 1935: 127 (in key),

1971b: 131 [♂♀].

Asiometopia (s. str.) uzbeka: Verves, 1984: 539 [faunistic]; 1986: 102 [catalogue]; Koçak and Kemal, 2015: 343 [faunistic].

Phrosinella uzbeka: Pape, 1996: 125 [catalogue].Holotype (male): Uzbekistan, Bukhara region, Kumak.

Deposited in ZIN.Distribution: Palearctic: Kazakhstan, Turkmenia,

Uzbekistan.Habits: Psammophilous species.Subgenus Caspiomyia Rohdendorf, 1935Caspiomyia Rohdendorf 1935: 126 (as subgenus of

Asiometopia). Type species: Asiometopia (Caspiomyia) persa Rohdendorf, 1935; by original designation and monotypy.

Phrosinella (Caspiomyia): Verves, 1986: 89 [catalogue]; Zerova et al., 2006: 109 [in key].

Bright colored midsized flies (5.0–8.0 mm in length). Frontal stripe distinctly narrower than one of parafrontalia, blackish, more or less lightly dusted; lower margin of head moderately shortened; abdomen with black dentate bands in hind 0.5–0.7 of 3rd and 4th tergites; 1+2nd and 5th tergites entirely shining black; ♂ fore tarsi without specialized setae; R1 dorsally with numerous black hairs in basal 0.5–0.8; R4+5 haired from base to the middle of its 2nd section. One Palearctic species.

Phrosinella (Caspiomyia) persa (Rohdendorf, 1935) (Figures 17–19)

Asiometopia (Caspiomyia) persa Rohdendorf, 1935: 127 [♂].

Asiometopia (Caspiomyia) persa: Rohdendorf and Verves, 1980: 498 [♀; faunistic]; Verves, 1984: 558 [faunistic]; Verves et al., 1984: 86 [habits & faunistic]; Verves, 1986: 89 [catalogue]; 1990: 553 [in key]; Chao and Zhang, 1998: 1532 [faunistic]; Koçak and Kemal, 2015: 343 [faunistic].

Phrosinella persa: Fan and Pape, 1996: 242 [faunistic]; Pape, 1996: 124 [catalogue]; Szpila, 2010: 52 [1st stage larva].

Phrosinella (Caspiomyia) persa: Verves, 1998: 51 [faunistic]; Zerova et al., 2006: 108 [in key & faunistic].

Holotype (male): Iran: Malyi Ashur I. [= Aşyr Ada I.]. Deposited in ZIN.

Distribution: Palearctic: China (Neimenggu), Iran, Mongolia, Romania, Ukraine.

Habits: Adult flies prefer sandy areas with Phragmites bushes on shores of sea and others reservoirs (Verves et al., 1984).

Subgenus Euhilarella Townsend, 1915Euhilarella Townsend, 1915: 22. Type species:

Gymnoprosopa fulvicornis Coquillett, 1895; by original designation.

Phrosinella (Euhilarella): Verves, 1980: 30.Phrosinella: Allen, 1926: 68; Shewell, 1987: 1185 [in

key]; Pape and Dahlem, 2010: 1318.

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Dark colored midsized flies (5.0–8.0 mm in length). Frontal stripe distinctly more widened than one of parafrontalia; lower margin of head moderately shortened; abdominal tergites with dark spots and stripes; 2nd

tarsomere of ♂ fore tarsi with a tuft of long hairs; R1 bare dorsally; R4+5 haired from base to the middle of its 2nd section. Seven species in Holarctic.

Habits: The flies occupy the ground-subterranean stratum, being found in the adult stage largely on barren ground; in and about the holes of burrowing Hymenoptera, rarely on low foliage. Larvae develop in hymenopteran ground nests in sandy areas (Allen, 1926; Evans and O’Neil, 1988).

Phrosinella (Euhilarella) aldrichi Allen, 1926Phrosinella aldrichi Allen, 1926: 75 [♂♀].Phrosinella aldrichi: Criddle, 1927: 47 [faunistic];

Pape, 1996: 123 [catalogue]; Pape and Dahlen, 2010: 1331 [faunistic].

Holotype (male): USA, Idaho, Lewiston. Deposited in SMNS.

Distribution: Nearctic: Canada (Alberta, British Columbia, Ontario), Mexico (Sonora), USA (California, Idaho, Iowa, Nevada, Washington).

Phrosinella (Euhilarella) aurifacies Downes, 1985Phrosinella aurifacies Downes, 1985: 269 [♂♀].Phrosinella aurifacies: Spofford and Kurczewski, 1985:

273 [habits]; Hager and Kurczewski, 1986: 453 [habits]; Kurczewski and Spofford, 1986: 13 [habits]; Spofford et al., 1986: 350 [habits]; Kurczewski, 1989: 397 [habits]; Spofford et al., 1989: 256 [hosts]; Spofford and Kurczewski, 1990: 731 [habits]; Kurczewski, 1991: 300 [habits]; Spofford and Kurczewski, 1992: 995 [hosts]; Pape, 1996: 123 [catalogue]; O’Hara et al., 2000: 172 [faunistic].

Holotype (male): USA, Michigan, Shiawassee Co., Rose Lake Conservation Area. Deposited in USNM.

Phrosinella fulvicornis [misidentification: not Gymnoprosopa fulvicornis Coquillett, 1895]: Brown, 1934: 249 [faunistic]; Cole and Lovett, 1921: 303 [faunistic]; Cole, 1923: 205 [faunistic]; Criddle, 1928: 92 [faunistic]; Knowton, 1936: 237 [faunistic]; Ristich, 1956: 271 [habits]; Evans and Lin, 1959: 115 [habits]; Krombein and Kurczewski, 1963: 139 [faunistic, habits]; Evans, 1966b: 35 [habits]; 1970: 451 [habits]; 1973: 888 [habits]; 1975: 888 [habits]; Peckham, 1977: 823 [habits]; Evans et al., 1980: 865; O’Hara et al., 2000 [faunistic, habits], Pickering, 2011: 1627, 1631.

Distribution: Nearctic: Canada (Alberta, Labrador, Manitoba, New Brunswick, Ontario, Quebec, Yukon Territory), USA (Colorado, Connecticut, Florida7, Illinois, Iowa, Kansas, Massachusetts, Michigan, Minnesota,

Mississippi, Missouri, Nebraska, New Jersey, New York, Ohio, Oregon, Pennsylvania, South Carolina, South Dakota, Texas, Wyoming8).

Habits: This species is one of the most common sarcophagids in sandy xerophytic places in North America, and it is a “hole searcher”; therefore, the larviposition response could be initiated in situations other than a wasp bringing in its prey (Downes, 1985). Both sexes feed on honeydew of aphid colonies in psammophilic xerophilous vegetation. Courtship and compound mating behavior are highly specific for this species (Spofford and Kurczewski, 1985). Gravid ♀♀ zigzag several millimeters above the surface of sand and sometimes land to investigate temporarily closed entrances into wasps’ ground nests. Then ♀♀ larviposit one or several maggots on the depression over the closed entrance, and larvae wriggle quickly through the entrance stopper into the burrow (Spofford et al., 1986). In another way, gravid flies pursue wasps lugging prey and enter into open burrows immediately after hosts (Kurczewski and Spofford, 1986; Kurczewski, 1989, 1991). Sometimes flies larviposit into the burrow before the beginning of wasp foraging (Peckham, 1977). Larvae develop in ground nests in sandy areas of different sphecid and occasionally pompiloid wasps, where they feed at host prey (paralyzed or freshly killed insects or spiders), viz:

Pompilidae. Episyron quinquenotatus at paralyzed orb weaver spiders, Araneidae (Spofford et al., 1989; Spofford and Kurczewski, 1992).

Sphecidae. 1. At paralyzed adult beetles (Buprestidae): Cerceris fumipennis (Spofford and Kurczewski, 1992).

2. At paralyzed caterpillars (Geometridae, Gelechiidae, Noctuidae, Pyralidae): Ammophila harti (Hager and Kurczewski, 1986).

3. At freshly killed adult flies: Crabro argusinus (Spofford and Kurczewski, 1992), C. monticola (Evans et al., 1980; Spofford and Kurczewski, 1992), C. virgatus (Evans et al., 1980), Lindenius armaticeps (Spofford and Kurczewski, 1992); L. columbianus (Spofford and Kurczewski, 1992; Lucas, 1997); Oxybelus bipunctatus (Spofford and Kurczewski, 1992), O. emarginatus (Krombein and Kurczewski, 1963; Peckham, 1977), O. subulatus (mainly Therevidae: Peckham, 1977; Spofford and Kurczewski, 1992); O. uniglumis (mainly Muscidae: Evans, 1970; Lucas, 1997).

4. At freshly killed solitary and honey bees: Phylanthus crabroniformis (Evans, 1970); P. gibbosus (Evans, 1970; Spofford and Kurczewski, 1992; Lucas, 1997); P. lepidus (Evans, 1970; Spofford and Kurczewski, 1992); P. politus (Spofford and Kurczewski, 1992); P. solivagus (Ristich,

7 after Krombein and Kurczewski, 19638 after Evans, 1970

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1956; Spofford et al., 1989; Spofford and Kurczewski, 1992); P. zebratus (Evans, 1970).

5. At paralyzed nymphs and adult Orthoptera: Tachysphex acutus (Melanoplus spp.: Kurczewski, 1964, 1989, 1991); T. antennatus (Acridoidea: Spofford and Kurczewski, 1992); T. intermedius (Acridoidea: Spofford and Kurczewski, 1992); T. similis (Acridoidea: Kurczewski, 1964), T. tarsatus (Acridoidea: Kurczewski, 1964; Spofford and Kurczewski, 1992), T. terminatus (Acridoidea: Krombein and Kurczewski, 1963; Kurczewski, 1964; Spofford and Kurczewski, 1992), Tachytes mergus (Tridactylus spp.: Krombein and Kurczewski, 1963), T. parvus (Tetrix ornata: Kurczewski and Spofford, 1986; Spofford and Kurczewski, 1992).

6. At paralyzed or freshly killed adult Homoptera: Clitemnestra bipunctata (Cicadoidea, Psiloidea: Spofford and Kurczewski, 1992; Evans and O’Neill, 2007); Gorytes canaliculatus (Cicadellidae: Evans, 1970; Spofford and Kurczewski, 1992); Sphecius speciosus (great-sized Cicadidae species from genera Diceroprocta,  Magicicada,  Neocicada,  Quesada, and Tibicen: Kurczewski and Spofford, 1987). 

7. At paralyzed Hemiptera: Plenoculus davisi (tarnished plant bug, Lygus lineolaris: Spofford and Kurczewski, 1992).

Phrosinella (Euhilarella) fulvicornis (Coquillett, 1895)Gymnoprosopa fulvicornis Coquillett, 1895: 106 [♂♀].Hilarella fulvicornis: Coquillett, 1897: 128 [notes];

Barber, 1915: 187 [habits]; Britton, 1920: 192 [faunistic]; Brimley, 1922: 23 [faunistic]; Strickland, 1938: 175, [faunistic]; Davis and Turner, 1978: 111 [faunistic].

Phrosinella fulvicornis: Allen, 1924: 92 [notes]; 1926: 70 [redescriptions of ♂♀, habits, faunistic]; Rowe, 1932 [faunistic]; Krombein, 1936: 93 [habits]; Séguy, 1941: 305 [hosts]; Strandtmann, 1945: 305–308 [habits]; Krombein, 1955: 223 [habits]; Downes, 1965: 938 [catalogue]; Kurczewski and Harris, 1968: 81 [habits]; Miller and Kurczewski, 1973: 365 [habits]; Peckham et al., 1973: 647 [habits]; Miller and Kurczewski, 1975: 82 [habits]; Peckham, 1977: 823 [habits]; Downes, 1985: 271 [taxonomy & faunistic]; Spofford et al., 1989: 257 [hosts]; Pape, 1996: 123 [catalogue]; Pickering, 2011: 1631, 1653, 1657, 1660, 1661, 1685, 1693, 1712, 1713, 1721, 1724, 1725 [habits].

Holotype (male): USA, New Jersey, Avalon. Holotype in USNM.

Distribution: Nearctic: Canada (New Brunswick: Prince Edward I.)9, USA (Connecticut, Florida, Kansas, Massachusetts, Michigan, Mississippi, New Jersey, New York, Pennsylvania, South Carolina, Texas).

Habits: Larvae develop in ground nests of different sphecid wasps, where they feed at host prey (paralyzed or freshly killed insects), viz:

1. At freshly killed adult flies: Bembix americana (Barber, 1915; Krombein, 1936; Evans, 1966a; Peckham and Peckham, 1898), Bembix spinolae  (Allen, 1926; Séguy, 1941; Evans, 1957), Lindenius armaticeps (Miller and Kurczewski, 1973, 1975), Oxybelus bipunctatus (Kurczewski and Harris, 1968; Peckham et al., 1973), O. emarginatus (Krombein and Kurczewski, 1963; Peckham, 1977), O. quadrinotatus (Séguy, 1941), O. subulatus (Peckham et al., 1973; Peckham, 1977), O. uniglumis (Allen, 1926; Strandtmann, 1945; Krombein, 1955; Evans and Lin, 1959; Evans, 1970; Peckham et al., 1973).

2. At freshly killed adult solitary wasps, bees, and honey bees: Philanthus albopilosus (Spofford et al., 1989), P. crabroniformis (Spofford et al., 1989), P. gibbosus (Evans and O’Neil, 1988), P. politus (Evans and O’Neil, 1988), P. pulcher (Evans, 1966b, 1970), P. sanbornii (Evans and O’Neil, 1988), P. solivagus (Ristich, 1956), P. zebratus (Spofford et al., 1989).

3. At paralyzed nymphs and adults of Orthoptera: Tachysphex acutus (Kurczewski, 1964), Tachysphex mergus (Krombein and Kurczewski, 1963), T. similis, T. tarsatus (Kurczewski, 1964), T. terminatus (Krombein and Kurczewski, 1963; Kurczewski, 1964; Kurczewski and Harris, 1968).

4. At paralyzed or freshly killed nymphs and adults of Homoptera, Cicadoidea: Alysson melleus (Evans, 1966a), Gorytes canaliculatus (Evans, 1966a, 1970, 1973)

5. At freshly killed adult and nymphs of Hemiptera, Homoptera, adult Hymenoptera and Diptera: Lindnerius columbianus (Miller and Kurczewski, 1973; Pickering, 2011).

Phrosinella (Euhilarella) fumosa Allen, 1926 Phrosinella fumosa Allen, 1926: 74 [♂♀].Phrosinella fumosa: Batra, 1980: 509 [habits]; Evans,

1960: 123 [habits]; Downes, 1965: 938 [catalogue]; Evans, 1970: 451[habits]; 1971: 500 [habits]; Downes, 1985: 271 [faunistic]; Spofford et al., 1989: 257, 259 [hosts]; Pape, 1996: 123 [catalogue]; Pickering, 2011: 1674, 1721 [habits].

Holotype (male): USA, Virginia: Falls Church. Deposited in USNM.

Distribution: Nearctic: USA (Delaware, Maryland, Massachusetts, Michigan, New Jersey, New York, Virginia).

Habits: Larvae develop in ground nests of different sphecid wasps and solitary bees, where they feed at host prey (paralyzed or freshly killed insects; nectar and blossom dust in nests of Colletidae), viz:

1. At freshly killed adult flies: Crabro argusinus (Evans, 1960).

9 after Downes, 1985

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2. At freshly killed adult solitary wasps, bees, and honey bees: Philanthus albopictus (Pickering, 2009), P. crabroniformis, P. pulchellus, P. zebratus (Spofford et al., 1989).

3. At paralyzed adult bees: Cerceris fumipennis (Pickering, 2009).

4. At paralyzed nymphs and adults of Homoptera, Cycadoidea: Gorytes canaliculatus (Evans, 1966, 1970).

5. At nectar and blossom dust: Colletes inaequalis, C. thoracicus, C. validus (Batra, 1980).

Phrosinella (Euhilarella) pilosifrons Allen, 1926Phrosinella pilosifrons Allen, 1926: 76 [♂♀].Phrosinella pilosifrons: Criddle, 1927: 47 [faunistic];

Downes, 1965: 938 [catalogue]; Evans, 1966: 35 [habits]; 1970: 465 [habits]; Bohart and Grissell, 1975: 19 [habits]; Spofford et al., 1989: 257 [hosts]; Pape, 1996: 125 [catalogue]; Valenti et al., 1997: 167 [faunistic]; Pickering, 2009: 1722, 1726 [habits].

Holotype (male): USA, Washington, Hood River. Deposited in USNM.

Distribution: Nearctic: Canada (British Columbia, Northwest Territories, Ontario10); USA (California, Oregon, Washington, Wyoming11).

Habits: Larvae are specialized kleptoparasites in ground nests of sphecid wasps from the genus Philanthus [P. barbiger (Evans and O’Neil, 1988), P. crabroniformis (Evans, 1970; Evans and O’Neil, 1988; Pickering, 2009), P. nitens (Evans, 1966, 1970), P. pulcher (Evans, 1970; Evans and O’Neil, 1988), P. zebratus (Evans, 1970; Bohart and Grissell, 1975; Evans and O’Neil, 1988; Pickering, 2009)], where they feed at different freshly killed solitary wasps and bees.

Phrosinella (Euhilarella) sannio (Zetterstedt, 1838) (Figure 16)

Tachina sannio Zetterstedt, 1838: Insecta Lapp.: 636 [♀]

Phrosinella sannio: Pape, 1986: 309 [type revised]; Rognes, 1986: 4 [redescriptions of ♂♀]; Pape, 1987: 66 [redescriptions of ♂♀]; Verves, 1990: 552 [in key]; Pape, 1996: 125 [catalogue]; Verves, 1998: 51 [faunistic]; Verves and Khrokalo, 2006: 101 [in key]; Zerova et al., 2006: 108 [in key]; Szpila, 2010: 53 [1st stage larva]; Pohjoismäki and Kahanpää, 2014: 388 [faunistic].

Holotype (female): Sweden, Åsele Lappmark, Åsele. Deposited in MZLU.

Synonyms: Tachina pilitarsis Zetterstedt, 1844: 1021[♂].Tachina pilitarsis: Pape, 1986: 308 [type revised].Holotype (male): Norway, Verdal, Østre Nes.

Deposited in MZLU.

Phrosinella septentrionalis Rohdendorf, 1970: 643 [♀]. Holotype (female): Norway, Dovre. Deposited in ZIN.Phrosinella septentrionalis: Rohdendorf, 1971a: 137

[redescription of ♀]; 1971b: 451 [♀]; Verves, 1980: 30 [description of ♂]; 1986: 102 [catalogue].

Distribution: Nearctic: Canada (British Columbia, Yukon Territory); USA (Alaska). Palearctic: Finland; Norway; Russia (Karelia, Leningrad Region, Taymyr, Kamchatka); Sweden; Ukraine (Cherkasy Region).

Phrosinella (Euhilarella) talpina Reinhard, 1961Phrosinella talpina Reinhard, 1961: 212 [♂].Phrosinella talpina: Downes, 1965: 938 [catalogue];

Pape, 1996: 125 [catalogue].Holotype (male): USA, California, San Diego Co.

Deposited in CNC.Distribution: Nearctic: USA (California).Subgenus Phrosinella s. str.Bright colored small or midsized flies (3.5–7.5 mm in

length). Frontal stripe as wide or slightly more widened than one of parafrontalia; lower margin of head strongly shortened; ♂ fore tarsi not modified, without specialized setae; R1 bare dorsally; abdominal tergites silvery dusted, with distinct hind dark spots and (or) stripes. Nine species in Palearctic and northern part of Oriental (Jammu & Kashmir) regions.

Phrosinella (s. str.) fedtshenkoi (Rohdendorf, 1925)Pedimyia fedtshenkoi Rohdendorf, 1925: 82 [♂♀]Phrosinella fedtshenkoi: Charykuliev, 1964: [faunistic];

Charykuliev and Myartzeva, 1964: [habits & faunistic]; Rohdendorf, 1971a: 448 [redescriptions of ♂♀, faunistic]; 1971b: 134 [redescriptions of ♂♀, faunistic]; Myartzeva, 1972a: 101 [habits & faunistic]; Verves, 1986: 102 [catalogue]; Pape, 1996: 123 [catalogue]; Verves, 2001: 236 [faunistic]; Nandi, 2002: 123–125 [redescription of ♂, faunistic].

Holotype (male): Uzbekistan, Samarkand Region, Zheravshan. Holotype in ZIN.

Distribution: Palearctic: Tajikistan; Turkmenia; Uzbekistan. Oriental: India (Jammu & Kashmir).

Habits: Larvae were developed in the nests of sphecid wasps Ammophila sp. (Myartzeva, 1972a) and A. gracillima on paralyzed caterpillars Pericyma albidentaria (Charykuliev and Myartzeva, 1964).

Phrosinella (s. str.) gussakovskii Rohdendorf, 1971Phrosinella gussakovskii Rohdendorf, 1971a: 447 (in

key), 448 [♂♀].Phrosinella gussakovskii: Rohdendorf, 1971b: 134

[redescriptions of ♂♀, faunistic]; Verves, 1986: 102 [catalogue]; Pape, 1996: 124 [catalogue].

10 after Criddle, 192711 after Evans, 1966

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Holotype (male): Tajikistan, Staraia Pristan at Wakhsh River near Dzhilicul. Deposited in ZIN.

Distribution: Palearctic: Tajikistan; Turkmenia.Phrosinella (s. str.) mongolica Rohdendorf & Verves,

1980Phrosinella mongolica: Rohdendorf and Verves, 1980:

501 [♂].Phrosinella mongolica: Verves, 1986: 102 [catalogue];

1990: 553 [in key]; Pape, 1996: 124 [catalogue].Holotype (male). Mongolia, Kobd aimak, 15 km S

Bulgan. Deposited in ZIN.Distribution: Palearctic: Mongolia.Phrosinella (s. str.) nasuta (Meigen, 1824) (Figures 14

and 15)Tachina nasuta Meigen, 1824: 374. Phrosinella nasuta: Séguy, 1941: 304 [redescriptions

of ♂♀, habits, faunistic]; Venturi, 1947: 126 [in key, faunistic]; 1960: 82, [redescriptions of ♂♀, faunistic]; Brander, 1964: 51 [faunistic]; Rohdendorf, 1970: 643 [in key]; 1971a: 447 [in key], 449 [redescriptions of ♂♀, faunistic]; 1971b: 135 [redescriptions of ♂♀, faunistic]; Mihályi, 1979: 85 [redescriptions of ♂♀ faunistic]; Verves, 1980b: 923, [faunistic]; 1986: 102 [catalogue]; Pape, 1987: 68, [faunistic]; Verves, 1990: 553 [in key]; Rognes, 1993: 43 [faunistic]; Fan and Pape, 1996: 241 [faunistic]; Pape, 1996: 124 [catalogue]; Povolný, 1997: 98 [faunistic]; Povolný and Verves, 1997: 108 [redescriptions of ♂♀, habits, faunistic]; Pape and Merz, 1998: 338 [faunistic]; Verves, 1998: 5 [faunistic]; 2000: 123 [faunistic]; Papp, 2001: 429 [faunistic]; Kara and Pape, 2002: 291 [faunistic]; Verves, 2003: 44 [faunistic]; Verves and Khrokalo, 2006: 101 [in key, habits, & faunistic]; Zerova et al., 2006: 108 [in key]; Draber-Mońko, 2007: 232 [faunistic]; Kaczorowska, 2009: 63 [habits & faunistic]; Raffone, 2009: 107 [faunistic]; Szpila, 2010: 51 [1st stage larva]; Koçak and Kemal, 2013: 137 [faunistic]; Koçak and Kemal, 2015: 347 [faunistic].

Holotype (male): Europe (probably Germany). Type material lost.

Synonyms:Miltogramma strenua Perris, 1852: 209. Holotype (male): France, Landes. Deposited in

MNHM.Brachicoma metopiella Rondani, 1859: 205 [♀]

Brachicoma metopiella: Venturi, 1947: 126 [taxonomy]; Pape, 1988: 10 [holotype revised]

Holotype (female): Italy, Parma. Deposited in MZUF. Phrosina argyrina Robineau-Desvoidy, 1983: 102,

Holotype: France: environs of Paris. Type material lost.Phrosinella pictipennis Rohdendorf, 1971a: 447 [in

key], 450 [♂♀] Phrosinella pictipennis: Rohdendorf, 1971b: 136

[redescriptions of ♂♀, faunistic]; Kolomyietz, 1979: 139 [faunistic]; Artamonov, 1980: 32 [faunistic]; Verves, 1986: 103 [catalogue].

Holotype (male): Russia, Primorye, Grossevichi near Soviet Havan. Deposited in ZIN.

Distribution: Palearctic: Armenia; Austria; Azerbaijan; China (Neimenggu); Czech Republic; Finland; France (mainland and Corsica); Germany; Hungary; Iran; Italy (mainland); Kazakhstan; Kyrgyzstan; Libya; Macedonia; Moldova; Mongolia; Russia (European part: Ivanovo, Leningrad, Lipetzk, Volgograd, Voronezh; West Siberia: Tomsk; East Siberia: Chita, Krasnoyarsk; Far East: Khabarovsk, Magadan, Primorye); Slovakia (Povolný, 1997); Spain; Sweden; Switzerland; the Netherlands; Turkey; Ukraine.

Habits: Very psammophilous and xerophilous species (Séguy, 1941; Zerova et al., 2006; Kaczorowska, 2009). Hosts unknown: the data of Povolný and Verves (1997: 109) about larval kleptoparasitism of P. nasuta in nests of sphecoid wasps Nitella [or Bembex] spinolae and Oxybelus quadrimaculatus really refer to North American species P. fulvicornis, as was noted by Séguy (1941: 305).

Phrosinella (s. str.) nigripunctata Rohdendorf, 1971Phrosinella nigripunctata Rohdendorf, 1971a: 447 [in

key], 449 [♀]. Phrosinella nigripunctata: Rohdendorf, 1971b: 136

[redescription, faunistic]; Rohdendorf and Verves, 1980: 500 [faunistic]; Verves, 1986: 103 [catalogue]; 1990: 552 [in key]; Pape, 1996: 124, [catalogue].

Holotype (female): Mongolia: Gobi-Altay aimak, Shargyn Gobi, 40 km SW Altay. Deposited in ZIN

Distribution: Palearctic: Mongolia; Russia (West Siberia: Tomsk).

Phrosinella (s. str.) sibirica Rohdendorf, 1971Phrosinella sibirica Rohdendorf, 1971a: 448 [in key],

451 [♀].Phrosinella sibirica: Rohdendorf, 1971b: 138

[redescription of ♀, faunistic]; Verves, 1986: 103 [catalogue]; 1990: 553 [in key]; Pape, 1996: 125 [catalogue].

Holotype (female): Russia: Altay, Kosh-Agach, Chuia steppe, 1750 m a.s.l. Deposited in ZIN.

Distribution: Palearctic: Russia (West Siberia: Altay, Krasnoyarsk, Tomsk).

Phrosinella (s. str.) similis (Rohdendorf, 1925)Pedimyia similis Rohdendorf, 1925: 83 [♂♀]Phrosinella similis: Rohdendorf, 1971a: 477 [in

key], 452 [redescriptions of ♂♀, faunistic]; 1971b: 139 [redescriptions of ♂♀, faunistic]; Charykuliev, 1986: 141 [faunistic]; Verves, 1986: 103 [catalogue]; Pape, 1996: 125 [catalogue].

Holotype (male): Uzbekistan: Katatkurgan. Deposited in ZIN.

Distribution: Palearctic: Kyrgyzstan; Tajikistan; Turkmenia; Uzbekistan.

Phrosinella (s. str.) simillima Rohdendorf, 1971 Phrosinella simillima Rohdendorf, 1971a: 452 [♀]

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Phrosinella simillima: Rohdendorf, 1971b: 139 [redescription of ♀, faunistic]; Verves, 1986: 103 [catalogue]; Pape, 1996: 125 [catalogue].

Holotype (female): Turkmenia: Krasnovodsk. Deposited in ZIN.

Distribution: Palearctic: Turkmenia.Phrosinella (s. str.) zarudnoji Rohdendorf, 1971Phrosinella zarudnoji Rohdendorf, 1971a: 448 [in key],

452 [♂♀]Phrosinella zarudnoji: Rohdendorf, 1971b: 140

[redescriptions of ♂♀, faunistic]; Rohdendorf and Verves, 1980: 502 [faunistic]; Verves, 1986: 103 [catalogue]; 1990: 553 [in key]; Pape, 1996: 125 [catalogue]; Koçak and Kemal, 2015: 347 [faunistic].

Holotype (male): Iran: Beluchistan, Bagu-Kelat. Deposited in ZIN

Distribution: Palearctic: Azerbaijan; Iran; Kazakhstan; Mongolia; Tajikistan; Turkmenistan.

Nomenclatural acts: This work and the nomenclatural acts it contains have been registered in ZooBank. The ZooBank Life Science Identifier (LSID) for this publication is:  http://zoobank.org/urn:lsid:zoobank.org:pub:A87957B6-B51C-4B90-ADCA-36F0EAD5572E.

AcknowledgmentsWe are grateful to Prof A Freidberg, Zoological Museum of Tel Aviv University, Israel, and Mr Adrian Pont, Natural History Museum, London, United Kingdom, for their kindness in sending material; and Dr K Szpila, Faculty of Biology and Environmental Protection, Nicolaus Copernicus University, Torun, Poland, for preparation of photos for Figures 1–6.

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