P O L I S H J O U R N A L OF ENTOMOLOG Y

P O L I S H J O U R N A L OF ENTOMOLOG Y P O L S K I E P I S M O E N T O M O L O G I C Z N E 3 VOL. 82: 287-302 Gdańsk 31 December 2013 DOI: 10.2478/v10200-012-0043-6 The Old World biting midge, Forcipomyia (Lepidohelea) pulcherrima SANTOS ABREU, new to the fauna of the United States (Diptera: Ceratopogonidae) WILLIAM L. GROGAN JR. 1, LAWRENCE J. HRIBAR 2 & FRANCIS G. HOWARTH 3 1 Research Associate, Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, Gainesville, Florida 32614, USA, e-mail: groganw@doacs.state.fl.us; 2 Florida Keys Mosquito Control District, 503 107 th Street, Marathon, Florida 33050, USA, e-mail: lhribar@keysmosquito.org; 3 Distinguished Associate, Department of Natural Sciences, Bishop Museum, 1525 Bernice Street, Honolulu, Hawaii 96817-2704, USA, e-mail: fhowarth@bishopmuseum.org ABSTRACT. We provide the first United States records of the Old World biting midge, Forcipomyia (Lepidohelea) pulcherrima SANTOS ABREU (Diptera: Ceratopogonidae), from California, Florida and Hawaii. The fourth instar larva of F. pulcherrima is also described and illustrated for the first time. KEY WORDS: Diptera, Ceratopogonidae, biting midge, Forcipomyia, California, Florida, Hawaii, new records, fourth instar larva, breeding site. INTRODUCTION The biting and predaceous midge (Diptera: Ceratopogonidae) fauna of the continental United States and Canada is very diverse with over 600 recorded species (BORKENT & GROGAN 2009); however, several of these are primarily Neotropical species that range north of Mexico in Arizona, California and Texas (BORKENT & SPINELLI 2000). Similarly, several predominatly Neotropical species that primarily inhabit the Caribbean Region also occur in Florida and some adjacent states (WILKENING et al. 1985, GROGAN et al. 2010).

288 Polish Journal of Entomology 82 (4) Some species in the Nearctic have apparently been introduced from elsewhere. For example, the Old World species, Forcipomyia (F.) swezeyana TOKUNAGA & MURACHI, 1959, was documented from Florida by WIRTH & SPINELLI (1992b), who reported this Australasian species from decaying Philodendron and banana (Musa) plants. More recently, GROGAN & HRIBAR (2006) reported the Neotropical species, Forcipomyia (Phytohelea) bromelicola (LUTZ), 1913, in Florida based on adults reared from larvae and pupae inhabiting bromeliads in the Florida Keys. During 2006, WLG and LJH examined specimens of an unusual species of biting midge in the subgenus Lepidohelea KIEFFER, 1917, of the genus Forcipomyia MEIGEN, 1818, which were collected during routine mosquito light trap surveys in the Florida Keys (Monroe Co.). Adult males of this species have greatly modified apices of their gonostyli that are expanded, flattened and foot-shaped, and adults of both sexes have dark brown or black femora and tibiae with a narrow to broad pale band at mid-length of these leg segments. Neither of these characters are present in any of the currently known Nearctic or Neotropical species in this subgenus, and we could not confidently key it to any New World species of F. (Lepidohelea) in the keys provided by WIRTH (1991b) and WIRTH & SPINELLI (1992a, b, 1993a, b). Therefore, we considered the possibility that it might represent an undescribed species. Subsequently, we identified several adults and their associated larvae of this same species that were intercepted by personnel of the Florida Department of Agriculture and Consumer Services from the soil of potted orchids that originated from a nursery in San Joaquin Co., California in late 2011. During a conversation by WLG about these specimens with Robert Woodruff (pers. comm.), Woodruff suggested that because they were associated with the orchid industry, it was likely a widespread, previously described species. Woodruff s suggestion proved accurate, as we found illustrations of the male genitalia of this species in articles by CLASTRIER (1956) and DESSART (1963), as well as habitus illustrations of adults of both sexes and their leg patterns in DESSART S (1963) review of African Forcipomyia. Finally, during early 2013, FGH contacted WLG about specimens collected in Hawaii on the islands of Maui and Oahu that HOWARTH & PRESTON (2007) had previously identified as Forcipomyia (Lepidohelea) chrysolopha (KIEFFER), 1911, which proved to be conspecific with our material from California and Florida. For most of the 20 th century, this species was known as Ceratopogon chrysolophus KIEFFER, 1911, or Forcipomyia chrysolopha; Lepidohelea ornatipes KIEFFER, 1921, or F. ornatipes; Lepidohelea formosae KIEFFER, 1922, or F. formosae; F. lepidota INGRAM & MACFIE, 1924; and F. variegeta GOETGHEBUER, 1933 (CLASTRIER 1956, 1983, SZADZIEWSKI 1983, WIRTH et al. 1980). Confusion with names of this species was finally resolved by BORKENT (1997) when he examined type material of SANTOS ABREU, who was actually the first to describe this species as Forcipomyia pulcherrima SANTOS ABREU,

GROGAN W.L. JR. et al.: The Old World biting midge, Forcipomyia pulcherrima 289 1918, in his review of the Chironomidae of the Canary Islands (SANTOS ABREU 1918). More recently, GHONAIM et al. (2001) described and illustrated adults of F. pulcherrima based on 84 males and 3 females from Marsafa, Qalyubiya, Egypt, and F. (Lepidohelea) marsafae, new species, based on 5 males and 1 female from the same locality in Egypt. However, SZADZIEWSKI et al. (2011) regarded F. marsafae GHONAIM et al. 2001 as a new synonym of F. pulcherrima, and, subsequently, BORKENT (2013) also listed F. marsafae as a synonym of F. pulcherrima. Here we provide the first United States records of this Old World biting midge from California, Florida and Hawaii, as well as the first description with accompanying scanning electron micrographs and automontage photographs of the fourth instar larva of this species and its mouthparts. We also reproduce habitus illustrations of adults by DESSART (1963) and male genitalia by CLASTRIER (1956), and include photomicrographs of the male genitalia and female spermathecae. We are pleased to dedicate this article to our colleague, Ryszard Szadziewski, in recognition of his numerous contributions to the systematics and biology of Palaearctic, Afrotropical and Asian Ceratopogonidae. In addition, he has also published many excellent articles on fossil biting midges from several amber deposits in the New and Old World. Acknowledgements We thank David J. DeMay, Florida Keys Mosquito Control District, for the Florida specimens, and Dyranna Russell-Hughes and colleagues, Florida Department of Agriculture and Consumer Services, Division of Plant Industry, for the California specimens. FGH extends special thanks to Bernarr Kumashiro, insect taxonomist and curator, Hawaii Department of Agriculture, for use of that department's insect collection. We extend special thanks to Gary Steck for the SEM photomicrographs and automontage colour photographs of larvae, and to Julieta Brambila for preparing the digital automontage photographs of the larval mouthparts. We are especially grateful to Art Borkent and Gustavo Spinelli for their reviews of an earlier draft of the manuscript. We also thank Patrick Grootaert, Institut Royal des Sciences Naturelles de Belgique, for permission to reproduce Dessart s habitus illustrations of adults. MATERIALS AND METHODS Adults were collected with light traps in the Florida Keys, whereas larvae and adults were collected by personnel employed by the Florida Department of Agriculture and Consumer Services from the soil of potted orchids (Phalaenopsis sp.) in a truck from Lodi, California en route to Miami, Florida. Adults from Hawaii were collected by Malaise trap, MV light and sweeping over a stream and a mating swarm. Specimens were preserved

290 Polish Journal of Entomology 82 (4) in 70-75% ethanol, and unless otherwise indicated, subsequently cleared in a solution of phenol crystals dissolved in 100% ethanol, and mounted on microscope slides in a mixture of the ethanol-phenol solution and Canada balsam by the methods described by WIRTH & MARSTON (1968). Specimens were compared with slide-mounted specimens of all Nearctic and some Neotropical species in F. (Lepidohelea), as well as published descriptions and illustrations of all Neotropical species. Terminology of adult Ceratopogonidae are those in DOWNES & WIRTH (1981), whereas the terminology of larvae follow those in SAUNDERS (1925, 1959), HRIBAR (1993), HRIBAR & MULLEN (1991) with updated changes of some mouthpart terms suggested by FÜRST v. LIEVEN (1998). Voucher specimens are deposited in the Florida State Collection of Arthropods, Gainesville (FSCA); United States National Museum of Natural History, Washington, D. C. (USNM); Canadian National Collection of Insects, Ottawa (CNCI); Museo de La Plata, Argentina (MLPA); Bishop Museum, Honolulu, Hawaii (BPBM); and the Florida Keys Mosquito Control District, Marathon (FLKC). Global Information Systems (GIS) data of specimens from California and Hawaii are all WGS-84 map datum. SYSTEMATICS DEBENHAM (1987) reviewed the Australasian species in Forcipomyia sensu stricto and F. (Lepidohelea). She concluded that previous studies of some subgenera of Forcipomyia by DE MEILLON et al. (1982) as well as the key to subgenera by WIRTH & RATANAWORABHAN (1978) were usually accurate for identifying adult males, but often totally inadequate for distinguishing adult females in these two subgenera. DEBENHAM S diagnosis of the subgenus Lepidohelea included several characters which, in combination, usually provided accurate assignment of adult males and most adult females to this subgenus. We list below what we consider the most important of these characters, which in combination, usually distinguish adult males and females of species in F. (Lepidohelea) from species in other subgenera of Forcipomyia: 1. Body usually with flattened, striated scales that are densest on the wings and legs, and very broad on legs; 2. Female mandible vestigial, without functional teeth; 3. Femora and tibiae with complex patterns of light and dark bands or spots; 4. Wing distinctly patterned due to the presence of pale and dark scales concentrated on discrete areas of veins and membrane; 5. Females with 2 (rarely 1) spermathecae that are spherical or slightly ovoid with distinct necks and often surface punctations; 6. Male genitalia with divided parameres of various shapes, aedeagi with unique shapes depending on the group they belong to, and gonostyli that are straight to sinuous with greatly modified apices in some groups.

GROGAN W.L. JR. et al.: The Old World biting midge, Forcipomyia pulcherrima 291 Forcipomyia (Lepidohelea) pulcherrima SANTOS ABREU, 1918 (Figs 1-17) Forcipomyia pulcherrima SANTOS ABREU, 1918: 272. (Canary Islands); BORKENT 1997: 13 (designated male as lectotype; listed all subsequent descriptions of this species as new synonyms). Forcipomyia (Lepidohelea) pulcherrima: BORKENT & WIRTH 1997: 41 (in World Catalogue); GHONAIM et al. 2001: 43 (records from Egypt; description, figs). Lepidohelea ornatipes KIEFFER, 1921: 1. (Sudan). [Preoccupied by Forcipomyia ornatipes (KIEFFER), 1918. Cameroon]; BORKENT 1997: 13 (as new synonym of F. pulcherrima SANTOS ABREU). Forcipomyia (Lepidohelea) ornatipes: WIRTH et al. 1980: 156 (in Afrotropical catalogue; as synonym of Ceratopogon chrysolopha KIEFFER); BORKENT & WIRTH 1997: 41 (in World Catalogue). Lepidohelea formosae KIEFFER, 19 22: 153. (Taiwan); BORKENT 1997: 13 (as new synonym of F. pulcherrima SANTOS ABREU). Forcipomyia formosae: TOKUNAGA 1940: 73 (records from Japan; fig. male genitalia). Forcipomyia (Lepidohelea) formosae: REMM 1967: 4 (in review of Ceratopogonidae of the Caucasus; record from USSR; fig. male genitalia; Forcipomyia lepidota INGRAM & MACFIE as probable synonym); SZADZIEWSKI 1983: 381 (Algeria record); BORKENT 1997: 13 (as new synonym of F. pulcherrima SANTOS ABREU); BORKENT & WIRTH 1997: 41 (in World Catalogue; as synonym of F. pulcherrima SANTOS ABREU). Forcipomyia lepidota INGRAM & MACFIE, 1924: 566. (Ghana); DESSART 1961: 362 (in review of species of Forcipomyia described by GOETGHEBUER from the Congo); SZADZIEWSKI 1983: 381 (as new synonym of Lepidohelea formosae KIEFFER); BORKENT 1997: 13 (as new synonym of F. pulcherrima SANTOS ABREU). Forcipomyia (Forcipomyia) lepidota: DESSART 1963: 82 (description; figs; distribution in Africa); CLASTRIER 1966: 694 (records from the Canary Islands). Forcipomyia (Lepidohelea) lepidota: CLASTRIER 1956: 506 (redescription; figs male genitalia, female wing, spermathecae; records from Algeria and Tunisia); SZADZIEWSKI 1983: 381 (Algeria; as synonym of Lepidohelea formosae KIEFFER); WIRTH 1990: 232 (in review of biting midges from Aldabra Atoll; distribution); BORKENT & WIRTH 1997: 41 (in World Catalogue; as synonym of F. pulcherrima SANTOS ABREU). Forcipomyia variegata GOETGHEBUER, 1933: 133. (Congo); DESSART 1961: 362 (as synonym of F. lepidota INGRAM and MACFIE); BORKENT 1997: 13 (as new synonym of F. pulcherrima SANTOS ABREU). Forcipomyia (Lepidohelea) variegata: BORKENT & WIRTH 1997: 41 (in World Catalogue; as synonym of F. pulcherrima SANTOS ABREU).

292 Polish Journal of Entomology 82 (4) Forcipomyia (Lepidohelea) marsafae GHONAIM, IBRAHIM & ALI, 2001: 42. (Egypt); SZADZIEWSKI et al., 2011: 638 (as new synonym of F. pulcherrima SANTOS ABREU; in review of Ceratopogonidae of the United Arab Emirates; records from UAE); BORKENT 2013: 47 (in online World Catalogue; as synonym of F. pulcherrima SANTOS ABREU). Ceratopogon chrysolophus KIEFFER, 1911: 333 (Seychelles). Forcipomyia (Lepidohelea) chrysolopha: WIRTH & MESSERSMITH 1977: 297 (redescription; figs; records from Seychelles and Mauritius; Lepidohelea ornatipes KIEFFER, Forcipomyia variegata GOETGHEBUER, Forcipomyia grata GOETGHEBUER, Forcipomyia guttatella GOETGHEBUER, as new synonyms of F. chrysolopha); WINDER 1978: 11 (in list of World Cacao pollinators; Forcipomyia lepidota INGRAM & MACFIE as synonym); WIRTH et al. 1980: 156 (in Afrotropical catalogue; L. ornatipes KIEFFER, F. lepidota INGRAM & MACFIE, and, F. variegata GOETGHEBUER as synonyms of F. chrysolopha); CLASTRIER 1983: 52 (Seychelles; designated male syntype as lectotype redescription; figs.); BORKENT & WIRTH 1997: 41 (in World Catalogue; Forcipomyia grata GOETGHEBUER, and, Forcipomyia guttatella GOETGHEBUER, as synonyms of F. chrysolopha). Diagnosis of adult males and females Adults (Fig. 1) differ from all New World species of F. (Lepidohelea) in having dark brown to black femora and tibiae with narrow to moderately broad pale bands at mid-length of these leg segments. Adult males differ from all New World species of F. (Lepidohelea) in having a gonostylus with a highly modified apex that is greatly expanded, flattened and boot- or foot-shaped (Figs 2-3). Adult females possess two spherical to slightly ovoid spermathecae (Fig. 4) with moderately long, tapering conical necks, which readily distinguishes them from females of all New World species in the annulatipes group of F. (Lepidohelea), which only have a single spermatheca (WIRTH & SPINELLI 1993a). Adult females of New World species in the seminole (WIRTH 1976, WIRTH & SPINELLI 1992a) and bicolor (WIRTH 1991a, b, WIRTH & SPINELLI 1993b) groups differ from adult females of F. pulcherrima in having ovoid spermathecae with narrow, short to very short necks. Description of fourth instar larva Total length 3.49 (3.17-3.76, n=7) mm. Colour translucent milky white; head capsule brown to greyish-brown. Head (Figs 5, 9). Hypognathous, moderately sclerotized, connected at angle of 20-30 with prothoracic segment; eyes circular to slightly ovoid, located posterior to antenna; antenna stout, tapering slightly distally to sharply pointed tip; p seta situated dorsad of eye, greatly elongated, broad and flattened basally, distal portion filiform, round in X-section,

GROGAN W.L. JR. et al.: The Old World biting midge, Forcipomyia pulcherrima 293 tapering gradually distally to very slender apex, surface apparently without spicules; q seta slightly below level of antenna, well developed, very long, virtually identical to p seta in shape and length; u seta situated considerably below eye, very long, tapering gradually distally; t seta short, 0.15 length of q seta. Several external mouthparts visible (Fig. 10), including the epipharynx (ep) and maxillary palpus (mp). Internal mouthparts (Figs 13-17) dark brown to blackish, very heavily sclerotized. Mandible (Figs 13-14) oriented to function in sagittal plane; triangular-shaped in three dimensional space at base, becoming laterally compressed, somewhat scoop-like toward distal portion with three well developed teeth in lateral view; anterior tooth broadly rounded, median tooth bluntly truncate, posterior tooth small, sharply pointed; large lateral protuberance visible from base to about midlength of mandible; length 0.122-0.125 mm (n=2), greatest breadth (at base) 0.038-0.043 mm (n=2). Messor (Figs 13, 15) blunt, sinuate, somewhat L-shaped, moving in sagittal plane. Mola cibarialis (JOBLING 1953; = epipharynx of other authors) (Fig. 16) large, heavily sclerotized, with several rows of slender, large, coarse teeth (Fig. 17). Four pairs of sensilla on epipharynx (= labrum of authors; palatum of HRIBAR & MULLEN 1991) visible under oil immersion (1000X), arranged anterior to posterior along midline. Hypostoma smooth, without teeth. Thorax. Prothoracic segment (Figs 5-6, 9) with pseudopod divided for about ½ of length, covered with fine spicules, annulations; a seta very short, non-hastate, basal ½ broad, distal portion tapers to fine point; b seta 3X longer than a seta, similar in shape to p seta of head; c seta (Fig. 6) slender, shorter than a seta. Mesothoracic, metathoracic segments (Figs 5-6) with short, slender, hastate a setae. Abdomen. All segments with slender, hastate a setae (Figs 7-8 ); terminal segment with non-hastate a setae identical to b setae in shape but slightly shorter, both setal types in slightly U-shaped pattern on dorsal surface (Fig. 7); anal pseudopod (Figs 7-8) elongate with apical whorl of bifid hooks. Thoracic, abdominal segments covered dorsally, laterally with fine, curved spicules (Figs 11-12). Material examined (all are New USA Records) United States: CALIFORNIA, San Joaquin Co., Lodi, 38.099 N, 121.317 W, 15-XII- 2011, ex. Phalaenopsis, 4 males, 1 female, 16 larvae. FLORIDA, Monroe Co., Key Largo, 17 Aug. 2006, D. DeMay, light trap, 3 females; same data except 26 Aug. 2006, 1 male; Tavernier, Taveraero Airport, 30 June 2009, D. DeMay, light trap, 1 male, 1 female; Upper Matecumbe Key, 4 Nov. 2008, D. DeMay, CO 2 -baited light trap, 1 female; Windley Key, 17 Nov. 2008, D. DeMay, CO 2 -baited light trap, 2 males. HAWAII, Maui I., Kahului Airport environs, 6 m elev., 20.903 N, 156.429 W, 14 Jan.-1 Feb. 2000, F.G. Howarth, D.J. Preston & J.E. Dockall, Malaise Trap, Leucaena shrubland, 4 females in ethanol [BL2002-184]; same data except 5 m elev., 20.904 N, 156.428 W, 21 Oct.-3. Nov. 2006, F. & K. Starr, F.G. Howarth, Malaise trap, Prosopis woodland, 3 females, 1 male and

294 Polish Journal of Entomology 82 (4) 12 females, 1 male in ethanol (BPBM), 2 females (FSCA) [KA2007-171]; same data except 2 m elev., 20.904 N, 156.431 W, 14 Nov. 2006, F.G. Howarth, D.J. Preston, F. & K. Starr, MV Bulb, Prosopis-Casuarina woodland, 2 males in fluid (BPBM) [KA2007-169]; same data except swarming at dusk near MV Bulb set up, >50 males in ethanol (BPBM) [KA2007-169A]. Oahu I., Honolulu, Kalihi Valley, 120 m elev, 21.344 N, 157.860 W, 7 Dec. 2007, FG Howarth, at light, 1 male (BPBM); same data except 25 Jan. 2009, 1 male, 3 females; same data except 28 Jan. 2009, 1 male; same data except 31 Jan. 2009, 3 males, 1 female (BPBM); Moanalua Valley, 21.3667 N; 157.8783 W, 80 m elev., 25 Jan. 2009, F.G. Howarth, sweeping stream, 2 females (BPBM). Hawaii I.., Koloko National Park, Aimakapa Pond, 19.67675 N, 156.02417 W, 15-19 Jun. 2012, Malaise trap, R. Englund, 3 females in ethanol (BPBM). Distribution Forcipomyia pulcherrima is broadly distributed in the Old World in the Canary Islands (type locality), southern Palaearctic in Algeria, Tunisia, Egypt, Spain and Caucasus, Afrotropical Region, the Seychelles, Aldabra, Reunion, and in Asia in the United Arab Emirates, China, Japan, Malaysia and Taiwan (WIRTH et al. 1980, GHONAIM et al. 2001, YU et al. 2005, SZADZIEWSKI et al. 2011). We are confident that it was recently introduced into and is now established in at least three states in the USA for three reasons: 1. It was not included in the New World studies by WIRTH (1991b) and WIRTH & SPINELLI (1992a, b, 1993a, b), 2. It is surely a recently introduced species in Hawaii, where four other nonnative species of ceratopogonids have also recently become established (HOWARTH & PRESTON 2007); and 3. Specimens were intercepted in a shipment of orchids en route from California to Miami, Florida. This latter fact suggests that it was introduced from Asia to Hawaii prior to 2000, then from that state into California, and subsequently from California into Florida also prior to 2005. DISCUSSION GHONAIM et al. (2001) indicated that their new species from Egypt, Forcipomyia (Lepidohelea) marsafae, differed from specimens of F. pulcherrima in its darker colour, having more scattered (wing) macrotrichia, smaller oval female spermathecae, having rather long and strong bristles on femora, tibiae, and 1 st and 2 nd tarsomeres of all legs, and stouter gonocoxite without folds. The more divergent distal end of the gonostylus is not expanded, and basal part of parameres is more inflated. In their review of the ceratopogonids of the United Arab Emirates, SZADZIEWSKI et al. (2011) noted that F. marsafae described from Egypt does not differ from F. pulcherrima now examined. Characters depicted as diagnostic by GHONAIM et al. (2001): Tibia and first tarsomere with long bristles, more hairy wing and less expanded tip of gonostylus, we found

GROGAN W.L. JR. et al.: The Old World biting midge, Forcipomyia pulcherrima 295 as infraspecific variation depending also on the state of preservation of specimen. As a result we propose to place F. marsafae in the list of junior synonyms of well-known and broadly distributed F. pulcherrima. Figs 1-4. Forcipomyia (Lepidohelea) pulcherrima SANTOS ABREU: 1 habitus of adults, lateral view (from DESSART 1963); 2 male genitalia, ventral view (from CLASTRIER 1956); 3 photograph of male genitalia, ventral view; 4 photograph of apical region of female abdomen and spermathecae, ventral view.

296 Polish Journal of Entomology 82 (4) Figs 5-8. Forcipomyia (Lepidohelea) pulcherrima SANTOS ABREU, photos of larva in ethanol: 5 head and thoracic segments, lateral view; 6 head and thoracic segments, dorsal view; 7 last 3 abdominal segments, dorsal view; 8 last 2 abdominal segments, lateral view. We too noted that most of the numerical values that GHONAIM et al. (2001) listed for adults of F. marsafae and F. pulcherrima are similar; for example, they provided wing lengths of 1.3-1.6 mm for males of F. marsafae, and 1.2-1.48 mm for males of F. pulcherrima. Furthermore, their illustrations of a male wing and that of the only female (allotype) of F. marsafae, are of specimens with nearly all macrotrichia present, whereas their wing illustrations of both sexes of F. pulcherrima are obviously of specimens

GROGAN W.L. JR. et al.: The Old World biting midge, Forcipomyia pulcherrima 297 that have lost nearly all of their macrotrichia due to preservation in ethanol, clearing in a solution of ethanol-phenol or dissection and slide-mounting in Canada balsam. We also noted a similar loss of the elongate setae on the legs and wings of some of our specimens of F. pulcherrima when they were preserved and/or subsequently cleared prior to, or, during dissection and slide-mounting. Furthermore, despite the fact that GHONAIM et al. (2001) stated that the apical region of the gonostylus was not expanded in their specimens of F. marsafae, their Fig. 1J clearly depicts an expanded apex, though not as much as in their Fig. 2J of F. pulcherrima; however, this may be due to an artifact of mounting. Finally, we consider that other differences they pointed out in these two forms, such as the shapes of the spermathecae and aedeagi, are due to intraspecific variation. Figs 9-12. Forcipomyia (Lepidohelea) pulcherrima SANTOS ABREU, SEM photographs of larva: 9 head and prothoracic segment, lateral view; 10 mouth opening, subventral view: mp = maxillary palpus, ep = epipharynx with sensilla; 11 abdominal segments, dorsal view; 12 thoracic and abdominal spicules, dorsal view, at 900X. Finally, as noted in the Introduction and under the synonymy of F. pulcherrima, this species has a confusing history of being recognized as several other species, including

298 Polish Journal of Entomology 82 (4) F. chrysolopha (e. g., WIRTH & MESSERSMITH 1977). However, in his excellent review of the Ceratopogonidae of the Seychelles, CLASTRIER (1983) noted that both F. pulcherrima and F. chrysolopha occur on these islands. CLASTRIER (1983) also provided detailed redescriptions of both sexes of F. chrysolopha, and illustrations of the male genitalia and leg banding patterns as well as female antennal flagella, palpus, apical abdominal genital sclerotizations and spermathecae. It is now clear that the banding patterns on the femora and tibiae of F. chrysolopha differ from those of F. pulcherrima in several ways, especially on the middle leg. Figs 13-17. Forcipomyia (Lepidohelea) pulcherrima SANTOS ABREU, automontage photographs of larval mouthparts dissected from head capsule: 13 messor (ms) and mandible (md), lateral view; 14 mandible, lateral inner view; 15 messor (ms) and its articulation with the torma, lateral view; 16 mola cibarialis, ventral view; 17 detail of mola cibarialis and teeth (tc) on comb.

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302 Polish Journal of Entomology 82 (4) YU Y.-X., LIU J.-H., LIU G.-P., LIU Z.-J, HAO B.-S., YAN G., ZHAO T.-S. 2005. Ceratopogonidae of China, Insecta, Diptera. Military Medical Science Press, Beijing. Vols. 1-2. 1699 pp. [In Chinese]. Received: 7 October 2013 Accepted: 23 October 2013