New and Little Known Species of the Dance-Fly Subgenus Xanthempis Bezzi, Genus Empis L. (Diptera, Empididae), from the Caucasus

ISSN 0013-8738, Entomological Review, 2008, Vol. 88, No. 9, pp. 1115 1126. Pleiades Publishing, Inc., 2008. Original Russian Text I.V. Shamshev, S.Yu. Kustov, 2008, published in Entomologicheskoe Obozrenie, 2008, Vol. 87, No. 4, pp. 776 790. New and Little Known Species of the Dance-Fly Subgenus Xanthempis Bezzi, Genus Empis L. (Diptera, Empididae), from the Caucasus I. V. Shamshev and S. Yu. Kustov All-Russia Institute for Plant Protection, St. Petersburg, Russia Kuban State University, Krasnodar, Russia Received October 30, 2007 Abstract Five new species of the subgenus Xanthempis Bezzi are described from the Caucasus: Empis (Xanthempis) annae sp. n. (Russia: Krasnodar Territory), E. (X.) grichanovi sp. n. (Russia: Krasnodar Territory; Georgia), E. (X.) pseudoconcolor sp. n. (Russia: Krasnodar and Stavropol territories; Georgia: Abkhazia), E. (X.) teberdaensis sp. n. (Russia: Karachay-Cherkessia), and E. (X.) zamotajlovi sp. n. (Russia: Krasnodar Territory and Adygea). The females of E. (X.) alanica Shamshev and E. (X.) kovalevi Shamshev are described for the first time. New data on the distribution of some previously described species are reported. The geographical distribution of Xanthempis is discussed. A key to Xanthempis species from the Caucasus is compiled. DOI: 10.1134/S0013873808090108 The subgenus Xanthempis Bezzi, genus Empis Linnaeus, comprises species characterized by a relatively large size (4 8 mm), yellow coloration of the body, posterior elongation of the head, relatively small eyes separated by a wide frons in both sexes, long antennal scape, strong reduction of the body chaetotaxy, and some other features (Chvála, 1996). These flies usually occur in grass or on leaves of trees and bushes, preferring shaded, rather moist biotopes (Chvála, 1994). Adults of both sexes retain for term of life the mixed feeding, including predation and nectarophagy. In contrast to other representatives of the genus Empis, the species of Xanthempis do not form swarms, and encounter and copulation of sexes occur on the surface of various substrata. Males before copulation make characteristic, species-specific court to females, which is an exception among empidids (Hamm, 1933; Chvála, 1976; Preston-Mafham, 1999). The subgenus Xanthempis is known only from the Palaearctic and includes 51 species (Daugeron, 2000). The maximum number of species (32 or 63%) were recorded in the fauna of Central and Southern Europe, three species, in North Africa (Morocco). It is indicative that the species richness of the group sharply decreases eastwards. Only five species of the subgenus occur in Siberia and the Far East, and only one has been described from Japan. Comparative morphological analysis of Xanthempis species has shown that all the East-Asian members of the group must have descended from E. stercorea L., the only species of the subgenus with the Transpalaearctic distribution range. One species of the genus, E. assalemensis Daugeron, was reported for the fauna of Middle Asia (Daugeron, 2000). However, in our opinion, this indication is not quite exact, since this species was collected at the boundary between Iran and Azerbaijan, i.e., actually, the subgenus Xanthempis was not found in Middle Asia. Ten species of Xanthempis (Shamshev, 1998; Shamshev and Kustov, 2006) have been recorded in the Caucasus. It is of interest that among these species, only E. stercorea occurs in other regions, and the rest are most likely endemics to the Caucasus (Kustov and Shamshev, 2007). Thus, we can assume that the subgenus Xanthempis has the three centers of speciation: central-southern European, Caucasian, and Far Eastern. This assumption can be corroborated by the fact that the degree of similarity between these centers in the number of species is extremely low. In the present communication, five new Caucasian species of the subgenus Xanthempis Bezzi are described. Moreover, additional data on the distribution 1115

1116 SHAMSHEV, KUSTOV of some previously known species are reported and the females of two species are described for the first time. All the species of the subgenus Xanthempis known from the Caucasus, including the new ones described here, are listed in the key below. The only exception is E. trigramma Wiedemann. Engel (1943) reported this species for the fauna of Azerbaijan. However, in our opinion, Engel erroneously identified the material which most likely belonged to E. adriani Chvála (Shamshev, 1998). Nevertheless, commonness of E. trigramma in many regions of European Russia and records of the species in Voronezh Province and the environs of Volgograd point to a high possibility of its distribution in the Northern Caucasus. Shamshev (1998) compiled a key to the Palaearctic species of the subgenus Xanthempis, but the key did not include seven species described later from Europe and Iran (Daugeron, 2000). It should be noted that many species of the subgenus Xanthempis are rather similar in appearance, and their identification can be difficult because of being based on the use of a small number of coloration characters. In such cases, the shape of the phallus, greatly varying in this group of Empididae, is the most reliable (sometimes, the only) criterion for identification of the species. The subgenus Xanthempis includes 14 species known from the Caucasus, which is the maximum among the Caucasian subgenera of the genus Empis (7 species in Empis s. str., 4 in Euempis Frey, 4 in Leptempis Collin, 5 in Pachymeria Stephens, and 5 in Polyblepharis Bezzi). Five species of Xanthempis have been found in the territory of European Russia, except for the Caucasus, and only E. stercorea is shared by the faunas of this region and the Caucasus. MATERIALS AND METHODS The present paper is based on examination of Empididae collections deposited in the Zoological Institute, Russian Academy of Sciences, St. Petersburg (ZIN); Zoological Museum, Moscow State University (ZM MSU); Biological Department, Kuban State University, Krasnodar (KubSU); and University of Oxford (Hope Entomological Collections, University Museum, Oxford, UK). The terminology used in the descriptions mostly follows that proposed by McAlpine (1981), and, for the antennae, by Stuckenberg (1999). The homology of sclerites of the male genitalia follows that by Sinclair (2000). The male terminalia were macerated in hot 85- percent lactic acid for 10 15 minutes and then were placed in glycerol to be used for making descriptions and figures. RESULTS AND DISCUSSION A Key to the Caucasian Species of the Subgenus Xanthempis 1. Scutum entirely yellow... 2. Scutum with dark stripes, spots, or entirely dark... 11. 2. Postsutural supraalar seta present. Occiput entirely yellow. Male: phallus thickened between base and preapical bend... E. kovalevi Shamshev. Postsutural supraalar seta absent. Another combination of characters... 3. 3. Anterior spiracle of thorax yellow... 4. Anterior spiracle of thorax brownish... 7. 4. Occiput (in dorsal view) with large dark diamondshaped spot behind ocellar tubercle...... E. pseudoconcolor sp. n. Occiput (in dorsal view) entirely yellowish or brownish yellow behind ocellar tubercle... 5. 5. Dorsocentral setae strong and rather long, subequal in length to presutural supraalar seta. Male: phallus arcuately curved... E. caucasica Bezzi. Dorsocentral setae fine, shorter than presutural supraalar seta... 6. 6. Male: phallus ansately curved in apical half...... E. grichanovi sp. n. Male: phallus arcuately smoothly curved...... E. assalemensis Daugeron. (Distinctions between females of these species not quite clear yet.) 7. Scutum with 5 6 pairs of dorsocentral setae. Male: phallus with bowl-shaped apex... E. ponti Chvála. Scutum with 7 9 pairs of dorsocentral setae. Male: phallus with beak-shaped apex... 8. 8. Occiput (in dorsal view) behind ocellar tubercle with large dark spot densely covered with grayish pruinosity... E. teberdaensis sp. n. Occiput (in dorsal view) behind ocellar tubercle entirely yellowish or brownish yellow... 9.

NEW AND LITTLE KNOWN SPECIES OF THE DANCE-FLY SUBGENUS 1117 9. Antenna entirely black... E. annae sp. n. Scape and pedicel pale brown, postpedicel and stylus black... 10. 10. Dorsocentral setae fine, shorter than presutural supraalar seta. Scutellum usually with 2 setae. Male: phallus sinuously curved...... E. pavli Shamshev. Dorsocentral setae strong and rather long, subequal in length to presutural supraalar seta. Scutellum usually with 4 setae. Male: phallus arcuately curved... E. caucasica Bezzi. 11. Postsutural supraalar seta present. Scutum with 3 stripes, 2 pairs of spots, or 1 wide stripe occupying entire space between dorsocentral setae... 12. Postsutural supraalar seta absent. Scutum with 1 narrow blackish median stripe... 16. 12. Scutum with 3 wide blackish longitudinal stripes... 13. Scutum with different pattern... 14. 13. Black spot on occiput almost shiny, covered with delicate grayish pruinosity. Abdomen almost entirely (except for segment VIII) covered with pale setae. Male: phallus very long and slender...... E. trigramma Wiedemann. Black spot on occiput covered with dense, pale gray pruinosity. Abdomen entirely covered with black setae. Male: phallus short and relatively thick...... E. adriani Chvála. 14. Scutum with 2 pairs of relatively large, oval, brownish spots (posterior pair of spots usually vague, occasionally inconspicuous). Abdomen covered with black setae... 15. Scutum with 1 wide stripe occupying entire space between dorsocentral setae. Abdomen covered with pale yellow setae... E. zinovjevae Shamshev. 15. Anterior spiracle of thorax brownish. Dorsocentral setae very short and fine (except for 1 anterior pair and 2 prescutellar pairs)... E. adzharica Shamshev. Anterior spiracle of thorax yellow. Dorsocentral setae equally strong and relatively long...... E. zamotajlovi sp. n. 16. Median stripe of scutum reaching margin of prescutellar depression, scutellum yellow. Anepisternum brownish along dorsal margin...... E. alanica Shamshev. Median stripe of scutum running along its entire length and continuing onto anterior surface of scutellum. Anepisternum entirely yellow...... E. stercorea Linnaeus. Empis (Xanthempis) adriani Chvála, 1996 Material. Russia: 5, 3, Northwestern Caucasus, 70 km S of Maikop, Guzeripl, 3 7.VI.1970, V.G. Kovalev; 2, Northwestern Caucasus, Guzeripl, 26 28.V.1970, V.G. Kovalev. In collection of ZM MSU. 2, Krasnodar Territory, Apsheronskii Distr., Kamyshanova Polyana, 14.V.2005, S.Yu. Kustov; 1, Krasnodar Terr., Severskii Distr., near Stavropol skaya St., 13.V.2006, S.Yu. Kustov. In collection of KubSU. Notes. Chvála (1996) described E. adriani from two specimens collected near Kazbegi (Georgia). Later, this species was found in North Ossetia, Adygea, in the territory of the Caucasian Nature Reserve, and in Azerbaijan (Shamshev, 1998). New data show that E. adriani also inhabits the foothills of the northern side of the Caucasian Mt. Range (Severskii Distr., Krasnodar Terr.). The species occurs in forests on mountain slopes in a wide range of altitudes from 150 m (near Stavropol skaya St.) to 2100 2200 m (Kazbegi). The flight period is rather long, lasts from early May to mid-july. In male specimens of the series collected in Guzeripl, the katepisternum and meron bear a brownish spot in the ventral part, and the anterior spiracle of the thorax is occasionally brownish yellow in the dorsal part (katepisternum, meron, and spiracles of the thorax are entirely yellow in the holotype). However, the other characters of these specimens are similar to those given in the original description of E. adriani and to those in specimens collected in other places. In addition, females of this series demonstrate no color deviations from the original description. Chvála (1996) and Shamshev (1998) mentioned the black antennae of E. adriani. However, the coloration of the scape and pedicel in this species seems to be widely varying, in particular, in the specimens examined by us, these segments are brown or yellowish brown. Distribution. Russia (Adygea, Krasnodar Terr., North Ossetia), Azerbaijan, Georgia. Empis (Xanthempis) adzharica Shamshev, 1998 Material. Georgia: 1, Adzharia, Kobuleti, Kintrishskii Nature Reserve, 13.V.1971, V.G. Kovalev;

1118 SHAMSHEV, KUSTOV 2, same locality, 26.V.1971; 2, same locality, 8.V.1971. In collection of ZM MSU. Notes. The species is known from three specimens collected in Adzharia (Shamshev, 1998). We report additional material from the type locality of E. adzharica. Distribution. Georgia (Adzharia). Empis (Xanthempis) alanica Shamshev, 1998 Material. Russia: 1, Northwestern Caucasus, Guzeripl, 26.V.1970, V.G. Kovalev; 2, 70 km S of Maikop, Guzeripl, 7.VI.1970, V.G. Kovalev; 1, Krasnodar Terr., near Severskaya St., Ubinskaya St., 8.V.1970, V.G. Kovalev; 1, Krasnodar Terr., near Ubinskaya St., 17.V.1970, V.G. Kovalev. [Hereinafter, V.G. Kovalev s phrase near Severskaya St. most likely means Severskii Distr.]. In collection of ZM MSU. 2, Krasnodar Terr., Belaya River bed, Shevchenko, 16.V.2004, S.Yu. Kustov; 1, Krasnodar Terr., Apsheronskii Distr., Kamyshanova Polyana, 12 14.VI.2004, S.Yu. Kustov. In collection of KubSU. Description. Female (described for the first time) entirely corresponding to description of male (Shamshev, 1998 : 134). In addition, abdominal segment VIII along posterior margin, segment X, and entire cerci brown; cerci long, narrow, covered with short black setae. Notes. The species was known from two specimens collected in Stavropol Terr. (environs of Pyatigorsk) and Karachay-Cherkessia (Arkhyz) (Shamshev, 1998). It occurs from the foothills to the medium mountain belt of the Northwestern and Central Caucasus, at altitudes of 150 1500 m. E. alanica is endemic to the Caucasus, an Euxian species. Distribution. Russia (Krasnodar and Stavropol territories). Empis (Xanthempis) annae Shamshev et Kustov, sp. n. (Figs. 1, 2) Diagnosis. Occiput and scutum yellow, antennae black, postsutural supraalar seta absent, 7 8 dorsocentral setae present, anterior spiracle of thorax brown. Male: phallus long, slender, arcuately curved, with short beak-shaped apex. Description. Male. Face, frons, and occiput entirely yellow or brownish yellow, covered with delicate grayish pruinosity. Ocellar triangle with small brownish spot and 2 setae. Occiput with 4 strong long, and several short setae in dorsal part; more closely to neck, numerous long setae forming nearly regular transverse row; postocular setae microscopic. Antenna entirely black. Scape about 4 times as long as wide; postpedicel about 5 times as long as wide; stylus very short. Length of labrum about twice height of head. Palpus yellow, covered with pale microtrichiae and bearing several short fine brownish setae. Thorax almost entirely yellow, only notopleural line and dorsal margin of anepisternum brownish; scutum (in dorsal view) nearly shiny, with delicate, pale gray pruinosity; mesopleuron entirely covered with denser pale gray pruinosity. Prosternum glabrous. Proepisternum with 2 3 short hair-like setae in dorsal part and several similar setae in ventral part. Antepronotum concave, with 4 6 short strong setae on each side. Postpronotal tubercle with 1 long and 3 4 short setae. Mesonotum with 1 presutural supraalar, 1 long notopleural, 1 postalar, and 4 scutellar setae (apical scutellar setae subequal in length to notopleural ones, subapical setae short and fine); acrostichial setae absent; 7 8 dorsocentral setae forming one row, 5 6 anterior setae short and fine (shorter than presutural supraalar seta), 2 prescutellar setae long. Laterotergite with several black setae varying in length. Anterior and posterior spiracles brown. Legs long, slender, mostly yellow; tarsi brownish. Fore femur ventrally covered with pale microtrichiae. Fore tibia with 3 short dorsal setae in medial part. Middle and hind femora ventrally covered with dense dark microscopic setae. Middle and hind tibiae with several short antero- and posterodorsal setae. Wing hyaline, with yellowish brown, heavily sclerotized, entire (except for Sc) veins. One short costal seta present. Stigma yellowish. Veins R 4+5 and M 1 parallel at apex of wing. Radial fork wide, veins R 4 and R 5 forming right angle, vein R 4 undulate. Anal lobe forming obtuse angle. Haltere yellow. Abdomen yellowish, with vague brownish spots on anterior margin of tergites, nearly shiny (in dorsal view), with delicate, pale gray pruinosity, covered with fine black, mostly short setae; tergites I III and segment VIII with longer posteromarginal setae. Sclerites of segment VIII partly fused. Terminalia (Fig. 1) almost entirely yellow, only cercus narrowly brown along dorsal margin. Cercus with vague posteromarginal emargination; lower process

NEW AND LITTLE KNOWN SPECIES OF THE DANCE-FLY SUBGENUS 1119 Figs. 1, 2. Empis (Xanthempis) annae sp. n., male, paratype: (1) terminalia, lateral view; (2) phallus, lateral view. Scale: 0.1 mm. apod. ej, Apodeme of ejaculator; hyp. apod, hypandrial apodeme; hyp, hypandrium; fal, phallus; cer, cercus; epand, epandrium. barely visible, with 1 long seta. Lobe of epandrium trapeziform, wide, covered with numerous black setae varying in length. Hypandrium glabrous. Phallus (Fig. 2) long, slender, smoothly arcuately curved, with short beak-shaped apex. Body length 4.9 5.2 mm, length of wing 5.7 5.9 mm. Female unknown. Material. Holotype:, Russia, Krasnodar Terr., Mt. Bol shoi Tkhach, 2300 2350 m, 5.VIII.2006, S.Yu. Kustov. In ZIN collection. Paratype 1, as holotype. In ZIN collection. Notes. Empis annae sp. n. is most closely related to E. caucasica and E. pavli, but differs in the coloration of the antennae and the shape of sclerites of the male genitalia, as shown in the key above. The new species has been collected on subalpine meadows of the northern slope of Mt. Bol shoi Tkhach, seems to be a local endemic to the Northwestern Caucasus, an Euxian species. Etymology. The species name originates from the feminine name Anna. Distribution. Russia (Krasnodar Terr.). Empis (Xanthempis) grichanovi Shamshev et Kustov, sp. n. (Figs. 3, 4) Diagnosis. Occiput and scutum entirely yellow; postsutural supraalar seta absent; dorsocentral setae short, fine; anterior spiracle of thorax yellow. Male: phallus sharply ansately curved in apical part, with short beak-shaped apex. Description. Male. Face, frons, and occiput entirely yellow, covered with delicate grayish pruinosity. Ocellar triangle brownish, with 2 short fine setae. Occiput with 2 pairs of strong black setae in dorsal part; more closely to neck, numerous similar setae forming nearly regular transverse row; postocular setae microscopic. Scape and pedicel yellowish brown, postpedicel and stylus blackish. Scape about 4 times as long as wide; postpedicel about 6 times as long as wide; stylus very short. Length of labrum about twice height of head. Palpus yellow, covered with pale microtrichiae and several short fine brownish setae. Thorax almost entirely yellow, only notopleural line blackish; scutum (in dorsal view) shiny; mesopleuron entirely covered with delicate, pale gray pruinosity. Prosternum glabrous. Proepisternum with 2 3 short hair-like setae in dorsal part and several similar setae in ventral part. Antepronotum concave, with 4 5 short strong setae on each side. Postpronotal tubercle with 1 long and 3 4 short setae. Mesonotum with 1 presutural supraalar, 1 very long notopleural, 1 postalar, and 4 scutellar setae (apical scutellar setae subequal in length to notopleural setae, subapical setae short and fine); acrostichial setae absent; 7 8 dorsocentral setae forming single row, 5 6 anterior setae short and fine (shorter than presutural supraalar seta), 2 prescutellar setae long. Laterotergite with several black setae varying in length. Anterior and posterior spiracles yellowish. Legs long, slender, mostly yellow; tarsi brownish yellow. Fore femur ventrally covered with pale microtrichiae. Fore tibia with 2 short dorsal setae in apical half (position of setae widely varying). Middle and

1120 SHAMSHEV, KUSTOV Figs. 3, 4. Empis (Xanthempis) grichanovi sp. n., male, paratype: (3) terminalia, lateral view; (4) phallus, lateral view. Scale: 0.1 mm. hind femora ventrally covered with dense dark microscopic setae. Middle tibia with 1 2 obsolete dorsal setae. Hind tibia with 2 3 short anterodorsal, and 2 3 similar posterodorsal setae (number and position of setae widely varying). Wing hyaline, with yellowish brown, heavily sclerotized, entire (except for Sc) veins. One short costal seta present. Stigma yellowish. Veins R 4+5 and M 1 parallel at apex of wing. Radial fork wide, veins R 4 and R 5 forming right angle, vein R 4 undulate. Anal lobe forming obtuse angle. Haltere yellow. Abdomen yellowish or brownish yellow, with vague brownish spots on anterior margin of tergites, nearly shiny (in dorsal view), with delicate, pale gray pruinosity, covered with fine black, mostly short setae; tergites I III and segment VIII with longer posteromarginal setae. Sclerites of segment VIII partly fused. Terminalia (Fig. 3) almost entirely yellow, only cercus narrowly brown or blackish along dorsal margin. Cercus with deep posteromarginal emargination dividing it into 2 processes; lower process narrow, digitate, with 1 long seta at base. Lobe of epandrium nearly triangular, covered with numerous black setae varying in length. Hypandrium with 2 short fine apical, and 3 5 long lateral setae on each side. Phallus (Fig. 4) relatively slender, long, sharply ansately curved in apical part, with short beak-shaped apex. Body length 4.9 5.4 mm, length of wing 6.4 6.6 mm. Female. Setae of abdomen shorter than those in male; abdominal segment VII brownish yellow; tergite VIII along posterior margin, and segment X entirely brownish. Cercus brown, long, narrow, covered with short black setae. Other characters as those in male. Material. Holotype:, Russia: Krasnodar Terr., near Severskaya St., Ubinskaya, 23.V.1970, V.G. Kovalev. In collection of ZM MSU. Paratypes: 2, 1, Russia: Krasnodar Terr., near Severskaya St., Ubinskaya, 7.V.1970, V.G. Kovalev; 1, same locality, 7.V.1970; 1, same locality, 21.V.1970; 1, same locality, 5.V.1971; 1, same locality, 8.VI.1971; 1, same locality, 9.VI.1971; 1, same locality, 10.VI.1971; 1, same locality, 12.VI.1971; 1, 1. same locality, 16.VI.1971; 1, same locality, 18.VI.1971; 1, 1, same locality, 19.VI.1971; 1, Western Caucasus, Krasnaya Polyana, 18.VI.1970, V.G. Kovalev. In collection of ZM MSU. 1, Krasnodar Terr., Goryachii Klyuch, 12.VI.2000, I.Ya. Grichanov. In ZIN collection. Georgia: 1, western Georgia, Borzhom. zap. [in Cyrillic, = Borzhomskii Nature Reserve], 17.VII.1969, V.G. Kovalev; 1, 2, same locality, 21.VII.1969; 1, same locality, 6.VIII.1969; 1, same locality, 10.VIII.1969; 1, 3, eastern Georgia, Lagotekh. zap. [in Cyrillic, = Lagodekhskii Nature Reserve], 25.VII.1969, V.G. Kovalev. In collection of ZM MSU. Notes. Specimens of this series, collected in different areas of Krasnodar Terr. and Georgia, almost do not differ from the original description of E. assalemensis known from the northern slopes of Elbrus Iran (Daugeron, 2000 : 373, fig. 1). However, the differences in the shape of the phallus, found between E. assalemensis and specimens of this series, are so significant and obvious that we distinguish these

NEW AND LITTLE KNOWN SPECIES OF THE DANCE-FLY SUBGENUS 1121 specimens into a separate species. In E. assalemensis, the phallus is arcuately, smoothly curved, and that in the new species forms a sharp, ansate curvature in the apical part (Fig. 4). In addition, the lateral process of the cerci is short and wide in E. assalemensis and narrow and digitate in the new species. Already there have been known pairs of the species in the subgenus Xanthempis, which reliably differ only in the shape of sclerites of the male genitalia (especially, sclerites of the phallus). The best known example of such a pair is E. stercorea and E. aemula Loew. The new species is also closely related to E. ponti Chvála, but differs in the yellow spiracles of the thorax, more numerous and finer dorsocentral setae, a different shape of the phallus, and some other characters. The main differences of the new species from E. caucasica Bezzi are listed in the key. Among the European species of the subgenus Xanthempis, the new species is similar to E. algecirasensis Strobl, E. concolor Verrall, E. lagoensis Chvála, and E. nevadensis Chvála, which also possess an entirely yellow scutum and no postsutural supraalar seta. However, in E. algecirasensis, E. nevadensis (both are known only from Spain), and E. concolor (widely distributed in Europe), the occiput bears a dark spot behind the ocellar tubercle. In E. lagoensis (known only from the Italian Alps), the occiput is entirely yellow, and the scape and pedicel are black. In addition, all the species mentioned differ from the new species in the shape of the phallus and other structures of the male genitalia. According to the available data, E. grichanovi sp. n. is distributed from the lower to the upper mountain belts. The flight period originates in May June in the lower mountain belt and in June July in the middle and upper mountain belts. E. grichanovi is endemic to the Caucasus, an Euxian mountain species. Etymology. The species is named after the dipterologist I.Ya. Grichanov (St. Petersburg). Distribution. Russia (Krasnodar Terr.), Georgia. Empis (Xanthempis) kovalevi Shamshev, 1998 Material. Georgia: 1, 1, western Georgia, Borzhomskii Nature Reserve, 9 10.VIII.1969, V.G. Kovalev; Russia: 3, 1, western Caucasus, Krasnaya Polyana, 15.VI.1971 V.G. Kovalev. In collection of ZM MSU. Female (described for the first time) entirely corresponding to description of male (Shamshev, 1998 : 138). In addition, abdominal sternite VIII brownish yellow, tergite VIII and segment X yellow; cerci long, narrow, brownish yellow, covered with short black setae. Notes. The species was known from a single specimen collected in western Georgia (Borzhomskii Nature Reserve) (Shamshev, 1998) We report additional material from the type locality of E kovalevi; in addition, this is the first record of the species from Krasnodar Terr. (Krasnaya Polyana). Distribution. Russia (Krasnodar Terr.), Georgia. Empis (Xanthempis) pavli Shamshev, 1998 Material. Russia: 1, Krasnodar Terr., Caucasian State Nature-Biosphere Reserve, Mt. Aishkha, 1700 2200 m, 7 8.VIII.2004, S.Yu. Kustov. In collection of KubSU. Georgia: 2, 1, western Georgia, Borzhomskii Nature Reserve 7 21.VII.1969, V.G. Kovalev; 1, same locality, 13.VIII.1969; 1, 1, western Georgia, Bakuriani, 18.VIII.1969, V.G. Kovalev. In collection of ZM MSU. Notes. The species was known from three specimens collected in the environs of Sukhumi and Bakuriani (Shamshev, 1998). It has been found in Krasnodar Terr. for the first time. E. pavli is endemic to the Caucasus, an Euxian species. Distribution. Russia (Krasnodar Terr.), Georgia. Empis (Xanthempis) ponti Chvála, 1996 Material. Georgia: 6, 7, eastern Georgia, Lagodekhskii Nature Reserve, alpine-subalpine, 26 28.III.1969, V.G. Kovalev. In collection of ZM MSU. Notes. Chvála (1996) described this species from Georgia (environs of Kazbegi). Later, Shamshev (1998) found E. ponti in collections from North Ossetia. We report additional material collected in the Lagodekhskii Nature Reserve (Eastern Georgia). The species occurs in July at altitudes of 1700 2000 m. Distribution. Russia (North Ossetia), Georgia. Empis (Xanthempis) pseudoconcolor Shamshev et Kustov, sp. n. (Figs. 5, 6) Diagnosis. Occiput with large diamond-shaped spot; scutum entirely yellow, postsutural supraalar seta

1122 SHAMSHEV, KUSTOV Figs. 5, 6. Empis (Xanthempis) pseudoconcolor sp. n., male, paratype: (5) terminalia, lateral view; (6) phallus, lateral view. Scale: 0.1 mm. absent, 8 10 dorsocentral setae present; anterior spiracle of thorax yellow. Male: phallus long, smoothly curved, with long beak-shaped apex. Description. Male. Face, frons, and most of occiput yellow, covered with dense pale pruinosity; occiput with large brownish diamond-shaped spot in dorsal part. Ocellar triangle brownish, with 2 fine short setae. Occiput with 2 4 rather long, and several short setae in dorsal part; more closely to neck, numerous long setae forming nearly regular transverse row; postocular setae microscopic. Scape and pedicel yellowish brown, postpedicel and stylus blackish. Scape about 4 times as long as wide; postpedicel 5.5 6.0 times as long as wide; stylus very short. Length of labrum about twice height of head. Palpus yellow, covered with pale microtrichiae and bearing several short fine brownish setae. Thorax entirely yellow (in some paratypes, notopleural line darkened posteriorly); scutum (in dorsal view) matt, covered with delicate pale pruinosity (better visible in front view); mesopleuron entirely covered with denser pale gray pruinosity. Prosternum glabrous. Proepisternum with 2 3 short hairlike setae in dorsal part and several similar setae in ventral part. Antepronotum concave, with 4 6 short strong setae on each side. Postpronotal tubercle with 1 long, and 2 3 short setae. Mesonotum with 1 presutural supraalar, 1 long notopleural, 1 postalar, and 4 scutellar setae (apical scutellar setae subequal in length to notopleural setae, subapical setae short and fine); acrostichial setae absent; 8 10 dorsocentral setae forming single, 6 8 anterior setae short and fine (shorter than presutural supraalar seta), 2 prescutellar setae long. Laterotergite with several black setae varying in length. Anterior and posterior spiracles yellow. Legs long, slender, mostly yellow; tarsi brownish yellow. Fore femur ventrally covered with pale microtrichiae. Fore tibia with 2 short dorsal setae in apical half (position of setae widely varying). Middle and hind femora ventrally covered with dense dark microscopic setae. Middle tibia without setae (except for preapical setae). Hind tibia with 2 3 short anterodorsal, and 2 3 similar posterodorsal setae (number and position of setae widely varying). Wing hyaline, with yellowish brown, heavily sclerotized, entire (except for Sc) veins. One short costal seta present. Stigma yellowish. Veins R 4+5 and M 1 parallel at apex of wing. Radial fork wide, veins R 4 and R 5 forming right angle, vein R 4 undulate. Anal lobe forming obtuse angle. Haltere yellow. Abdomen mostly yellowish, nearly shiny (in dorsal view), with delicate, pale gray pruinosity; tergites I VII with wide brownish dorsal stripe and vague lateral spots; abdomen covered with fine black, mostly short setae; tergites I III and segment VIII with longer posteromarginal setae. Sclerites of segment VIII fused. Terminalia (Fig. 5) almost entirely yellow, only cercus narrowly brownish along dorsal margin. Cercus with deep posteromarginal emargination dividing it into 2 processes; dorsal process long, digitate, covered with short setae; lower process wide, short, with 1 long, and several short marginal setae. Lobe of epandrium nearly triangular, relatively wide, covered with numerous black setae varying in length. Hypandrium with 2 short spiniform apical, and 3 4 long lateral setae on each side. Phallus (Fig. 6) relatively slender, long, smoothly curved, with long beak-shaped apex. Body length 5.4 5.7 mm, length of wing 6.8 7 mm. Female. Tergite VIII along posterior margin and segment X entirely brownish. Cercus long, narrow,

NEW AND LITTLE KNOWN SPECIES OF THE DANCE-FLY SUBGENUS 1123 Figs. 7, 8. Empis (Xanthempis) teberdaensis sp. n., male, paratype: (7) terminalia, lateral view; (8) phallus, lateral view. Scale: 0.1 mm. brown, covered with short black setae. Other characters as those in male. Material. Holotype:, Georgia: Sukhumi, Kelasuri, oak forest, 18.V.1969, K.B. Gorodkov; Empis concolor Ver., Shamshev det. In ZIN collection. Paratypes: Georgia: 2, 2, as holotype. In ZIN collection. 1, western Georgia, Borzhomskii Nature Reserve, 13.VII.1969, V.G. Kovalev; 1, same locality, 19.VII.1969; 1, same locality, 6.VII.1969; 2, Abkhazia, environs of Sukhumi, Kashtak Gorge, 29.V.1979, V.G. Kovalev. In collection of ZM MSU. Russia: 1, Tatarka, 10 km S of Stavropol, 16.VI.1968, K.B. Gorodkov; 1, Northern Caucasus, Karachaevsk, 900 m, beech forest in a ravine, 13.IX.1965, K.B. Gorodkov; 1, Gagry (Black Sea coast, 16 30.V.1916, I. Sokolov; 2, Mt. Strizhament, environs of Stavropol, 1.VII.1973, Trjapitzin. In ZIN collection. 1, western Caucasus, Krasnaya Polyana, 19.VI.1970, V.G. Kovalev. In collection of ZM MSU. Notes. Shamshev (1998) identified as E. concolor several specimens collected in Stavropol and Krasnodar territories and Georgia. However, comparison of the species with the lectotype of E. concolor [from G.H. Verrall s collection deposited in the museum of the University of Oxford) has shown that these specimens belong to a separate species (described in the present communication). Empis pseudoconcolor sp. n. is very closely related to E. concolor, but differs in a smaller size of the dark spot on the occiput, greater number of the dorsocentral setae, and shape of the cerci and phallus. At present, E. concolor is known only from Europe (including the British isles). The new species is also closely related to E. caucasica, E. ponti, and E. grichanovi sp. n., but all these species, in contrast to E. pseudoconcolor, possess an entirely yellow occiput. The same character distinguishes the new species from E. lagoensis. In E. algecirasensis, E. nevadensis, and E. kozlovi Shamshev (known from Transbaikalia), the dark spot on the occiput differs in size and shape. Etymology. The species name points to similarity of the new species to E. concolor. Distribution. Russia (Krasnodar and Stavropol territories), Georgia (Abkhazia). Empis (Xanthempis) teberdaensis Shamshev et Kustov, sp. n. (Figs. 7, 8) Diagnosis. Occiput with large dark spot; scutum entirely yellow, postsutural supraalar seta absent, 7 8 short fine (except for prescutellar ones) dorsocentral setae present, anterior spiracle of thorax brownish. Male: phallus long, slender, sinuously curved in apical part, with long narrow beak-shaped apex. Description. Male. Face and frons brownish yellow, covered with grayish pruinosity. Occiput mostly yellow, covered with grayish pruinosity, its dorsal part with large dark spot also covering ocellar triangle but not reaching margin of eyes. Ocellar triangle with 2 fine, rather long setae. Occiput with 4 long, and several short setae in dorsal part; more closely to neck, numerous long setae forming nearly regular transverse row; postocular setae short. Scape and pedicel brownish, postpedicel and stylus black. Scape about 4 times as long as wide; postpedicel about 6 times as long as wide; stylus very short. Length of labrum about twice height of head. Palpus yellow, covered with pale microtrichiae and bearing several short fine brownish setae.

1124 SHAMSHEV, KUSTOV Thorax almost entirely yellow, only notopleural line brownish; scutum (in dorsal view) nearly shiny, covered with delicate, pale gray pruinosity; mesopleuron entirely covered with denser pale gray pruinosity. Prosternum glabrous. Proepisternum with 2 3 short hair-like setae in dorsal part and several similar setae in ventral part. Antepronotum concave, with 6 7 short strong setae on each side. Postpronotal tubercle with 1 long and 3 4 short setae. Mesonotum with 1 presutural supraalar, 1 notopleural, 1 postalar, and 4 scutellar setae (apical setae long, subapical ones short); acrostichial setae absent; 7 8 dorsocentral setae forming single row, 5 6 anterior setae short and fine (shorter than presutural supraalar seta), 2 prescutellar setae long. Laterotergite with several black setae varying in length. Anterior and posterior spiracles brownish. Legs long, slender, mostly yellow; tarsi brownish yellow. Fore femur ventrally covered with pale microtrichiae. Fore tibia with 2 3 (2, in holotype) short dorsal setae in apical half. Middle and hind femora ventrally covered with dense dark microscopic setae. Middle and hind tibiae with 2 3 short anterodorsal, and 2 3 posterodorsal setae (position of setae widely varying). Wing hyaline, with yellowish brown, heavily sclerotized, entire (except for Sc) veins. One short costal seta present. Stigma yellowish. Veins R 4+5 and M 1 parallel at apex of wing. Radial fork wide, veins R 4 and R 5 forming right angle, vein R 4 undulate. Anal lobe forming obtuse angle. Haltere yellow. Abdomen yellowish, nearly shiny (in dorsal view), with delicate, pale gray pruinosity, covered with fine black, mostly short setae; tergites I III and segment VIII with longer posteromarginal setae. Sclerites of segment VIII fused. Terminalia (Fig. 7) entirely yellow. Cercus with deep posteromarginal emargination dividing it into 2 processes; lower process short, wide, with 1 long and several short marginal setae. Lobe of epandrium nearly triangular, narrow in apical half, covered with numerous black setae varying in length. Hypandrium with 2 short and fine apical, and 3 rather long lateral setae on each side. Phallus (Fig. 8) long, relatively fine, sinuously curved in apical part, with long narrow beakshaped apex. Length of wing 6.9 7.3 mm. Female unknown. Material. Holotype:, Russia: Teberda, 7.VII.1963, A. Tikhomirova. In collection of ZM MSU. Paratype: 1, Northern Caucasus, M. Khatipara, 2000 m, Teberdinskii Nature Reserve, 29.VI.1982. E.P. Nartshuk. In ZIN collection. Additional material. 1 specimen (sex?), as holotype. Notes. The new species belongs to a complex of species with an entirely yellow scutum and without postsutural supraalar seta. This complex includes the following Caucasian species of the subgenus Xanthempis: E. caucasica, E. grichanovi sp. n., Е. pseudoconcolor sp. n., E. ponti, E. pavli, and E. zamotajlovi sp. n. The new species can be readily distinguished from E. caucasica, E. grichanovi sp. n., and Е. pseudoconcolor sp. n. in the brownish anterior spiracles of the thorax. Some specimens of E. caucasica possess darkened spiracles, but the occiput in this species is always entirely yellow. The spiracles of the thorax in Empis ponti, E. pavli, and E. zamotajlovi sp. n. are brownish, as those in the new species. However, E. pavli and E. zamotajlovi sp. n. differ from the new species in the entirely yellow occiput, and Е. ponti differs in the sparser and longer dorsocentral setae. Among the European species of the subgenus Xanthempis, the new species can be compared with E. algecirasensis, E. concolor, E. lagoensis, and E. nevadensis. However, in all the species listed, the spiracles of the thorax are yellow. The same character distinguishes the new species from E. kozlovi. The collection of ZM MSU includes one specimen of this species, labeled as the holotype. However, we do not list the specimen among paratypes because of the almost total absence of its abdomen. The species seems to be characteristic of the medium- and high-mountain forest zones of the Central Caucasus. This is a local Euxian endemic to the Caucasus. Etymology. The species name originates from the name of the type locality (Teberda). Distribution. Russia (Karachay-Cherkessia). Empis (Xanthempis) zamotajlovi Shamshev et Kustov, sp. n. (Figs. 9, 10) Diagnosis. Occiput without dark spot; scutum with 2 pairs of brownish spots; postsutural supraalar seta

NEW AND LITTLE KNOWN SPECIES OF THE DANCE-FLY SUBGENUS 1125 present; anterior spiracle of thorax yellow. Male: phallus nearly straight, with short cup-shaped apex. Description. Male. Face, frons, and occiput entirely yellow or reddish yellow, covered with delicate grayish pruinosity. Ocellar triangle brownish or brownish yellow, with 2 relatively strong setae. Occiput with several strong setae in dorsal part, setae varying in length; more closely to neck, numerous setae forming nearly regular transverse row; postocular setae microscopic. Scape and pedicel yellowish brown, postpedicel and stylus blackish (postpedicel occasionally paler at base). Scape about 4 times as long as wide; postpedicel about 5 times as long as wide; stylus very short. Length of labrum about twice height of head. Palpus yellow, covered with pale microtrichiae and with several short fine brownish setae varying in length. Thorax almost entirely yellow; scutum (in dorsal view) with 2 pairs of relatively large, oval, brownish spots (posterior pair of spots usually vague, occasionally barely visible), nearly shiny, covered with delicate, pale gray pruinosity; notopleural line brownish yellow or blackish; mesopleuron entirely covered with delicate, pale gray pruinosity. Prosternum glabrous. Proepisternum with several short, fine, black setae in ventral part. Antepronotum concave, with 6 8 short strong setae on each side. Postpronotal tubercle with 1 long and 2 3 short setae. Mesonotum with 1 presutural supraalar, 2 notopleural (anterior one longer), 1 postsutural supraalar, 1 postalar, and 4 scutellar setae (apical scutellar setae subequal in length to anterior notopleural ones, subapical setae shorter and finer); acrostichial setae absent; 8 9 dorsocentral setae forming single row, strong, relatively short (slightly shorter than presutural supraalar seta), 2 prescutellar setae longer. Laterotergite with numerous black setae varying in length. Anterior and posterior spiracles yellowish. Legs long, slender, almost entirely yellow, only 5th tarsal segment brownish. Fore femur ventrally without pale pubescence, covered with rather dense microscopic setae. Fore tibia with 2 4 short dorsal setae in apical half (number and especially position of setae widely varying). Middle and hind femora ventrally covered with dense dark microscopic setae. Middle tibia without pronounced setae (except for preapical ones). Hind tibia with 1 3 short dorsal setae (number and position of setae widely varying). First segment of hind tarsus slightly thickened in basal half. Figs. 9, 10. Empis (Xanthempis) zamotajlovi sp. n., male, paratype: (9) terminalia, lateral view; (10) phallus, lateral view. Scale: 0.1 mm. Wing hyaline, with yellowish brown, heavily sclerotized, entire (except for Sc) veins. One short costal seta present. Stigma yellowish. Veins R 4+5 and M 1 parallel at apex of wing. Radial fork wide, veins R 4 and R 5 forming right angle, vein R 4 undulate. Anal lobe forming obtuse angle. Haltere yellow. Abdomen yellowish, with vague triangular brownish spots on anterior margin of tergites, nearly shiny (in dorsal view), with delicate, pale gray pruinosity, covered with fine black, mostly short setae (posteromarginal setae longer). Sclerites of segment VIII partly fused. Terminalia (Fig. 9) almost entirely yellow, only cercus widely brownish along dorsal margin. Cercus with vague posteromarginal emargination dividing it into 2 processes; dorsal process with narrow elongate apex, covered with numerous marginal setae; lower process wide, with 1 long seta. Lobe of epandrium trapeziform, covered with numerous black setae varying in length. Hypandrium glabrous, with triangular apex. Phallus (Fig. 10) long, relatively slender, mainly almost straight, with short cup-shaped apex. Body length 5.9 6.4 mm, length of wing 6.9 7.2 mm. Female. First segment of hind tarsus fine. Setae of abdomen shorter than those in male; segment VIII along posterior margin and segment X entirely brownish. Cercus long, narrow, brown, covered with short black setae. Other characters as those in male. Material. Holotype:, Russia: Krasnodar Terr., Afips River bed, 7 km S of Krepostnaya St., 3.V.2004, S.Yu. Kustov. ZIN collection. Paratypes: 1, 1,

1126 SHAMSHEV, KUSTOV as holotype; 1, 1, Krasnodar Terr., upper Afips River, Verkhneafipskoe hunt ground, 9.V.2004, S.Yu. Kustov; 1, Krasnodar Terr., Kamyshanova Polyana, 28.V.2000, S.Yu. Kustov; 1, same locality, 15.V.2004; 2, 1, same locality, 12 14.VI.2004. In collections of ZIN and KubSU. 1, northwestern Caucasus, Guzeripl, 11.V.1970, V.G. Kovalev; 1, same locality, 1.VI.1970; 2, Krasnodar Terr., near Severskaya St., Ubinskaya, 12.VI.1970, V.G. Kovalev. In collection of ZM MSU. Notes. The new species is closely related to E. adzharica known from the Caucasus (Shamshev, 1998). Both the species possess a postsutural supraalar seta and two pairs of brownish spots on the scutum (an unique character of the subgenus Xanthempis). The main differences between these species are listed in the key. Empis zamotajlovi sp. n. occurs in broadleaf and mixed forests, from the foothills to the medium mountain belt (150 1500 m). The flight period lasts from late April early May to mid-june. Apparently, E. zamotajlovi is endemic to the northwestern Caucasus, an Euxian species. Etymology. The species is named after A.S. Zamotajlov (Krasnodar), who made a great contribution to organization of investigations of insects of the Caucasus and considerably extended the knowledge of the coleopteran fauna of this region. Distribution. Russia (Krasnodar Terr., Adygea). ACKNOWLEDGMENTS The authors are grateful to A.L. Ozerov (Moscow) for the Empididae material from the Zoological Museum of the Moscow State University, supplied for examination. I.V. Shamshev thanks A. Pont (Oxford) for his help in organization of the visit to the University of Oxford. REFERENCES 1. Chvála, M., Swarming, Mating and Feeding Habits in Empididae (Diptera), and Their Significance in Evolution of the Family, Acta Entomol. Bohemoslov. 73, 353 366 (1976). 2. Chvála, M., The Empidoidea (Diptera) of Fennoscandia and Denmark: III. Genus (Empis), Fauna Entomol. Scand., No. 29, 1 192 (1994). 3. Chvála, M., Classification and Phylogeny of European Empis Subgenus Xanthempis Bezzi (Diptera, Empididae), Studia Dipterol. 3 (1), 3 18 (1996). 4. Daugeron, C., The Subgenus Xanthempis: New Species and Taxonomical Data (Diptera: Empididae), Ann. Soc. Entomol. France (N. S.) 36 (4), 371 388 (2000). 5. Engel, E.O., Xanthempis Bezzi, in Die Fliegen der Palaearktischen Region, Ed. by Lindner, E. (Stuttgart, 1943), Vol. IV (4), pp. 192 281. 6. Hamm, A.H., The Epigamic Behaviour and Courtship of Three Species of Empididae, Entomol. Mont. Mag. 69, 113 117 (1933). 7. Kustov, S.Yu. and Shamshev, I.V., Zoogeographical Characteristic of the Fauna of Flies the Families Hybotidae and Empididae (Diptera) of the Caucasus, in Abstracts of Papers of the XIII Congress of the Russian Entomol. Soc. Problems and Trends of General Entomology, Krasnodar, September 9 15, 2007 (Krasnodar, 2007), pp. 184 185. 8. McAlpine, J.F., Morphology and Terminology Adults. [Chapter] 2 in Manual of Nearctic Diptera, Ed. by McAlpine, J.F. (1981), Vol. 1, pp. 9 63 [Research branch agriculture Canada. Monograph 27; Ottawa]. 9. Preston-Mafham, K.G., Courtship and Mating in Empis (Xanthempis) trigramma Meig., E. tessellata F. and E. (Polyblepharis) opaca F. (Diptera: Empididae) and the Possible Implication of Cheating Behaviours, J. Zool. 247, 239 246 (1999). 10. Shamshev, I.V., Revision of the Genus Empis Linnaeus (Diptera: Empididae) from Russia and Neighbouring Lands: I. Subgenus Xanthempis Bezzi, Int. J. Dipterol. Res. 9 (2), 127 170 (1998). 11. Shamshev, I.V. and Kustov, S.Yu., A List of Species of the Families Hybotidae and Empididae (Diptera) of the Caucasus, in Kavkaz. Entomol. Byull. 2 (2), 221 230 (2006). 12. Sinclair, B.J., Morphology and Terminology of Diptera Male Terminalia. [Chapter] 1, 2 in Contributions to a Manual of Palaearctic Diptera (with Special Reference to Flies of Economic Importance), Vol. 1, General and Applied Dipterology, Ed. by Papp, L, Darvas, B. (Science Herald, Budapest, 2000), pp. 53 74. 13. Stuckenberg, B.R., Antennal Evolution in the Brachycera (Diptera), with a Reassessment of Terminology Relating to the Flagellum, Studia Dipterol. 61 (1), 33 48 (1999).