Revision of the Lispe longicollis-group (Diptera, Muscidae)

Abstract The Lispe longicollis species-group is revised. Lispe ethiopica sp. n. is described. The following 3 new synonyms are established: Lispe assimilis Wiedemann, 1824 (syn: cyrtoneurina Stein, 1900 and modesta Stein, 1913); Lispe manicata Wiedemann, 1830 (syn: forficata Kurahashi & Shinonaga, 2009). Female of Lispe microptera Seguy, 1937 is described for the first time. Identification key for known Eurasian and African species is given.


Introduction
The Lispe longicollis species-group was proposed by Hennig (1960) based on the characteristic shape of a vein M which is distinctly curved forward at apex. The species of this group also share these additional characters: t3 with submedian av, ad and pd setae; abdomen with large, more or less fused trapezoid spots; frontal triangle narrow; dc 2+4, usually only 2 posterior pairs are strong (in L. glabra Wiedemann and L. manicata Wiedemann dc setae should be described as 0+2, but careful examination shows that 4 anterior pairs of minute dc setulae are normally present but broken in most specimens). Hennig divided the group into two subgroups.
Subgroup 1 included the species with ventral seta on t2 and consisted of L. longicollis Meigen, 1826 (S Palaearctic) and L. cilitarsis Loew, 1856 (North Africa and Near East). In this paper another three species are added to subgroup 1: L. microptera Seguy, 1937 known from Pakistan and India and two Afrotropical species, L. barbipes Stein, 1908 and the here described L. ethiopica sp. n. The main subgroup character is the presence of fine hairs on the meron above the hind coxa; other characters: t2 with ventral seta (except for L. microptera); male hind basitarsus curved and bears long ventral hairs (except for L. longicollis); halves of cercal plate of a subquadrate shape and strongly conjoined with each other (less so in L. longicollis); the flies inhabit banks of salted to fresh water.
Hennig's subgroup 2 included widespread L. assimilis Wiedemann, 1824 and African L. nuba Wiedemann, 1830 which lack the ventral seta on t2. In this paper another three Oriental species are added to the subgroup 2: L. glabra Wiedemann, 1824, L. manicata Wiedemann, 1830 and L. pacifica Shinonaga & Pont, 1992. Subgroup 2 is characterized as follows: meron bare; t2 without ventral seta; male hind basitarsus unmodified; halves of cercal plate of subtriangular shape and less conjoined with each other; the flies inhabit banks of fresh water only.
This revision considers Palaearctic, Afrotropical and Oriental species of the Lispe longicollis group. The group is absent in the Nearctic region, there are two Australian species not seen by the author, namely Lispe weschei Malloch, 1922 andLispe xenochaeta Malloch, 1923, which also belong to the same group.

Material and methods
The majority of the specimens studied are deposited in the Zoological Museum of Lomonosov Moscow State University, Moscow, Russia, in this case not indicated in text. Other collections are abbreviated as follows: BMNH Natural History Museum, London, UK; TAUI Tel-Aviv University, Israel; ZIN Zoological Institute of the Russian Academy of Sciences, St Petersburg, Russia; ZMHU Museum für Naturkunde, Humboldt-Universität zu Berlin, Berlin, Germany.
The abbreviation for the tarsi as tar followed by a pair of digits separated by a hyphen was proposed by Vikhrev (2011): the first digit (1 to 3) gives the leg number and the second digit (1 to 5) the number of the tarsal segment. For example, tar1-4 = 4th segment of fore tarsus; tar3-1 = hind basitarsus.
Geographical coordinates are given in the Decimal Degrees format. Synonymies are listed only for the species to which the new synonymies are considered, for full lists of synonymies see the regional Diptera Catalogues: Pont 1977Pont , 1980Pont and 1986 Identification key to Eurasian and African species of Lispe longicollis-group t2 with 1 p(pd) seta. ♂ (Fig. 3): f3 with 4-5 fine long pv in basal half and 1(2) av in basal 1/3; tar3-1 slightly laterally compressed and outward curved, with waved ventral setulae more dense at base and at apex; tar3-2 with waved v setulae; cercal plate as in Fig. 8. ♀: f3 on av surface usually with a short av seta before middle (in some specimens this seta absent  Distribution. S Palaearctic, Afrotropical and Oriental regions, Australia, Oceania. Synonymies. The taxonomy of L. assimilis was considered by Shinonaga and Pont (1992). In that paper the synonymy of L. quadrilineata, L. incerta and L. inexpectata with L. assimilis was established and the related Oriental species with long ventral hairs on mid femur was described as Lispe pacifica Shinonaga & Pont, 1992, it was shown that L. assimilis in the sense of old authors is L. pacifica, while later authors followed this misinterpretation.
Lispe cyrtoneurina Stein, 1900 -syn. nov. of L. assimilis. Stein's (1900) original description completely fits L. assimilis, the only difference found is 3 (instead of 4) post dc. The male lectotype of L. cyrtoneurina (stored in Genoa, Museo Civico di Storia Naturale di Genova) was reexamined by Adrian Pont. The lectotype is in poor condition, damped and mostly squashed; 4 post dc; everything else fit L. assimilis (Pont, pers. com. and unpublished notes).
Lispe modesta Stein, 1913 -syn. nov. of L. assimilis. The very short Stein's (1913) description fits L. assimilis. Examined by me specimens from Asia and Africa were found similar, the specimens from Ziway Lake in Ethiopia are especially interesting as it is the type locality of L. modesta. In a later paper (Stein, 1918: 175) Stein himself listed L. assimilis from Rangoon (Yangoon, Myanmar) as "Lispe assimilis Wied. var. modesta Stein" and wrote that the male of L. assimilis var. modesta (=L. assimilis in the present interpretation) differs from L. assimilis (=L. pacifica in the present interpretation) only by the absence of long ventral hairs on f2.
Lispe pacifica Shinonaga & Pont, 1992. According to the remark cited above Stein in 1918 started to regard L. assimilis and L. pacifica as variations of the same species. In fact, the separation of these species in female sex is sometimes doubtful and males have similar genitalia. Note also that in both species the pollinosity is very variable: dusting on face, parafacialia and parafrontalia from pure white to deep yellow, dusting on parafrontalia and frontal triangle from weak to strong, dusting of scutum from grey to brown, the colour of the tibiae from almost entirely yellow to almost entirely dark. I would like to report that my observations made around Pai (Thailand, Mae Hong Son province) somewhat support the valid taxonomic status of Lispe pacifica Shinonaga & Pont, 1992. Pai town is so far the only locality I know where both species L. assimilis and L. pacifica were found together at the same time and usually at the same pools. A series of 17 males of L. pacifica and 14 males of L. assimilis were collected. All examined males have distinct characters either of one or other species, with no intermediate specimens recorded. Thus in a sympatric condition no trace of crossbreeding between the two species has been found.
So As it was reported by Pont and Werner (2006): "there must be some doubt as to whether this is actually a syntype, since the locality [of ♂ syntype] was not mentioned by Stein (1908) and is on the coast of Namibia rather than at the eastern edge of the Kalahari desert in Botswana shorter than antenna width, in apical half bare. Palpus yellow. Scutum and scutellum brownish dusted with indistinct vittae, pleura brownish-grey dusted. dc 2+4 (strong-strong+medium-medium-strong-strong); intraalars 1+2; supraalars 1+2, the posterior one weak. Meron with setulae above hind coxa. Wing with vein R4+5 distinctly curved forward. Legs dark, but femora at apex and tibiae in basal half yellow. f1 with a row of 6-7 strong av setae in apical 3/5. t1 with a row of 7-8 short but strong d setae and with submedian p seta. f2 with 2(3) strong, straight and long (2 times as long as femur width) ventral spines; other setae: 1(2) median a seta(e), 2 p at apex. t2 with 1 submedian v seta (which is slightly shifted from true v position onto posterior surface and may be named "pvv seta"). f3 curved; with 1-2 av setae and 1 long but fine pv in basal 1/3, pv preapical present, av preapical absent. t3 with submedian 1 av, 1 ad and 1 pd setae; below middle with a row of 3-4 straight ad; at apical 1/3 with long waved setae on ad to av surface. Hind tarsus modified: tar3-1 elongated, downward curved; with waved ventral setulae, these in apical 1/3 especially long; tar3-2 thickened. Abdomen grey dusted with large lateral black spots, these on tergites 3 and 4 separated by grey vitta, on tergite 5 fused.
Females differs from male as follows: spines on f2 absent; t2 with 2 approximated submedian setae, shorter p-pd and longer v-pv, f3 without av and pv setae (these characters did not mention in Stein's (1908) original description), t3 without long setae at apex; hind tarsus unmodified.    (Pont 1991). In Ethiopia L. cilitarsis seems uncommon and restricted to northern regions in comparison with resembling L. ethiopica sp. nov., so specimens from Africa should be re-examined and so far I regard other Afrotropical records as doubtful. Description. Male, body length 6.5-7.5 mm. Head. Frontal triangle remarkably narrow, brownish in posterior half, yellowishgrey dusted in anterior half. Interfrontalia blackish-brown. Fronto-orbital plate blakish-brown in posterior third, yellowish-grey dusted anteriorly; with 3-5 inclinate and 2 proclinate setae and dense hairs in outer row. Parafacial and cheek whitish dusted, occiput grey, parafacial with a row of hairs. Antenna black, postpedicel short, only 2 times longer than pedicel. Arista with hairs half as long as antenna width. Vibrissae medium strong. Palpi blackish.
Legs black with grey dusting, but knees and base of tibiae yellowish. f1 with a row of pd setae and a row of pv setulae; t1 with submedian p seta. f2 with a seta at middle and 2 pd preapicals; t2 with p seta at middle and av seta in apical third; mid tarsus simple. f3 with 1-2 fine v setulae at base, at apex with 1 short av and 1 short pv; t3 with submedian ad and pd setae and with long fine av at apical third, setulae in the ad row elongated. Hind tarsus modified: tar3-1 dorso-ventrally flattened, distinctly wider than width of t3, on av surface with a dense row of fine curled setulae.
Abdomen with dense whitish dusting; tergites 3 to 5 with a pair of large black fused spots each. Cercal plate and sternite 5 as in Figs 5 and 6.
Female differs from male as follows: body length 7-8 mm; t3 with av seta strong; hind tarsus simple.
Diagnosis. Lispe ethiopica sp.n. is related to L. cilitarsis Loew, 1856 and probably was overlooked due to that resemblance. These two species may be reliably distinguished in both sexes as recommended in the identification key above.
Etymology. Named after the locality of the type series.  Descriptive notes. Body length 8.5-9.5 mm. Wings slightly brownish infuscated. Vein M gradually curved forward from level of crossvein dm-cu, cell r4+5 is almost closed and distance between veins M and R4+5 at wing margin is shorter than crossvein rm. Vein CuA2 not reaching wing margin, extending only to crossvein dm-cu; crossvein dm-cu skewed, it reaches vein M at acute angle of about 45˚. There is a downcurved fold surrounded by long microtrichia along posterior margin of wing between veins M and A2, microtrichia directed outward to the fold. Mid legs: f2 with remarkable row of very dense curled Velcro fastener-like setae in pv position in basal 2/3; t2 in apical 1/4 with a row of long ventral hairs; tar2-1 with a complete row of long curved pv setulae. Male is unmistakable due to modified wings and mid legs. Female differs from female of L. manicata as given in the key.   Distribution. South-East Asia: Cambodia, Malaysia (Borneo), Indonesia (Java), Singapore (Séguy 1948), S Thailand, S Vietnam (Séguy 1948). Synonymy. The illustrations of wonderfully modified male mid tarsi and characteristic genitalia given by Kurahashi and Shinonaga (2009: fig. 1 c-d, 3 a-d) for L. forficata suggest it's conspecifity with L. manicata. So, Lispe manicata Wiedemann, 1830 = Lispe forficata Kurahashi & Shinonaga, 2009, syn. n.

Lispe longicollis
The characters of female of L. forficata were shortly mentioned by Kurahashi and Shinonaga (2009), but these authors did not compare L. forficata with the most related species L. glabra and, in my opinion, the diagnostically important characters were either not mentioned or given erroneously. Therefore I find it necessary to provide the description of the female below.
Description of female. Body length 8.5-9.5 mm.
Thorax. Pleura and dorsolateral area, including postpronotal lobe and notopleuron, densely grey dusted. Disc of scutum mostly subshining brownish-black, with a pair of thinly dusted vittae situated mesad to dc rows, disc of scutellum subshiny brownish-black. ac hairs in 5-6 irregular rows; dc 0+2, only posterior pair strong (hair-like and indistinct posterior prst dc and 1-2 pairs anterior post dc may be found in some specimens); intraalars 0+1, presutural one absent; supraalars 1+2, the posterior one weak. Katepisternal setae 1+2; anepimeron with 9 (8-10) fine hairs, meron and katepimerom bare. Scutellum bare below at apex. Mesothoracic spiracle yellowish. Wings hyaline, slightly brownish infuscated, vein M distinctly curved forward at apex, calypters whitish, halteres yellow. Legs. Legs long, densely grey dusted; black including coxae, but knees and basal half of t2 and t3 brownish-yellow; f1 with a full row of 6 (5-7) pd setae and with a row of fine pv setulae, but in apical half 2-4 setae in pv row are strong (longer than tibial diameter, stronger than setae in pd row); t1 with p seta, preapical d and apical pd and pv; f2 thickened in basal half, f2 with row of 3-5 a-setae in basal 1/3 and 2 pd at apex; t2 with submedian pd; f3 with short av before middle and shorter than setae of ad row, with a full row of ad subequal to femur depth, preapicals: av and pv; t3 with submedian av, ad and pd setae; hind basitarsus with av seta at base.
Abdomen: black with grey pollen. Tergites 1+2 to 5 each with a large blackish spots on dorsal and lateral sides, these spots on tergites 3 and 4 divided at midline by grey dusted interrupted vitta.
Thorax. Scutum and scutellum brownish dusted with a pair of indistinct vittae, pleura grey dusted.
Legs. Femora dark with yellow apex, tibiae yellow in basal half and dark in apical half, tarsi black.
f1 with a complete row of 10-12 pv setae. t1 with submedian p seta. f2 with a row of short a setae in basal half and with 2 pd at apex. t2 with 1 submedian p seta. f3 slightly curved; with a short av seta at basal 1/3 (absent in some specimens) and short pv at apex, av preapical absent. t3 with 1 ad and 1 pd setae at middle. Hind tarsus unmodified.
Abdomen grey dusted with large dorsal black spots separated by anteriorly interrupted grey vitta.
Male differs from female as follows: body length 6.5-7 mm, wing length 5-5.5 mm; f2 with a complete row of fine v setae about as long as femur width; f3 in basal half with 4-5 fine long (2-2.5 femur width) pv setae and 1(2) av in basal 1/3; hind tarsus modified: tar3-1 slightly laterally compressed and outward curved, with waved ventral setulae more dense at base and at apex; tar3-2 with waved ventral setulae; male cercal plate as in Fig. 8. Remarks. Emden (1941) in the key to African Lispe wrote that in L. nuba "front tibiae without a pv". It is not correct, all examined specimens have t1 with pv seta in both sexes, though short in males. Distribution. Widespread in South-East Asia.

ecology
As it was mentioned above, the species of the subgroup 1 of the Lispe longicollis group may be found both on the freshwater and salted basins. L. ethiopica sp.n. was equally common on the freshwater Ziway Lake, on the brackish (2g/l) Langano Lake and on the salt (26g/l) Abijata Lake. L. microptera was collected at brackish lakes and ponds around Jaipur and on the hypersaline (70-300g/l depending on the season) Sambhar Lake. L. longicollis was collected in spring time on the hypersaline Baskunchak Lake, on the seashore salted marshes in Mersin province of Turkey and on the freshwater Titreyen Lake in Antalya province. L. cilitarsis in Israel near Eilat was also found at freshwater of cattle drinking bowl and on a hypersaline lake shore, although in the latter case L. cilitarsis avoided the sites covered with dry salt where only Lispe halophora Becker, 1903 was still present. In contrast to this salt-tolerance, the species of the subgroup 2 of the Lispe longicollis group were observed on freshwater basins only: river banks, rice fields or freshwater lakes/ponds. All species but one prefer open sites, L. manicata seems to be the species of the forest rivers and streams where it was collected in Thailand and Cambodia, the same natural habitat was reported by Kurahashi and Shinonaga (2009) (for L. forficata) in Malaysian Borneo.
The species of the Lispe longicollis group mostly feed on slow moving living prey like Nematocera larvae (Fig. 17), but also were observed feeding on dead arthropods (Fig. 18) or even successfully hunting on a small Diptera imago like Paracoenia, Ephydridae (Fig. 19) or Syntormon, Dolichopodidae.