Species Diversity of Japanese Clusiidae (Diptera: Acalyptrata) with description of 12 new species

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1 Ann. soc. entomol. Fr. (n.s.), 2006, 42 (1) : 1-26 ARTICLE Species Diversity of Japanese Clusiidae (Diptera: Acalyptrata) with description of 12 new species Masahiro Sueyoshi Department of Entomology, PO Box 37012, National Museum of Natural History, CE607, MRC169, Washington, DC USA (Postdoctoral Fellow of Research Abroad, Japan Society for the Promotion of Science) Present address: Forest Zoology Group, Kyushu Research Center, Forestry and Forest Products Research Institute, Kurokami, Kumamoto, Japan Abstract. The Japanese Clusiidae are revised and 23 species, including 12 species new to science, are recognized and keyed. Clusiodes angulosus n. sp., Clusiodes discostylus n. sp., Clusiodes tobi n. sp., Clusiodes usikumuri n. sp., Craspedochaeta varicolor n. sp., Heteromeringia crenulata n. sp., Heteromeringia quadrispinosa n. sp., Heteromeringia sexramifera n. sp., Heteromeringia yamata n. sp., Phylloclusia quadrivittata n. sp., Sobarocephala uncinata n. sp., and Tetrameringia borealis n. sp. were described as new. Distribution records were compiled and mapped. Transitional climatic zone between the warm and cool temperate zones is possibly suggested as one of the most richest temperate area in terms of Japanese clusiid species richness. Hendelia beckeri Czerny 1903, Paraclusia japonica Sasakawa 1957, and Clusiodes plumosus Sasakawa 1964 are broadly distributed over all the Japanese temperate forests, but these are not always most abundant species. The clusiid assemblage was heterogeneous in the species composition among various forest types. The femalebiased sample caught by Malaise traps and distinction of clusiid species composition between temperate and subtropical zones are also pointed out. Résumé. La diversité spécifique des Clusiidae au Japon (Diptera : Acalyptrata), avec la description de 12 espèces nouvelles. Une révision de la faune Japonaise de Clusiidae a été effectuée. Vingt-trois espèces sont reconnues, dont 12 sont nouvelles, et une clef d identification a été composée. Les espèces suivantes ont été décrites : Clusiodes angulosus sp. n., Clusiodes discostylus sp. n., Clusiodes tobi sp. n., Clusiodes usikumuri sp. n., Craspedochaeta varicolor sp. n., Heteromeringia crenulata sp. n., Heteromeringia quadrispinosa sp. n., Heteromeringia sexramifera sp. n., Heteromeringia yamata sp. n., Phylloclusia quadrivittata sp. n., Sobarocephala uncinata sp. n., Tetrameringia borealis sp. n. La distribution géographique de chaque espèce a été compilée et est représentée sur carte. La zone climatique transitoire entre les zones tempérées chaude et froides est considérée comme la plus riche en espèces pour les clusiides du Japon. Hendelia beckeri Czerny 1903, Paraclusia japonica Sasakawa 1957, and Clusiodes plumosus Sasakawa 1964 ont une distribution qui couvre la majeure partie de la foret tempérée Japonaise mais ces espèces ne sont pas toujours les plus abondantes. La composition d espèces est variable selont le type de forêt, formant un assemblage hétérogène pour la faune échantillonée. On remarque encore la forte prédominance des spécimens femelle capturés dans les pièges Malaise ainsi que la différence entre les zones tempérée et subtropicale dans la composition spécifique des clusiides. Keywords: Diptera, Clusiidae, species composition, Japan, new species. The dipterous family Clusiidae, comprising relatively few species (approximately 246 species in 18 genera), has received little scientific attention, including ecological studies. This relative neglect is attributable to two primary conditions. First, the family is of no known economic importance (Ferrar, 1987) and few resources have been available to support research on the family. Second, specimens are apparently uncommon in nature and are infrequently collected using standard sweeping methods (Roháček 1995). Over the last two decades, however, several attempts msuey@affrc.go.jp Accepté le 14 novembre 2005 using various trapping methods (Malaise traps by Sabrosky & Steyskal 1974, Greve & Midtgaard 1986, Sueyoshi et al. 2003; bait-traps by Sasakawa & Mitsuii 1995; flight intercepts by Barrientos et al. 1996) have resulted in larger collections of clusiid flies. Although a combination of taxonomic and ecological studies on the Clusiidae using trapping methods has facilitated research on the biodiversity of this family, few reports have been published. Sueyoshi et al. (2003), using Malaise traps, compared the species composition of clusiids among forest stands of varying ages in temperate zones and suggested that these flies are relatively rich in secondary forests. However, the number and identify of the species from their study exceeded the known fauna of Japanese clusiids, which then included only nine species. 1

2 Masahiro Sueyoshi To facilitate further research on the biodiversity of the Clusiidae, comprehensive taxonomic studies, including keys to clusiid species, are imperative. Six genera and nine species had been recorded from Japan (Sasakawa 1957, 1959, 1963, 1964, 1965a, 1986; Sasakawa & Mitsui 1995), and other unidentified species of Hendelia Czerny 1903, Clusiodes Coquillett 1904, and Paraclusia Czerny 1903 have also been reported (Tamaki 1997; Kubo 2000; Sueyoshi et al. 2003). Although Sueyoshi et al. (2003) recorded 11 species, eight were undetermined and all of their identifications should be reviewed. I also found several species that have not yet been described or recorded from Japan during my faunal research of Japanese Clusiidae. The undetermined specimens include species from the Ryukyus, where no clusiid species have been recorded. To understand the species diversity of Japanese Clusiidae, I studied a series of clusiid flies caught in Malaise traps that were installed in temperate forests of the Ibaraki Prefecture, Honshu, Japan in 1996 and 2002 by the Forestry and Forest Products Research Institute (FFPRI), together with unpublished and published specimens deposited in other museums and institutions. I also examined all undetermined specimens from previous studies (Tamaki, 1997; Kubo 2000; Sueyoshi et al. 2003). This paper has two objectives: 1) a taxonomic review of Japanese Clusiidae, describing new species and providing a key to all Japanese species; 2) clarification of the clusiid species diversity in temperate zones of Japan. Materials and Methods Study area The study includes all of Japan, from Hokkaido to the Ryukyus. The Golden Owl Publisher s Malaise traps of the Towns type were installed in the study site located at the southern edge of the Abukuma Mountains (approximately 36º56 N, 140º35 E, m) near the border between Ibaraki and Fukushima Prefectures in central Japan (fig. 1). This site is in the transitional climatic zone between warm and cool temperate zones. At present, Quercus serrata Thunberg, Fagus japonica Maximowiczi, and F. crenata Blume (Fagaceae) are the dominant species and account for 49 % of the oldgrowth forests (Masaki et al. 1992) in the study site. Six and ten deciduous broadleaf forest stands of different ages were selected in the study site in 1996 and 2002, respectively. These stands included eight secondary forests (0-70 year-old after clear-cutting: SA-SH), a plantation of Japanese cedar mixed with deciduous trees (57 year-old after clear-cutting: PA), three old-growth deciduous forest (120, 173, 177 year-old: OA-OC) (tab. 1). The six stands (SA, SB, SC, SD, PA, OA) in 1996 are the same as those described in Maeto & Makihara (1999) and Sueyoshi et al. (2003), and the remaining six (SB, SC, SD, OA) were studied in Additional specimens were obtained by general sweeping and yellow pan traps in old-growth and secondary forests throughout Japan from Hokkaido to the Ryukyus. Traps and collecting A trap was installed in each of the above stands from 19 Mar-10 Oct in 1996 and from 9 Apr-2 Nov in Specimens were stored in 70% ethanol or pure ethanol+propylene-glycol, with the traps emptied every two weeks. The specimens were sorted in 80% ethanol and then preserved in pure glycerol with the exception of two holotypes preserved dry on pins. The specimens collected by sweeping were preserved dry on pins. Figure 1 Location of study site for species composition of temperate forests in Japan Symbols indicate the sites studied in 1996 ( ), 2002 ( ), and in both 1996 and 2002 ( ). See text for explanation of site code. Number indicates stand age. Preparation of specimens Specimens were observed under a stereoscopic microscope. Male and female abdomens were treated with 10% KOH solution at 40 C for 10 hours, stained with Chlorazol Black E and then observed in pure glycerol. Dissected parts of the abdomen were preserved in pure glycerol in a plastic microvial pinned under the dry specimen. Right or left wings of the holotypes and other specimens were slide-mounted in Euparal. Body length was measured between the anterior apex of the head and the posterior margin of the abdomen, and wing length was measured between the base of the costa and the apex of vein R 4+5. Wing width was measured at widest point. 2

3 Clusiidae of Japan All specimens examined have a serial number label and are deposited in the Biosystematics Laboratory, Graduate Institute of Social and Cultural Studies, Kyushu University, Fukuoka (BLKU) (Cl , , 2125, 2126), the Forestry and Forest Products Research Institute, Tsukuba (FFPRI) (Cl , 2127), the Entomological Laboratory, Kyushu University, Fukuoka (ELKU) (Cl.3001), the Tamaki s personal collection, Moroyama (Cl ), the Entomological Laboratory, Ehime University, Matsuyama (ELEU) (Cl.5001), the Entomological Laboratory, Kyoto Prefectural University, Kyoto (KPU) (Cl ), the Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn (ZFMK) (Cl ), and the Carnegie Museum of Natural History, Pittsburgh (CMNH , , , , , , , , , , ). Terminology Terminology and abbreviations follow McAlpine (1981), excluding the following terms: postpedicel (Stuckenberg 1999; 1st flagellomere: McAlpine 1981), surstylus (Cumming et al. 1995; posterior surstylus: McAlpine 1981), postgonite (Cumming et al. 1995; paramere: McAlpine 1981), phallus (Cumming et al. 1995; aedeagus: McAlpine 1981), 10th abdominal tergite (Griffiths 1981; epiproct: McAlpine 1981), 10th abdominal sternite (Griffiths 1981; hypoproct: McAlpine 1981). The abdominal tergites and sternites are abbreviated T and S, respectively. Abbreviations of localities are as follows: C: City, P: Prefecture, T: Town, V: Village. Systematics A total of 198 specimens were examined. These were assigned to 23 species in 10 genera. Of these, 12 species are new to science and are described below. Craspedochaeta nigripalpis (Steyskal & Sasakawa 1966) and Craspedochaeta flavofemorata Sasakawa 1990, were newly recorded from Japan. The taxa are arranged alphabetically. Table 1. Location, forest type, and forest age of the investigated sites. See text for explanation of the site codes. Site Forest type Forest age (year) OA old growth OB old growth OC old growth PA Plantation of Japanese cedar 57 - SA secondary growth 70 - SB secondary growth SC secondary growth 5 12 SD secondary growth SE secondary growth - 70 SF secondary growth - 0 SG secondary growth - 23 SH secondary growth - 2 Key to the Japanese species of the Clusiidae (modified from Sasakawa 1964) 1. Anteromost orbital seta reclinate (Frey 1960, fig. 5)or lateroclinate (cf. Frey 1960, fig. 4)... 2 Anteromost orbital seta inclinate Mid and hind tibiae with preapical dorsal seta; last section of vein M at least 4 times length of penultimate section of M; 0+2, or 1+2 dorsocentral setae... 3 All tibiae without preapical dorsal seta; last section of vein M less than 4 times length of section of M between crossveins r-m and dm-cu; 0+3 dorsocentral setae.... Allometopon Kertesz 1906 (1 sp. in Japan: A. falcistylus Sasakawa 1964) 3. Two or three orbital setae present, all reclinate (fig. 4A, 7A, 9A, 11A, 13A)... 4 Four orbital setae present, all reclinate except anterior 2nd seta inclinate (fig. 14A, 16A, 17A)... Craspedochaeta Czerny 1903 (3 spp. in Japan) 9 4. Frons 1/3 times length (from anterior ocellus to anterior margin of frons) of width (widest distance between eyes); width between base of antennae more than 3 times diameter of scape (Sasakawa 1998: fig. 7); arista with last flagellomere broader than basal two flagellomeres.... Hendelia Czerny 1903 (1 sp. in Japan: H. beckeri Czerny 1903) Frons more than 1/3 times length of width; distance between base of antennae less than 2 times diameter of scape; arista with last flagellomere narrower or as wide as basal two flagellomeres (fig. 4A, 7A, 9A, 11A, 13A)... Clusiodes Coquillett 1904 (5 spp. in Japan) 5 5. Scutum entirely black.... Clusiodes usikumuri n. sp. Scutum yellowish-brown in ground color, with paired longitudinal black markings Two orbital setae; postocellar seta distinctly shorter than ocellar seta (fig. 4A); intrapostalar seta scarcely longer than acrostichal seta.... Clusiodes angulosus n. sp. Three orbital setae; postocellar seta as long as ocellar seta (fig. 7A, 11A); intrapostalar seta at least 1/4 long as posteromost dorsocentral seta Wing with pale brown marking along vein CuA 1 at posterior side of cell dm (fig. 5C); face or gena black.... Clusiodes plumosus Sasakawa 1964 Wing without pale brown marking along vein CuA 1 at posterior side of cell dm (fig. 5B, D), only at apical 1/3; face and gena yellow, not black Scutum without dark presutural markings; anteromost dorsocentral seta shorter than half length of posteromost dorsocentral seta, inserted posterior to transverse suture.... Clusiodes tobi n. sp. Scutum with dark presutural markings, connecting paired longitudinal stripes on scutum; anteromost dorsocentral seta longer than half length of posteromost dorsocentral seta, inserted anterior to transverse suture.... Clusiodes discostylus n. sp. 9. Scutum yellowish-brown in ground color, with black longitudinal markings; setae on head black.... Craspedochaeta varicolor n. sp. Scutum black in ground color; setae on head pale brown

4 Masahiro Sueyoshi 10. Postpedicel with apical dark markings.... Craspedochaeta nigripalpis (Steyskal & Sasakawa 1966) Postpedicel without dark markings Craspedochaeta flavofemorata Sasakawa Interfrontal seta absent (Figs. 18A, 21A, 23A, 24A, 28A, 30A); all tibiae without ormid tibia with preapical dorsal seta Interfrontal seta present (fig. 27A); all tibiae except fore tibiae without preapical dorsal setae Preapical dorsal seta on mid tibia present Sobarocephala Czerny 1903 (2 spp. in Japan) 13 Preapical dorsal seta on mid tibia absent Scutum with paired longitudinal dark makings in female; T1-5 yellow in both sexes, with medial dark stripe; scutum without dark postsutural markings; anepisternum and katepisternum yellow Sobarocephala mitsuii Sasakawa 1995 Scutum without paired longitudinal dark markings in female; T2-6 in female and T2-4 in male dark brown, with yellow lateral margin or submargin; scutum with broad longitudinal dark postsutural marking; anepisternum and katepisternum yellow, with dark markings.... Sobarocephala uncinata n. sp. 14. Ocellar seta shorter than postocellar seta; arista with last flagellomere narrower or as wide as basal two flagellomeres Ocellar seta as long as postocellar seta (fig. 27A); arista with last flagellomere broader than basal two flagellomeres... Phylloclusia Hendel 1913 (1 sp. in Japan: P. quadrivittata n. sp.) 15. Scutum entirely dark brown; less than four orbital setae present (Figs. 18A, 21A, 23A, 24A); presutural intra-alar seta absent Heteromeringia Czerny 1903 (4 spp. in Japan) 16 Scutum yellow-brown in ground color, with paired longitudinal dark markings; three long and 1 short orbital setae, between anterior 1st and 2nd long orbital setae, present (fig. 30A); presutural intra-alar present.... Tetrameringia McAlpine 1960 (1 sp. in Japan: T. borealis n. sp.) 16. Face yellow; fore femur yellow, at most with dark markings at apex Face black; fore femur entirely dark brown Heteromeringia quadrispinosa n. sp. 17. Postpedicel entirely yellow; notopleuron, meron, and hypopleuron dark brown to black Postpedicel yellow, with dark brown along anterior margin; notopleuron, meron, and hypopleuron yellow.... Heteromeringia yamata n. sp. 18. Apical half of palpus yellow, basal half of palpus black; katepisternum yellow ventrally; fore femur entirely yellow.... Heteromeringia sexramifera n. sp. Palpus entirely yellow, without black markings; katepisternum entirely black; fore femur yellow, with black markings on apical half Heteromeringia crenulata n. sp. 19. Postocellar and prescutellar acrostichal setae absent; proepisternal seta long; proepimeral seta absent Clusia Halliday 1838 (2 spp. in Japan) 20 Postocellar and prescutellar acrostichal setae present; proepisternal seta weak; proepimeral seta present Paraclusia Czerny 1903 (3 spp. in Japan) Face without any markings; T1-8 yellow, with median dark markings.... Clusia flava (Meigen 1830) Face with 2 vittae; T1-8 dark brown Clusia okadomei Sasakawa Scutum with paired lateral dark stripes; anepisternum with dark marking Scutum and anepisternum entirely yellow or testaceous.... Paraclusia ciliata Sasakawa Dark marking on anepisternum not extending to propleuron; katepisternum and meron yellow; wing marking restricted to apex of wing Paraclusia omogensis Sasakawa 1965 Dark marking on anepisternum extending to propleuron; katepisternum and meron dark brown; wing marking expanded to cell cua 1 (fig. 9E).... Paraclusia japonica Sasakawa 1957 Genus Allometopon Kertesz 1906 Allometopon falcistylus Sasakawa 1964 (fig. 2) Figure 2 Distribution map of Japanese Allometopon Kertesz and Clusiodes Coquillett Allometopon falcistylus Sasakawa, Clusiodes angulosus n. sp., C. discostylus n. sp., C. plumosus Sasakawa, C. tobi n. sp. Allometopon falcistylus Sasakawa 1964: 14. Specimens examined. Holotype : JAPAN/ HOKKAIDO/ KUSHIRO:, Nibushi (am See/ Kuttyaroko) / 19.VII.1953/ SYUSIRO Ito, HOLOTYPE/ Allometopon/ falcistylus n.sp./ Det. M. SASAKAWA, vinyl chloride plates, Cl.6001, KPU. Distribution. Japan (Hokkaido: fig. 2). Remarks. Allometopon falcistylus is assigned to the subgenus Allometopon because it has a pair of interfrontal setae and four orbital setae. Allometopon falcistylus is distinguished from other members of the nominal subgenus by a following combination 4

5 Clusiidae of Japan of characters: postpedicel and palpus entirely yellow; scutum brown, with two pairs of longitudinal dark brown markings; anepisternum without any dark markings; two pairs of dorsocentral setae present; no distinct acrostichal seta and presutural intra-alar seta; scutellum yellow; leg entirely yellow; surstylus strongly incurved, flattened, truncated, and triangular in shape apically (Fig. 2B in Sasakawa 1964) (cf. McAlpine 1960; Sasakawa 1974, 1993). The holotype is in good condition, without any lost parts. The abdomen posterior to the 5th abdominal segment has been removed and dissected. Genus Clusia Halliday 1838 Clusia flava (Meigen 1830) Heteromyza flava Meigen 1830: 46. Clusia flava: Schiner 1864: 36; Sasakawa 1963: 18. Remarks. Clusia flava was recorded from Japan by Sasakawa (1963) based on one female labelled as Honshu, Tochigi Prefecture, Kinu Vil (= Oyama City at the present time), 12.VII.1959, Tohru Kodama leg. and there have been no further records since then. I suspect that this female is conspecific with a male also determined by M. Sasakawa as Clusia flava labelled as Japan-Honsyu/ Totigi/ Kinunonuma/ 12.VII.1959/ T. Kodama, which I have found to be conspecific with Paraclusia ciliata Sasakawa The record of Clusia flava from Japan is doubtful, and it needs confirm. Minamiazumigun/ Nagano Pref./ 19,VII,1968/ T. Okadome leg., HOLOTYPE / Clusia/ okadomei n.sp./ Det. M.SASAKAWA, No. 240, Cl.6002, KPU. Distribution. Japan (Honshu: fig. 3). Remarks. Clusia okadomei is distinguished from congeners by the following combination of characters: face with pair of brown stripes; thoracic notopleuron, anepimeron, anepisternum with dark markings; posterior halves of 3rd to 5th abdominal tergites with dark transverse markings; wing with dark infuscation at apex, not at middle (cf. Melander & Argo 1924). It is very similar to C. lateralis (Walker 1849), in two stripes on the female face, the anepisternum with dark marking on upper margin, and wing markings. However, it is distinguished from C. lateralis by the 1st abdominal tergite largely yellow, the 3rd to 5th abdominal tergites largely dark brown. It is also distinguished from C. flava in having a pair of brown stripes on the face and a dark brown abdomen. The holotype is in good condition. The left anteromost orbital seta, right inner vertical seta, left anteromost dorsocentral seta, both inner postalar setae, right posterior notopleural seta, left apical half of the mid tibia and tarsus are lost. Clusia okadomei Sasakawa 1986 (fig. 3) Clusia okadomei Sasakawa 1986: 1. Specimens examined. Holotype : Iwanadome, Figure 3 Distribution map of Japanese Clusia Halliday, Hendelia Czerny, Paraclusia Czerny, and Phylloclusia Hendel Clusia okadomei Sasakawa, Hendelia beckeri Czerny, Paraclusia ciliata Sasakawa, Pa. japonica Sasakawa, Pa. omogensis Sasakawa, Phylloclusia quadrivittata n. sp. Figure 4 Male of Clusiodes angulosus n. sp. (holotype) A: Head in left lateral view; B, C: epandrium complex (epandrium and surstylus), B- left lateral view, C- posterior view, D: hypandrium complex (hypandrium, pregonite, postgonite, phallus) in left lateral view. Abbreviations. bp: basiphallus; c: cercus; dp: distiphallus; ep: epandrium; hyp: hypandrium; if: interfrontal seta; iv: inner vertical seta; oc: ocellar seta; or: orbital seta; ov: outer vertical seta; po: postocellar seta; pog: postgonite; prg: pregonite; s: surstylus; v: vibrissa. Scale bars: A = 0.30 mm; B, C = 0.20 mm; D = 0.10 mm. 5

6 Masahiro Sueyoshi Genus Clusiodes Coquillett 1904 Clusiodes angulosus n. sp. (figs. 2, 4A-D, 5A, 6A, B) Clusiodes sp. 1: Sueyoshi et al. 2003: 182 (in part). Description. Male. Head (fig. 4A) higher than long; frons entirely yellow; ocellar triangle black, partly covered with silvery pubescence; face, parafacial, gena, and postgena yellow; occiput yellow, with dark brown marking lateral to medial occipital sclerite. Head chaetotaxy: 2 orbital setae, 1 ocellar seta, 1 postocellar seta, 1 interfrontal seta, 1 inner vertical seta, 1 outer vertical seta, 1 paravertical seta, 9 postocular setae, 1 vibrissal seta, 4-5 subvibrissal setae, 2 postgenal setae, all pale brown; postocellar seta distinctly shorter than ocellar seta; no distinct genal seta. Scape and pedicel yellow, pedicel with pale brown setae along anterior margin and long, pale brown seta dorsally; postpedicel yellow; arista dark brown, with black hairs longer than width of arista. Mouthparts entirely yellow. Thorax yellow; scutum with dark brown stripe from anterior margin to posterior margin through dorsocentral seta, and stripe from scuto-scutellar suture to postalar wall; scutellum with dark brown marking at lateral corner; mediotergite Figure 5 Left wings of Japanese species of Clusiodes Coquillett and Craspedochaeta Czerny (holotypes) A: Clusiodes angulosus n. sp.; B: Cl. discostylus n. sp.; C: Cl. plumosus Sasakawa; D: Cl. tobi n. sp; E: Cl. usikumuri n. sp.; F: Craspedochaeta flavofemorata Sasakawa; G: Cr. nigripalpis (Steyskal & Sasakawa); H: Cr. varicolor n. sp. 6

7 Clusiidae of Japan dark brown laterally; laterotergite entirely dark brown; dorsal half of anepisternum and anepimeron dark brown. Thoracic chaetotaxy: 1 postpronotal seta, 1+2 dorsocentral setae, 1 postsutural intra-alar seta, 2 postalar setae, 1 scutellar seta, 2 notopleural setae, 1 anepisternal seta, 1 katepisternal seta, all pale brown; intrapostalar seta scarcely longer than acrostichal seta; no distinct acrostichal and presutural intra-alar seta; 2-3 additional setae anterior to scutellar seta. Wing (fig. 5A) pale brown, with dark brown markings on anterior half of apical 1/3 and along middle of veins R 4+5 and CuA 1 ; veins brown. Halter pale yellow. Legs yellow; fore femur with anterior row of ventral setae, mid femur with two posterior rows of ventral setae. Abdomen entirely dark brown. Genitalia (fig. 4B-D): epandrium (fig. 4B-C: ep) dome-like in posterior view; surstylus (fig. 4B-C: s) short, as long as 1/4 height of epandrium, situated ventral to epandrium; hypandrium (fig. 4D: hyp) with postgonite expanded broadly, 2 times higher than hypandrium, with 2 pairs of long setae on ventral margin; postgonite (fig. 4D: pog) as long as basiphallus, with triangular projection at middle bearing 5 minute setulae, apex tapered and acute anteriorly; basiphallus extending to phallus; distiphallus (fig. 4D: dp) entirely membranous. Body length 3.7 mm. Wing length 2.6 mm; width 0.9 mm. Female (fig. 6A-B). Similar to male in appearance except as follows: fore tarsus excluding basitarsus, clypeus, palpus, scape, and pedicel dark brown; frons dark brown along ocellar triangle; face tinged with brown; T6 and S6 slightly longer than T7 and S7, respectively; S6 with short taenia extending to S7; S7 divided medially into 2 sclerites; T8 and S8 (fig. 6A) weakly sclerotized. Genitalia (fig. 6B) entirely membranous, excluding sclerotized spermathecae; ventral receptacle (fig. 6B: vr) elongated, longer than vagina (fig. 6B: v), with round apex. Spermathecal duct approximately 3 times as long as spermatheca, consisting of slender basal and expanded distal sections; distal section 2-3 times as wide as and 4-5 times as long as basal section; spermatheca (fig. 6B: sp) knob-like, with cuff and apical depression. Figure 6 Female of Clusiodes angulosus n. sp. A: postabdomen in left lateral view (Cl.2078); B: genitalia in dorsal view (Cl.2082). Abbreviations. ag: accessory gland; T & S6-8, 10: 6-8th and 10th abdominal sternites and tergites; ag: accessory gland; ov: common oviduct; sp: spermatheca; v: vagina; vr: ventral receptacle. Scale bars: A = 0.25 mm, B = 0.20 mm. Specimens examined. Holotype : Sarugajo/ Kitaibaraki City/ Ibaraki Pref./ Honshu Japan/ 27.viii ix.2002/ H. Gotoh leg., Cl.2125, Holotype/ Clusiodes/ angulosus/ Sueyoshi (red label), BLKU. Paratypes: [HONSHU] Ibaraki P: 6 19, Ogawa, Kitaibaraki C, 14.V.-9.VII.1996 (Cl , , ), FFPRI; 5, same locality, 25.VII.-4.IX.1996 (Cl.2004, , 2032), FFPRI; 1 1, Kaizawa, Kitaibaraki C, 21.V-18.VI.2002 (Cl ), FFPRI; 3 4, Ogawa Reserve Forest, Kitaibaraki C, 21.V-18.VII.2002 (Cl , 2118), FFPRI; 2 2, same locality, 30.VII-28.VIII.2002 (Cl ), FFPRI; 5 2, Ogawa, Kitaibaraki C, 8.V-18.VII.2002 (Cl ), FFPRI; 1, same locality, 30.VII- 14.VIII.2002 (Cl.2095), FFPRI; 1 6, Sarugajo, Kitaibaraki C, 4.VI-3.VII.2002 (Cl ), FFPRI; 1, same locality, VIII.2002 (Cl.2094), FFPRI; 1, Mt.Osho, Kitaibaraki C, 17.VI-3.VII.2002 (Cl.2084), FFPRI; 1 2, same locality, 30.VII-28.VIII.2002 (Cl ), FFPRI. Etymology. The specific epithet is named for the angled postgonite. Distribution. Japan (Honshu: fig. 2). Remarks. Clusiodes angulosus is distinguished from congeners by the following combination of characters: presence of short postocellar setae; presutural dorsocentral seta approximately as large as postsutural dorsocentral seta; face yellow in ground color; two pairs of orbital setae (fig. 4A: or); antenna pale yellow; legs entirely yellow; scutum yellow, with paired lateral dark brown markings; surstylus short, triangular in lateral view, without spines (fig. 4B-C: s) (cf. Malloch 1918; Melander & Argo 1924; Sasakawa 1987; Caloren & Marshall 1998). It is distinguished from Japanese congeners in having a combination of two orbital setae, short postocellar setae (fig. 4A), and a presutural dorsocentral seta. The phylogenetic relationships of C. angulosus to other species are unclear because it has some diagnostic characters of the C. apiculatus and C. gradiator groups, which are two major monophyletic divisions of the genus, established by Caloren & Marshall (1998). It may be assigned to the C. albimanus subgroup of C. apiculata group, established by Caloren & Marshall (1998), by having fore femur with a anterior row of ventral setae, mid femur with two posterior rows of ventral setae, and spermathecal duct three times length of spermatheca (fig. 6B) but C. angulosus also shares a dome-like epandrium and the surstylus situated posteriorly with the C. gradiator group. Clusiodes angulosus also shares two orbital setae (fig. 4A: or), a narrow gena (fig. 4A), three subequal dorsocentral setae, short intrapostalar seta with C. apiculatus Malloch 1922 and C. gradiator McAlpine Although the body color is faded and the left vibrissal seta is absent, the holotype is in good condition. Furthermore, the abdomen is dissected and the left wing is slide-mounted. Some specimens reported in Sueyoshi et al. (2003) as an unidentified species of Clusiodes are assigned here to C. angulosus. Clusiodes discostylus n. sp. (figs. 2, 5B, 7A-D, 8A, B) Hendelia beckeri: Sueyoshi et al. 2003: 182 (in part) (misidentification). Description. Male. Head (fig. 7A) higher than long; frons entirely dark yellow; ocellar triangle black, partly covered with silvery 7

8 Masahiro Sueyoshi Figure 7 Male of Clusiodes discostylus n. sp. (holotype) A: Head in left lateral view; B, C: epandrium complex, B- left lateral view, C- posterior view, D: hypandrium complex in left lateral view. Refer to Fig. 3 for abbreviations. Scale bars: A = 0.30 mm, B, C = 0.20 mm, D = 0.10 mm. pubescence; face entirely dark yellow; parafacial and gena above genal groove dark yellow, with silvery pubescence; gena below genal groove dark yellow, without silvery pubescence; occiput and postgena entirely dark brown excluding yellow ventral half of postgena. Head chaetotaxy: 3 orbital setae, 1 ocellar seta, 1 postocellar seta, 1 interfrontal seta, 1 inner vertical seta, 1 outer vertical seta, 1 vibrissal seta, 1 subvibrissal seta, all black; 3-4 paravertical setae, 8 postocular setae, 2 postgenal Figure 8 Female of Clusiodes discostylus n. sp. (Cl.2097) A: Postabdomen in left lateral view; B: genitalia in left lateral view. Refer to fig. 6 for abbreviations. Scale bars = 0.20 mm. setae, all yellow; ocellar seta slightly longer than postocellar seta; no distinct genal seta. Scape and pedicel yellow; pedicel with black setae along anterior margin and long black seta dorsally and ventrally; postpedicel orange yellow; arista black, with black hairs as long as width of central rib. Mouthparts entirely orangeyellow. Thorax yellow; scutum entirely dark yellow excluding black stripe lateral to dorsocentral seta and black marking connecting these stripes on presutural region; scutellum dark yellow; mediotergite dark brown; dark brown stripe from proepisternum to katatergite through entire postpronotal lobe, ventral margin of notopleuron, and entire anepisternum and anepimeron. Thoracic chaetotaxy: 1 postpronotal seta, 1+2 dorsocentral setae, 1 postsutural intra-alar seta, 2 postalar setae, 2 notopleural setae, 1 scutellar seta, 1 anepisternal seta, 1 katepisternal seta, all black; no distinct acrostichal and presutural intra-alar setae; anteromost dorsocentral seta longer than half of posteromost dorsocentral seta; 1 additional seta anterior to scutellar seta present. Wing (fig. 5B) hyaline, with pale brown markings in anterior half of apical 1/3; veins dark brown. Halter entirely yellow. Legs yellow, with fore tibia slightly dark brown and basitarsus and basal 2 tarsomeres of fore leg entirely black. Abdomen entirely dark brown. Sixth tergum symmetrical, separate from syntergosternite 7+8. Genitalia (fig. 7C-E): epandrium (fig. 7C, D: ep) saddle-like in shape; surstylus (fig. 7C, D: s) shorter than epandrium, round in profile; hypandrium (fig. 6E: hyp) with postgonite triangular in profile, with 2 long setae on anterior margin and 2 at apex; postgonite (fig. 7E: pog) as long as basiphallus, without processes; basiphallus (fig. 7E: bp) extending to apex of phallus; distiphallus (fig. 7E: dp) entirely membranous. Body length 3.9 mm. Wing length 3.3 mm; width 1.2 mm. Female (fig. 8A, B). Similar to male in appearance except as follows: T6 and S6 (fig. 8A: T6, S6) as long as T7 and S7 (fig. 8A: T7, S7), respectively; T8 and S8 (fig. 8A: T8, S8) weakly sclerotized. Genitalia (fig. 8B) entirely membranous, excluding weakly sclerotized spermathecae; ventral receptacle (fig. 8B: vr) longer than vagina (fig. 8B: v), with bowl-like depression at subapical ventral side. Spermathecal duct consisting of slender basal and expanded distal sections; distal section 4-5 times as wide as and same as long as basal section; spermatheca knoblike, with cuff and apical depression. Specimens examined. Holotype : Shikaribetsu/ Shikaoi Town/ Hokkaido/ Hokkaido Japan/ 2.vii.2000/ H. Nakayama leg., Cl.1003, lime green circle, Holotype/ Clusiodes/ discostylus/ Sueyoshi (red label), BLKU. Paratypes: 4. [HONSHU] Ibaraki P: 1, Ogawa, Kitaibaraki C, 6-20.VIII.1996 (Cl.2059), FFPRI; 1, Sarugajo, Kitaibaraki C, 17.VI.-3.VII.2002 (Cl.2096), FFPRI; 1, same locality, VIII.2002 (Cl.2097), FFPRI; 1, Ogawa, Kitaibaraki C, 17.VI.-3.VII.2002 (Cl.2119), FFPRI. Etymology. The specific epithet is named for disc-like surstylus. Distribution. Japan (Hokkaido and Honshu: fig. 2). Remarks. Clusiodes discostylus is distinguished from congeners by the combination of the following characters: presutural dorsocentral seta approximately as large as postsutural dorsocentral seta; gena without modifications; scutum yellow brown, with dark brown stripes lateral to dorsocentral seta; outline of surstylus entire and rounded (fig. 7B, C); apical 1/3 of wing with markings between costa and vein M (fig. 5B); wing 8

9 Clusiidae of Japan cell dm without dark marking; wing vein R 2+3 slightly curved anteriorly but cell r 1 not narrowed at middle (fig. 5B) (cf. Malloch 1918; Melander & Argo 1924; Caloren & Marshall 1998). It is also very similar to C. tobi, described below, but differs from C. tobi by having a scutum without dark markings on the presutural region and a surstylus shorter than the epandrium (fig. 7B, C). Clusiodes discostylus probably belongs to the albimanus subgroup of the genus, established by Caloren & Marshall (1998), by having three orbital setae equally distant from each other, a narrow gena, long intrapostalar seta, three dorsocentral setae approximately equal in length, fore femur with a anterior row of ventral setae, and mid femur with two posterior rows of ventral setae. However, its phylogenetic position within the subgroup is unclear because C. discostylus has a long and broad surstylus and a distiphallus lacking the crescent sclerite at apex. It is similar to the Nearctic C. johnsoni Malloch 1922, and the Palaearcitc C. albimanus (Meigen 1830) in body color and broad surstylus (fig. 7B) and long presutural dorsocentral seta, but distinguished from the former species by well-developed postocellar setae (fig. 8A) and a spermathecal duct that is wider than the spermatheca (fig. 8B), and the latter species by entirely yellow mid and hind legs and yellow thoracic pleuron. The holotype is in good condition, although the left outer vertical seta, left postpronotal seta, right anteromost dorsocentral seta, left fore leg excluding coxa are lost. Furthermore, the abdomen is dissected and left wing is slide-mounted. One of the specimens reported in Sueyoshi et al. (2003) as Hendelia beckeri is identified as this species. Clusiodes plumosus Sasakawa 1964 (figs. 2, 5C, 9A-D, 10) Clusiodes plumosus Sasakawa 1964: 14; 1990: 540. Burmanocephala [sic] sp.: Sueyoshi et al. 2003: 182. Figure 9 Male of Clusiodes plumosus Sasakawa (Cl.1001) A: Head in left lateral view; B, C: epandrium complex, B- left lateral view, C- posterior view, D: hypandrium complex in left lateral view. Refer to fig. 4 for abbreviations. Scale bars: A = 0.50 mm, B, C = 0.20 mm, D = 0.10 mm. Figure 10 Distribution map of Japanese Clusiodes Coquillett, Heteromeringia Czerny, and Sobarocephara Czerny Clusiodes plumosus Sasakawa, C. usikumuri n. sp., Heteromeringia sexramifera n. sp., Sobarocephara uncinata n. sp. Specimens examined. Holotype : Honshu, Japan/ Asiu, Kyoto/ 7.VI.1964/ M. Sasakawa, HOLOTYPE / KPU0184/ Clusiodes/ (s.str.)/ plumosus/ SASAKAWA, Cl.6003, KPU. Paratype: 1. [HONSHU] Kyoto P: Ashiu, Miyama T, 7.VI.1964 (Cl.6008), KPU. Other specimens. [HOKKAIDO] 1, Hyakumatuzawa, Sapporo C, 22.VII.1996 (Cl.1005), BLKU; 1, Kamitoshibetsu, Ashoro C, 10.VIII.1996 (Cl.1007), BLKU. [HONSHU] Ibaraki P: 1, Ogawa, Kitaibaraki C, VII.1996 (Cl.2033), FFPRI; 1, same locality, 25.VII.-7.VIII.1996 (Cl.2034), FFPRI; 1, Ogawa, Kitaibaraki C, 2-18.VII.2002 (Cl.2061), FFPRI; 1, Sarugajo, Kitaibaraki C, same campaign (Cl.2062), FFPRI; 1, Sadanami, Kitaibaraki C, 9-23.IX.2002 (Cl.2060), FFPRI. Kanagawa P: 1, Terayama, Hadano C, 16.VI.2000 (Cl.4003), BLKU. [SHIKOKU] Tokushima P: 1, Mt. Tsurugi, Koyadaira V, 5.VII.1996 (Cl.1009), BLKU. Ehime P: 1, Kamegamori, Omogo T, 4.VII.1996 (Cl.1006), BLKU. [KYUSHU] Kumamoto P: 1, Haki, Izumi V, 23.V.1997 (Cl.1008), BLKU. [RYUKYUS] 2, Yona, Kunigami V, 23.III.2000 (Cl.1001, 1011, 1012), BLKU. Distribution. Japan (Hokkaido, Honshu, Shikoku, Kyushu, and Ryukyus: figs. 2, 10). Remarks. Clusiodes plumosus is distinguished from congeners by the following combination of characters: presence of presutural dorsocentral seta and postocellar seta; three pairs of orbital setae present; scutum and scutellum yellow-brown, with dark brown markings; legs yellow, excluding dark brown fore tarsus; surstylus short and triangular in lateral view, without spines (fig. 9B-C) (cf. Malloch 1918; Melander & Argo 1924; Sasakawa 1987; Caloren & Marshall 1998). Clusiodes plumosus may be assigned to the C. gladiator group of the genus, established by Caloren & Marshall (1998), by having the dorsal seta on the pedicel longer than ventral seta, a loosely plumose arista, deeply incised posterior margin 9

10 Masahiro Sueyoshi of the epandrium, short cerci, surstylus positioned on the posterior margin of the epandrium, 1+2 dorsocentral setae, long postsutural intra-alar seta, and a broad gena. The facial markings of male C. plumosus have two forms: entirely or laterally black (Cl.1001, 1011, 1012, 2033, 2061, 4003), or yellow excluding the black lateral sides and a median oval black marking (Cl.1008, 1009). The former occurs in Honshu and the Ryukyus, and the latter occurs in Shikoku and Kyushu. All females examined from Hokkaido, Honshu, and Shikoku have entirely black faces but no female specimens are known from the Ryukyus. All other morphological structures, including the male genitalia and coloration, are identical between both forms. The holotype is in good condition, although the left inner vertical seta, the left fore and mid tarsi, left hind leg, and right mid leg are lost. The specimens reported in Sueyoshi et al. (2003) as unidentified species of Burmanochorea Frey 1960 (= Burmanocephala [sic]) and Clusiodes, are assigned here to Clusiodes plumosus. Clusiodes tobi n. sp. (figs. 2, 5D, 11A-E, 12A, B) Description. Male. Head (fig. 11A) higher than long; frons yellow; ocellar triangle black, partly covered with silvery pubescence; face entirely yellow; parafacial and gena above genal groove yellow in ground color, with silvery pubescence; gena below genal groove yellow, without silvery pubescence; occiput and postgena yellow, with occiput lateral to median occipital sclerite and dorsal half of postgena dark brown. Head chaetotaxy: 3 orbital setae, 1 ocellar seta, 1 postocellar seta, 1 interfrontal seta, 1 inner vertical seta, 1 outer vertical seta, 1 vibrissal seta, 3-4 subvibrissal setae, all black; 3-4 paravertical setae, 8 postocular setae, postgenal seta, all pale yellow; ocellar seta slightly shorter than postocellar seta; no distinct genal seta. Scape and pedicel yellow; pedicel with pale yellow setae along anterior margin, and long black seta dorsally; postpedicel orange-yellow; arista entirely black except pale yellow on basal 1/5, with black hairs as long as width of arista. Mouthparts yellow. Thorax yellowish-brown, with longitudinal dark markings; scutum entirely yellowish-brown excluding dark brown stripe lateral to dorsocentral seta; scutellum yellow; mediotergite dark brown; narrow dark brown stripe from proepisternum to katatergite through ventral margin of postpronotal lobe and notopleuron, and dorsal margin of anepisternum and anepimeron. Thoracic chaetotaxy: 1 postpronotal seta, 0+3 dorsocentral setae, 1 postsutural intra-alar seta, 2 postalar setae, 2 notopleural setae, 1 scutellar seta, 1 anepisternal seta, 1 katepisternal seta, all black; no distinct acrostichal and presutural intra-alar setae; anteromost dorsocentral seta shorter than half of posteromost dorsocentral seta; 1 additional seta anterior to scutellar seta present. Wing (fig. 5D) hyaline, with dark brown marking on anterior half of apical 1/3; veins pale yellow. Halter entirely yellow. Legs: coxae and trochanters entirely yellow. Abdomen dark brown. Genitalia (fig. 11B-E): epandrium (fig. 11B-D: ep) spherical; surstylus (fig. 11B-D: s) longer than epandrium, bifid apically, with process anteromedially; apical processes of surstylus consisting of narrow anterior process and broad posterior process, anterior process with dense, minute pile-like setulae at apex; hypandrium (fig. 11E: hyp) with small bilobed pregonite and boot-shaped postgonite (fig. 11E: pog); basiphallus (fig. 11E: bp) extending to apex of phallus, longer than postgonite; distiphallus (fig. 11E: dp) entirely membranous. Body length 3.7 mm. Wing length 3.3 mm; width 1.3 mm. Female (fig. 12A, B). Similar to male in appearance except as follows: postpedicel with broad dark brown marking on lateral side: T6 and S6 (fig. 12A: T6, S6) slightly shorter than T7 and S7 (fig. 12A: T7, S7), respectively; S7 divided medially into 2 sclerites; T8 and S8 (fig. 12A: T8, S8) weakly sclerotized. Genitalia (fig. 12B) entirely membranous, excluding sclerotized spermathecae; ventral receptacle (fig. 12B: vr) elongate, longer than vagina (fig. 12B: v), with bowl-like apex. Spermathecal duct consisting of slender basal and expanded distal sections; distal section 4-5 times as wide as and equal in length to basal section; spermatheca knob-like, with cuff and apical depression. Specimens examined. Holotype : Shikaribetsu/ Shikaoi Town/ Hokkaido/ Hokkaido Japan/ 10.vii.2000/ H. Nakayama leg., Cl.1002, lime green circle, Holotype/ Clusiodes/ tobi/ Sueyoshi (red label), BLKU. Paratype: [HOKKAIDO] 1, same locality as holotype, 11.VII.2000 (Cl.1004), BLKU. Etymology. The specific epithet is in reference to the shape of the postgonite; derived from the Japanese term meaning a tool for dealing a log. Distribution. Japan (Hokkaido: fig. 2). Remarks. Clusiodes tobi is distinguished from congeners by the following combination of characters: absence of presutural dorsocentral seta and presence of distinct wing marking at apex (fig. 5D); scutum yellowish-brown in ground color, with dark brown stripes lateral to dorsocentral seta; male epandrium dome-shaped (fig. 11B), posterior margin entire (fig. 11B); surstylus divided into two lobes at apex, anterior lobe broader than half of width of posterior lobe, covered by many minute setae, not spines (fig. 11B-D); male postgonite boot-shaped in lateral view, without distinct incision (fig. 11E); female spermathecal duct short, 3 times as long as spermatheca (fig. 12B) (cf. Malloch 1918; Melander & Argo 1924; Caloren & Marshall 1998). The following characters suggest that Clusiodes tobi belongs to the C. melanostomus species group, established by Caloren & Marshall (1998): lateral position of surstylus with deep cleft at apex; anterior lobe of surstylus covered with stout setae on inner surface; pregonite divided into two lobes, each bearing two setae; apex of postgonite bent medially; female T6 with transverse sclerotized ridge; ventral receptacle without sclerotized disc; spermatheca cylindrical. Clusiodes tobi is very similar to C. clandestinus Caloren & Marshall 1998 in general appearance and genitalic structure, such as the dome-shaped epandrium, but differ in that the postgonite is lacking a notch or apical cleft with stout setae, but less developed than that of C. clandestinus on the inner surface of the anterior lobe of the surstylus. It is also very similar in appearance to C. discostylus, but differs from C. discostylus by the scutum lacking dark markings on the presutural region, the anteromost dorsocentral seta shorter than half of the posteromost dorsocentral seta, and the surstylus longer than the epandrium (fig. 11B-D). The holotype is in poor condition. Many setae on the head and thorax and all the femora, tibiae, and tarsi were lost on the holotype. Furthermore, the abdomen is dissected and left wing is slide-mounted. 10

11 Clusiidae of Japan Figure 11 Male of Clusiodes tobi n. sp. (holotype) A: Head in left lateral view; B-D: epandrium complex, B- left lateral view, C- posterior view, D- anterior view; E: hypandrium complex in left lateral view. Refer to fig. 4 for abbreviations. Scale bars: A = 0.40 mm, B-D = 0.20 mm, E = 0.10 mm. Clusiodes usikumuri n. sp. (figs. 5E, 10, 13A-C) Description. Male. Unknown. Female. Head (fig. 13A) higher than long, entirely black excluding dark yellow face; parafacial and gena without silvery pubescence; occiput and postgena black, with narrow yellow peristomal margin on postgena. Head chaetotaxy: 3 orbital setae, 1 ocellar seta, 1 postocellar seta, 1 interfrontal seta, 1 inner vertical seta, 1 outer vertical seta, 1 vibrissal seta, 4 subvibrissal setae, 3-4 paravertical setae, 8 postocular setae, postgenal setae, all pale brown; ocellar seta slightly longer than postocellar seta; no distinct genal seta. Scape and pedicel yellow, pedicel with black setae along anterior margin and long black seta dorsally; postpedicel orange-yellow; arista black, with black hairs longer than width of arista. Mouthparts: clypeus and palpus entirely dark brown, other parts entirely yellow. Thorax black and yellow; scutum, notopleuron, scutellum, mediotergite, laterotergite, postpronotal lobe, and anepisternum entirely black; anepimeron yellow excluding black posterodorsal margin; other thoracic sclerites yellow. Thoracic chaetotaxy: 1 postpronotal seta, 3 dorsocentral setae, 1 postsutural intraalar seta, 2 postalar setae, 2 notopleural setae, 2 scutellar setae, 1 anepisternal seta, 1 katepisternal seta, all black; no distinct acrostical and presutural intra-alar setae; anteromost dorsocentral seta longer than half of posteromost dorsocentral seta. Wing (fig. 5E) tinged with pale brown markings from cell dm to apex of cells r 1, r 2+3, and r 4+5 ; veins dark brown. Halter entirely pale yellow. Legs yellow, with fore tibia slightly and basitarsus entirely black. Abdomen entirely dark brown: T6 and S6 (fig. 13B: T6, S6) slightly shorter than T7 and S7 (fig. 13B: T7, S7), respectively; intersegmental membrane between S7 and S8 (fig. 13B: T8, S8) as long as S7; posterior margin of S7 deeply incised to anterior 1/3 of sclerite; T8 and S8 (fig. 13B: T8, S8) weakly sclerotized. Genitalia (fig. 13C) entirely membranous, excluding sclerotized spermathecae; ventral receptacle (fig. 13C: vr), longer than vagina, with apex rounded. Spermathecal duct consisting of slender basal and expanded distal sections; distal section 2-3 times as wide as and 6-7 times as long as basal section; spermatheca (fig. 13C: sp) knob-like, with cuff and apical depression. Body length 3.5 mm. Wing length 2.6 mm; width 0.9 mm. Specimens examined. Holotype : Ushikumori/ Taketomi Town/ Iriomote Isl./ Ryukyu Japan/ 7.vii.1998/ M. Sueyoshi leg., Cl.1010, Clusiodes/ sp./ det. B. Merz 2002, red circle, Holotype/ Clusiodes/ usikumuri/ Sueyoshi (red label), BLKU. Etymology. The specific epithet is derived from type locality meaning beautiful forest. Distribution. Japan (Ryukyus: fig. 10). Remarks. Clusiodes usikumuri is distinguished from congeners by the following combination of characters: presence of presutural dorsocentral seta and postocellar seta; three pairs of orbital setae present (fig. 13A); scutum and scutellum entirely black; legs yellow, excluding dark brown fore tibia and tarsus (cf. Malloch 1918; Melander & Argo 1924; Sasakawa 1987; Caloren & Marshall 1998). It can be distinguished from Japanese congeners by its black scutum. Clusiodes usikumuri may be assigned to the C. gladiator group, established by Caloren & Marshall (1998), by having the arista loosely plumose, the dorsal seta of the pedicel longer than the ventral seta, a broad gena, and 1+2 dorsocentral setae. Its phylogenetic relationship to the other members of the C. gladiator group is unclear, however, because the male is unknown. The holotype is in good condition. The left vibrissal seta, and left anteromost and posteromost dorsocentral setae are lost from the holotype, and left fore leg excluding coxa is removed Figure 12 Female of Clusiodes tobi n. sp. (Cl.1004) A: Postabdomen in left lateral view; B: genitalia in left lateral view. Refer to fig. 6 for abbreviations. Scale bars = 0.25 mm. 11

12 Masahiro Sueyoshi Figure 13 Female of Clusiodes usikumuri n. sp. (holotype) A: Head in left lateral view; B: postabdomen in left lateral view; C: genitalia in left lateral view. Refer to fig. 6 for abbreviations. Scale bars: A = 0.40 mm, B, C = 0.20 mm. Figure 15 Distribution map of Japanese Craspedochaeta Czerny Craspedochaeta flavofemorata Sasakawa, C. nigripalpis Steyskal & Sasakawa, C. varicolor n. sp. and glued to card point. Furthermore, the abdomen is dissected and the left wing is slide-mounted. Genus Craspedochaeta Czerny 1903 Craspedochaeta flavofemorata Sasakawa 1990 (figs. 5F, 14A-E, 15) Craspedochaeta flavofemorata Sasakawa, in Sasakawa & Pong 1990: 58. Specimens examined. [RYUKYUS] 1, Oku, Kunigami V, 27.VI.2001 (Cl.1017), BLKU; 2, Nakamagawa Pass, Taketomi T, 2.VII.2001 (Cl.1018, 1020), BLKU; 1, same locality, 5.VII.1998 (Cl. 1021), BLKU; 2, Mt.Urabe, Yonaguni T, 6.IV.1996 (Cl ), BLKU; 1, Haimita, Taketomi T, 3.VII.2001 (Cl.1024), BLKU. Distribution. Malaysia: Sabah (Sasakawa & Pong 1990); Japan (Ryukyus: fig. 15, New record). Remarks. Craspedochaeta flavofemorata is assigned to the C. biseta group of the genus, diagnosed by four pairs of orbital setae (fig. 14A: or) and two distinct pairs of dorsocentral setae (cf. Soós 1962). It is distinguished from other members of this species group by the following combination of characters: setae on head yellowish-brown, not black; posteromost orbital seta distinctly shorter than other orbital setae (fig. 14A); palpus yellow; katepisternum and anepimeron entirely dark brown; legs entirely yellow (cf. Sasakawa 1971; Sasakawa & Pong 1990). It is very similar to C. nigripalpis, but differs from C. nigripalpis by its entirely yellow palpus, postpedicel, and femora. Craspedochaeta nigripalpis (Steyskal & Sasakawa 1966) (figs. 5G, 15, 16A-C) Figure 14 Male of Craspedochaeta flavofemorata Sasakawa (Cl.1018) A: Head in left lateral view; B, C: epandrium complex, B- left lateral view, C- posterior view, D: hypandrium complex (excluding phallus) in left lateral view, E: phallus in ventral view. Refer to fig. 4 for abbreviations. Scale bars: A = 0.40 mm, B-E = 0.20 mm. Tonnoiria nigripalpis Steyskal & Sasakawa 1966: 246. Specimens examined. [RYUKYUS] 1, Yona, Kunigami V, Okinawa P, 11.VII.1998 (Cl. 1016), BLKU; 1, Oku, Kunigami V, Okinawa P, 29.VI.2001 (Cl. 1014), BLKU; 1, Omoto, Ishigaki C, Okinawa P, 29.VI.2001 (Cl. 1015), BLKU; 1, Nakamagawa Pass, Taketomi T, Okinawa P, 2.VII.2001 (Cl. 1019), BLKU. Distribution. Marianas (Saipan), North West New Guiana (Biak) (Steyskal & Sasakawa 1966); Bismarks (Sasakawa 1971); Japan (Ryukyus: fig. 15, New record). Remarks. Craspedochaeta nigripalpis is assigned to the C. biseta 12

13 Clusiidae of Japan Figure 16 Female of Craspedochaeta nigripalpis (Steyskal & Sasakawa) (Cl.1014) A: Head in left lateral view; B: postabdomen in lateral view; C: genitalia in dorsal view. Refer to fig. 6 for abbreviations. Scale bars: A = 0.40 mm, B = 0.25 mm, C = 0.10 mm. group of the genus because of having four pairs of orbital setae (Fig. 16A) and two distinct pairs of dorsocentral setae (cf. Soós 1962). It is distinguished from other members of this species group by the following combination of characters: face and gena dark yellow, not black; posteromost orbital seta distinctly shorter than other orbital setae (Fig. 16A); pedicel dark brown; palpus dark brown; katepisternum and anepimeron entirely dark brown; coxae yellow; femora and tibiae yellow, with dark brown markings (cf. Steyskal & Sasakawa 1966; Sasakawa 1971). It is very similar to C. flavofemorata, but differs from C. flavofemorata in that a dark apical marking on the postpedicel. anepisternum entirely dark brown, katepisternum with dark brown marking largely covering posterior 3/4, other thoracic sclerites dark yellow. Thoracic chaetotaxy: 1 postpronotal seta, 3 dorsocentral setae, 1 presutural intra-alar seta, 1 postsutural intra-alar seta, 2 postalar setae, 2 notopleural setae, 1 scutellar seta, 1 anepisternal seta, 1 katepisternal seta, all black; no distinct acrostichal seta; 1-2 additional setae anterior to scutellar seta present. Wing (fig. 5H) tinged with pale brown; veins dark brown. Halter pale yellow. Legs dark yellow in ground color; femora dark brown, with basal 1/5-1/4 dark yellow; tibiae dark brown, with basal 1/10 dark yellow. Abdomen dark brown. Genitalia (fig. 17C-F): epandrium (fig. 17C, D: ep) dome-like in shape; surstylus (fig. 17C, D: s) half length of epandrium, triangular in profile; pregonite (fig. 17E: prg) triangular, separate from hypandrium, united with postgonite posteriorly; postgonite (fig. 17E: pog) with short setulae ventrally and apically; basiphallus (fig. 17F: bp) cylindrical, with acute projection on posterior margin; distiphallus (fig. 17F: dp) entirely membranous, with narrow longitudinal sclerite extending to subapical portion and apical small complex sclerite. Body length 3.5 mm. Wing length 2.4 mm; width 1.1 mm. Female. Unknown. Specimens examined. Holotype : Omoto/ Ishigaki C/ Ishigaki Isl./ Ryukyu Japan/ 2.vii.2001/ M. Sueyoshi leg., Cl.1013, lime green circle, Holotype/ Craspedochaeta/ varicolor/ Sueyoshi (red label), BLKU. Etymology. The specific epithet is named for having yellow and dark brown colors on the postpedicel and thorax. Distribution. Japan (Ryukyus: fig. 15). Craspedochaeta varicolor n. sp. (figs. 5H, 15, 17A-F) Description. Male. Head (fig. 17A, B) higher than long; frons and ocellar triangle black, with margin along eye and anterior margin of frons dark brown; face dark yellow; parafacial and gena above genal groove dark yellow, with silvery pubescence; gena below genal groove brown, without silvery pubescence; occiput and postgena black, with narrow brown peristomal margin on postgena. Head chaetotaxy: 4 orbital setae, 1 ocellar seta, 1 postocellar seta, 1 inner vertical seta, 1 outer vertical seta, 1 vibrissal seta, 6 subvibrissal setae, paravertical setae, postocular setae, 1 genal seta, postgenal seta, all black; ocellar seta longer than postocellar seta. Scape and pedicel dark yellow; pedicel with black setae along apical margin and long black seta dorsally; postpedicel dark yellow, with apical half dark brown; arista black, with black hairs longer than width of arista. Mouthparts dark yellow excluding dark brown prementum. Thorax dark yellow, with dark brown markings; scutum with large dark brown marking in presutural region, with dark brown postsutural stripe between dorsocentral setae, stripe tapered anteriorly and united with dark marking in presutural region; scutum lateral to postsutural intra-alar seta and lateral postsutural seta and postalar wall slightly tinged with dark brown; scutellum dark brown; mediotergite with median dark brown stripe; laterotergite slightly tinged with dark brown; Figure 17 Male of Craspedochaeta varicolor n. sp. (holotype) A: Head in left lateral view; B: right antenna in left lateral view; C, D: epandrium complex, C- left lateral view, D- posterior view, E: hypandrium complex (excluding phallus) in left lateral view, F: phallus in ventral view. Refer to fig. 4 for abbreviations. Scale bars: A, B = 0.40 mm, C, D = 0.20 mm, E, F = 0.10 mm. 13

14 Masahiro Sueyoshi Remarks. Craspedochaeta varicolor is assigned to the C. biseta group because of having four pairs of orbital setae (fig. 17A) and two pairs of dorsocentral setae (cf. Soós 1962). It is distinguished from other members of this species group by the following combination of characters: frons black in ground color excluding dark brown margin along eye and anterior margin of frons; setae on head all black; posteromost orbital seta distinctly shorter than other orbital setae (fig. 17A); postpedicel dark brown on apical half; katepisternum dark brown in ground color except for dark yellow anterior margin; anepimeron entirely dark yellow (cf. Sasakawa 1971). It is also distinguished from Japanese congeners by its yellow scutum, with dark brown markings. The holotype is in good condition, although the left arista and right scutellar seta are lost. Furthermore, the abdomen is dissected and left wing is slide-mounted. Genus Hendelia Czerny 1903 Hendelia (Hendelia) beckeri Czerny 1903 (fig. 3) Hendelia beckeri Czerny 1903: 84; Sasakawa 1963: 18, 1965b: 230; Sueyoshi et al. 2003: 182 (in part). Hendelia sp.: Tamaki 1997: 148; Sueyoshi et al., 2003: 182. Clusiodes sp. 1: Sueyoshi et al. 2003: 182 (in part). Specimens examined. [HOKKAIDO] 1, Lake Akan, Akan C, 17.VII.1953 (Cl.6004), KPU. [HONSHU] Ibaraki P: 1 13, Ogawa, Kitaibaraki C, 30.IV.-9.VII.1996 (Cl.2024, , ), FFPRI; 2, Ogawa, Kitaibaraki C, 21.V.-5.VI.2002 (Cl ), FFPRI; 1, Kaizawa, Kitaibaraki C, same campaign (Cl.2100), FFPRI; 1, Ogawa Reserve Forest, Kitaibaraki C, same campaign (Cl.2101), FFPRI; 1 1, Mt.Osho, Kitaibaraki C, same campaign (Cl ), FFPRI; 1, Sarugajo, Kitaibaraki C, 17.VI.-3.VII.2002 (Cl.2102), FFPRI. Saitama P: 1, Asuwa, Moroyama T, 2.V.1994 (Cl.4001), Tamaki s personal collection. [KYUSHU] Fukuoka P: 1, Noko, Fukuoka C, 11.IV.1997 (Cl. 1025), BLKU; 1, same locality, 3.IV.1998 (Cl. 1026), BLKU. Distribution. Norway (Greve 1983), Austria, Finland (Soós 1984), Czech Republic (Roháček 1995); Japan (Hokkaido, Honshu, and Kyushu: fig. 3). Remarks. The genus Hendelia is divided into four subgenera, and the monotypic nominal subgenus represented by H. beckeri is distinguished from the other three subgenera by the following characters: only 2 pairs of orbital setae; 1 pair of setulae just behind anterior margin of the frons; postocellar setae absent; female palpus enlarged (Frey 1960). Some of the specimens reported in Tamaki (1997) and Sueyoshi et al. (2003) as unidentified species of Hendelia and Clusiodes are identified here as Hendelia beckeri. Genus Heteromeringia Czerny 1903 Heteromeringia crenulata n. sp. (figs. 18A-F, 19A, 20) Description. Male. Head (fig. 18A) higher than long; frons entirely black excluding yellow anterior 1/3; ocellar triangle black; face entirely yellow; parafacial and gena dark yellow, with silvery pubescence above genal groove; occiput and postgena black. Head chaetotaxy: 3 orbital setae, 1 ocellar seta, 1 postocellar seta, 1 inner vertical seta, 1 outer vertical seta, 1 vibrissal seta, 3 subvibrissal setae, 1-2 paravertical setae, 8 postocular setae, postgenal setae, all black; ocellar seta distinctly shorter than postocellar seta, as short as anteromost orbital seta; no distinct genal seta. Scape and pedicel yellow; pedicel with black setae along anterior margin and long black seta dorsally; postpedicel pale yellow; arista black, with hairs as long as width of arista. Mouthparts: clypeus dark brown; other part entirely yellow. Thorax entirely black; setulae on scutum pale brown. Thoracic chaetotaxy: 1 postpronotal seta, 3 dorsocentral setae, 1 postsutural intra-alar seta, 2 postalar setae, 2 notopleural setae, 1 scutellar seta, 1 anepisternal seta, 1 katepisternal seta, all black; no distinct presutural intra-alar and acrostical setae; anteromost dorsocentral seta shorter than half length of posterior 2 dorsocentral seta; 1 additional seta anterior to scutellar seta present. Wing (fig. 19A) hyaline, slightly tinged with pale brown, with dark marking on anterior half of apex; veins pale yellow. Halter entirely pale yellow. Legs yellow, with tibia and entire fore tarsus and apical half of fore femur black. Abdomen dark brown. Genitalia (fig. 18B-F): epandrium (fig. 18B, C: ep) dome-like in shape; cerci (fig. 18B, C: c) separate from each other; surstylus (fig. 18B, C: s) shorter than half length of epandrium, anterior margin with perpendicular ridge, apex with 7 short spines; pregonite (fig. 18D, E: prg) asymmetrical, with 2 lobes, anterior lobe large with 2 long setae at apex, posterior lobe small with minute setulae, 1 setula Figure 18 Male of Heteromeringia crenulata n. sp. (holotype) A: Head in left lateral view; B, C: epandrium complex, B- left lateral view, C- posterior view, D: hypandrium complex (excluding distiphallus) in left lateral view, E: right pregonite and postgonite in left lateral view; F: phallus in left lateral view. Refer to fig. 4 for abbreviations. Scale bars: A = 0.50 mm; B, C, F = 0.10 mm; D, E = 0.25 mm. 14

15 Clusiidae of Japan present between these 2 lobes; postgonite asymmetrical: left postgonite (fig. 18D: pog) quadrate, as long as anterior lobe of pregonite; right postgonite (fig. 18E: pog) tapering apically; basiphallus (fig. 18F: bp) cylindrical and asymmetrical, with acute projection on posterior right side; distiphallus (fig. 18F: dp) consisting of 2 longitudinal sclerites united with basiphallus basally and branched into 4 and 3 processes apically; right sclerite branched to 1st branch (fig. 18F: α) and 2nd branch (fig. 18F: β) at apex of distiphallus; 1st branch divided into 2 tube-like sclerites; 2nd branch divided into 2 plate-like sclerites and composing longest shaft of distiphallus together with longitudinal sclerite of left sclerite; left sclerite divided into 3 sclerites at middle of distiphallus; 2 sclerites expanded, sawlike (fig. 18F: γ), with remaining sclerite elongate, expanding to apex. Body length 3.0 mm. Wing length 2.3 mm; width 0.9 mm. Female. Unknown. Specimens examined. Holotype : Tomioka/ Yokohama City/ Kanagawa Pref./ Honshu Japan/ 21.v.2001/ K. Kubo leg., Cl.4004, lime green circle, Holotype/ Heteromeringia/ crenulata/ Sueyoshi (red label), BLKU. Paratype. [HONSHU] Chiba P: 1, Ichinomiya T, 12.VI.1999 (Cl.4010), BLKU. Figure 19 Left wings of Japanese species of Heteromeringia Czerny, Paraclusia Czerny, Phylloclusia Hendel, Sobarocephala Czerny, and Tetrameringia McAlpine A-D, F-H: holotype; E: Cl A: H. crenulata n. sp.; B: H. quadrispinosa n. sp.; C: H. sexramifera n. sp.; D: H. yamata n. sp.; E: Pa. japonica Sasakawa; F: Phylloclusia quadrivittata n. sp.; G: S. uncinata n. sp.; H: T. borealis n. sp. 15

16 Masahiro Sueyoshi Etymology. The specific epithet is named for having a saw-like sclerite on the phallus. Distribution. Japan (Honshu: fig. 20). Remarks. Heteromeringia crenulata is distinguished from congeners by the following combination of characters: gena and palpus entirely yellow; thorax entirely black; wing with dark marking along veins R 2+3 and R 4+5 (fig. 19A); halter entirely yellow; coxae entirely yellow; distal 1/3 and half of fore femur and tibia respectively and entire fore tarsus black; surstylus bent inwards, with round apex bearing short spine-like setae (fig. 18C); postgonite asymmetrical, quadrate and tapering in lateral view (fig. 18D, E); right branch of distiphallus divided into 4 branches at apex (fig. 18F); cerci separated from each other (fig. 18C) (cf. Melander & Argo 1924; McAlpine 1960; Sasakawa 1966). It is very similar to H. sexramifera, described below, but differs from the latter species by having black setae on the head, a yellow palpus, a black thorax, a black katepisternum, and black markings on the apical half of fore femur. The holotype is in good condition, but both vibrissal setae are lost. Furthermore, the abdomen is dissected and left wing is slide-mounted. Heteromeringia quadrispinosa n. sp. (figs. 19B, 20, 21A-F, 22A, B) Description. Male. Head (fig. 21A, B) higher than long, entirely black excluding lateral margin of frons, and gena above genal groove dark yellow, with silvery pubescence. Head chaetotaxy: 3 orbital setae, 1 ocellar seta, 1 postocellar seta, 1 inner vertical seta, 1 outer vertical seta, 1 vibrissal seta, 4 subvibrissal seta, paravertical setae, postocular setae, postgenal setae, all black; ocellar seta shorter than postocellar seta; no distinct genal seta. Scape and pedicel black; pedicel with black setae along anterior margin and long black seta dorsally; postpedicel orange-yellow; arista dark brown, with minute hairs shorter than width of arista. Mouthparts black excluding yellow labella. Thorax entirely dark brown; scutum entirely covered with black setulae. Thoracic chaetotaxy: 1 postpronotal, 3 dorsocentral, 1 postsutural intra-alar, 2 postalar, 2 notopleural, 1 scutellar, 1 anepisternal, 1 katepisternal setae, all black; no distinct acrostical and presutural intra-alar setae; anteromost dorsocentral seta shorter than half of posterior 2 dorsocentral setae; 1 additional seta anterior to scutellar seta present. Wing (fig. 19B) tinged with pale brown, and with dark marking along apical half of veins R 2+3, R 4+5, and M; veins pale brown. Halter pale yellow, with dark brown knob. Legs entirely yellow, except fore tibia and fore tarsus and apical half of fore femur black. Abdomen dark brown. Genitalia (fig. 21C-F): epandrium (fig. 21C, D: ep) dome-like in posterior view; cerci united, large and triangular; surstylus (fig. 21C, D: s) half length of epandrium, with blunt apex bearing short spine-like setae; pregonite united with hypandrium, with 6 setae on posterior margin, apex expanded ventrally, with 2 stout setae (fig. 21E: pog); postgonite shorter than pregonite, rod-like; basiphallus cylindrical, posterior margin expanded as plate-like sclerite, connecting with anterior margin of subepandrial sclerite; distiphallus (fig. 21F) elongate and strap-like, consisting of 2 lateral longitudinal sclerites, connecting with basiphallus at Figure 20 Distribution map of Japanese Heteteromeringia Czerny, Sobarocephala Czerny, and Tetrameringia McAlpine Heteteromeringia crenulata n. sp., H. quadrispinosa n. sp., H. yamata n. sp., Sobarocephala mitsuii Sasakawa, Tetrameringia borealis n. sp. base; right longitudinal sclerite branched into sclerite extended straightly to bell-like apex of distiphallus (fig. 21F: α); left sclerite plate-like, consisting of bifid apex with sickle-shaped (fig. 21F: β) and tube-like projections (fig. 21F: γ). Body length 4.3 mm. Wing length 2.7 mm; width 1.1 mm. Female (fig. 22A, B). Similar to male in appearance except as follows: scape and pedicel yellow, postpedicel with apical margin dark brown; T6 and S6 (fig. 22A: T6, S6) longer than T7 and S7 (fig. 22A: T7, S7), respectively; intersegmental membrane between S7 and S8 shorter than S8; T8 and S8 (fig. 22A: T8, S8) weakly sclerotized. Genitalia (fig. 22B) entirely membranous, excluding sclerotized spermathecae; main body of ventral receptacle short, as long as spermatheca, with remarkably elongate projection, longer than entire postabdomen (fig. 22B: vr). Spermathecal duct same diameter throughout; spermatheca (fig. 22B: sp) knob-like, with cuff and apical depression. Specimens examined. Holotype : Tomioka/ Yokohama City/ Kanagawa Pref./ Honshu Japan/ 4.vi.2002/ K. Kubo leg., Cl.4005, lime green circle, Holotype/ Heteromeringia/ quadrispinosa/ Sueyoshi (red label), BLKU. Paratypes 1 4. [HONSHU] Chiba P: 2, Mt. Gundari, Ichinomiya T, 27.V.2000 (Cl. 4008, 4009), BLKU. Kanagawa P: 1, Kamigo, Yokohama C, 1.VI.2000 (Cl.4006), BLKU; 1, Hitorizawa, Yokohama C, 14.VII.2002 (Cl.4007), BLKU. [SHIKOKU] Ehime P: 1, Omogo V, 14.VII.1952 (Cl.3001), ELKU. Etymology. The specific epithet is named for the four spines on the pregonite. Distribution. Japan (Honshu and Shikoku: fig. 20). Remarks. Heteromeringia quadrispinosa is distinguished from congeners by the combination of the following characters: ocellar seta short, less than 1/3 length of postocellar seta; arista with minute hairs shorter than width of arista (fig. 21A); 16