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Kato, Makoto, Kawakita, Atsushi, Goto, Ryutaro, Okamoto, Tomoko, Kobayashi, Chisato, Imada, Yume, Nakase, Yuta, Nishioka, Tatsuki, Chanthavong, Bakham, Keothumma, Khamsing, Kosaka, Yasuyuki (2021): Community-level plant-pollinator interactions in a Palaeotropical montane evergreen oak forest ecosystem. Journal of Natural History 54 (33-34): 2125-2176, DOI: 10.1080/00222933.2020.1837977, URL: http://dx.doi.org/10.1080/00222933.2020.1837977

Litoleptis kiiensis Imada & Kato sp. n. [Japanese name: kii-jingasa-shitone-abu] (Figs. 3 c, 3 d, 4 b, 6) Description. Body length: 2.6 mm (n= 6) in male, 3.0 mm (n= 7) in female. Wing length: 2.6 mm (n= 7) in male, 2.8 mm (n= 7) in female. Head Frons bare in male, bare or having a pair of setae in female. Antenna 0.55 mm (n= 3). Antenna 1.0 times in male and 1.3 times in female as long as diameter of eye. First flagellomere with enlarged base; with distinctive depression at middle part (at basal 1 / 3 ��� 2 / 3) of anterior margin; apex slightly enlarged (wider than midpoint width) (Fig. 6 a). Antenna covered with short setae consistently. Male (Fig. 6 b) Gonostylus with semicircle-shaped extension apically, pointed inwardly at apex; smooth in posterior edge of basal major part. Sperm sac bulbous hardly visible. Lateral ejaculatory process integrated into sperm sac basally, but freely elongate apically. Ejaculatory apodeme short, round at anterior margin of gonocoxite. Less than four bristles at inner corner of ventral surface of gonocoxite. Base of aedeagus connected with paramere at thick and wide joint. Paramere widest at connection with aedeagus, approximately three times as wide as narrowest point. Apex of aedeagus flat with shallow incision at apex. Female (Fig. 6 c) Spermatheca membranous, cone-shaped. Spermathecal duct accessory gland with short, straight sclerotized columnar duct, arising at less than two-thirds distal length from genital chamber to spermatheca. Common spermathecal duct absent. Genital chamber small. Type material. Holotype. JAPAN [HONSHU] 1 ♂, emerged on 28.III. 2011 from larva collected by MK on 22.XI. 2011 at Akabanegawa, Mie Pref (Fig. 1: 18), ���Rh 0109���, NMNS. Paratypes. 2 ♂, 9 ♀ (Rh 0110���0132) emerged from 27.III��� 30.IV. 2011, from the larvae collected by MK on 22.XI. 2011 at same locality as holotype, KUHE. Additional materials. In total, 58 specimens were collected. All following materials were obtained as larvae. All specimens are stored in Kyoto University (KUHE). JAPAN [HONSHU] 13 ♂, 15 ♀ (Rh 0133���0161) emerged on 30.III. 2014 from larvae collected by YI on 9.III. 2014 at Nanataki, Wakayama Pref (Fig. 1: 12). 7 ♂, 7 ♀ (Rh 0162���0175) emerged during 3.IV. 2014 from larvae collected by YI on 9.III. 2014 at Hyakkennzannkeikoku, Wakayama Pref (Fig. 1: 13). 4 ♂, 2 ♀ (Rh 0176���0181) emerged on 30.III. 2014 from larvae collected by YI on 10.III. 2014 at Takinohai, Wakayama Pref (Fig. 1: 15). 2 ♂ (Rh 0 182, 0183) emerged during 7���9.IV. 2009 from larvae collected by MK on 1.IV. 2009 at Youjigawa, Wakayama Pref (Fig. 1: 16). 4 ♂, 4 ♀ (Rh 0212���0223) emerged during 9���11.IV. 2010 from larvae collected by MK on 3. IV 2010 at Shimajigawa, Mie Pref (Fig. 1: 19). Etymology. The specific epithet is a noun in apposition, taken from the Kii peninsula where this species exists. Distribution. Japan (Honshu: Wakayama and Mie Prefectures) in Japan (Fig. 1). Natural history. Larvae of this species are thallus miners of Reboulia hemisphaerica (Marchantiales: Aytoniaceae) growing on rocky or clayey slopes in both evergreen and deciduous forests (Fig. 11 f). This species prefers sunny and relatively arid microhabitats compared to the habitat of L. japonica, depending on the microhabitat preference of its host-plant. Adults emerged in spring (March-July, in laboratory condition). Diagnosis. Litoleptis kiiensis resembles L. niyodoensis and himukaensis, and can be distinguished from them by the following characters: first flagellomere with distinctive depression at basal one-third to two-thirds of dorsal margin and swollen at apex; gonostylus with semicircle-shaped extension at apex (as opposed to square-shaped in the other two species), and smooth posterior edge of basal part (as opposed to undulate dorsal edge in the other two species); less than four bristles at posteromedial margin of ventral surface of gonocoxite; spermatheca conicalshaped in female (as opposed to round-shaped in L. niyodoensis)., Published as part of Imada, Yume & Kato, Makoto, 2016, Bryophyte-feeding of Litoleptis (Diptera: Rhagionidae) with descriptions of new species from Japan, pp. 41-58 in Zootaxa 4097 (1) on pages 47-48, DOI: 10.11646/zootaxa.4097.1.2, http://zenodo.org/record/271005

Biology of the Japanese Litoleptis All Japanese Litoleptis species were found to be thallus-miners of thallose liverworts belonging to Aytoniaceae and Conocephalaceae (Marchantiopsida, Marchantiophyta) growing on moist riverine slopes or cliffs in evergreen or deciduous forests (Fig. 11 a). All Litoleptis species in Japan are univoltine, and their larval stages spanned almost eleven months, from spring till late winter. The mines were formed within photosynthetic tissue of the thalli particularly near the upper side but not in the upper epidermis, and sometimes easily were seen through the upper thallus (Fig. 11 b). The mines were linear at first, but occasionally became radiate or marked with blotches later. The number of larvae per thallus was usually one. The larvae underwent pupation inside the mine in early spring (Fig. 11 c). The pupal stage spanned approximately a couple of weeks and then the adults emerged, which lived for only a few days. The adult flies were inactive and their feeding behavior has not been observed so far, nevertheless their labella were well-developed (Fig. 2 a). The adults were seldom found in the field; therefore, it was difficult to observe their behavior. We did however observe in the field a L. japonica female ovipositing at the apical-dorsal part of the thallus of their host-plant later in spring. Each Japanese species of Litoleptis fed on only a single liverwort genus as their host-plant: L. japonica feed on Conocephalum, L. asterellaphile on Asterella, and the other four species on Reboulia. Among the host-plants of the Japanese Litoleptis, Conocephalum conicum species complex and Reboulia hemisphaerica are common in Japan, whereas Asterella odora is extremely rare and whose distribution is restricted in the Kanto Region (Iwatsuki 2001). In general, the Japanese Litoleptis species appeared to be locally distributed in limited area except for L. japonica, nevertheless the mines of Litoleptis spp. were found relatively frequently in each population. In Japan, the association between liverworts and Litoleptis spp. seemed to be as common as it is in Micropterigidae, the most basal moth family (Kristensen 1984). The micropterigid moths in Japan feed on liverworts and collectively have a wide distribution in Japan (Imada et al. 2011). In particular, all the Japan endemic species of Micropterigidae belonging to four endemic genera, feed only on Conocephalum liverworts (Imada et al. 2011). The distribution pattern of these micropterigid species in Japan, which is local and allopatric (Imada et al. 2011), is in contrast to that of Litolepis japonica, which is widespread and covers the main islands of Japan (Hokkaido, Honshu, Shikoku, Kyushu), even though these two lineages share the same host-plant species., Published as part of Imada, Yume & Kato, Makoto, 2016, Bryophyte-feeding of Litoleptis (Diptera: Rhagionidae) with descriptions of new species from Japan, pp. 41-58 in Zootaxa 4097 (1) on page 54, DOI: 10.11646/zootaxa.4097.1.2, http://zenodo.org/record/271005, {"references":["Iwatsuki, Z. (2001) Mosses and liverworts of Japan. Heibonsha Ltd, Tokyo, 355 pp.","Kristensen, N. P. (1984) Studies on the morphology and systematics of primitive Lepidoptera (Insecta). Steenstrupia, 10, 141 - 191.","Imada, Y., Kawakita, A & Kato, M. (2011) Allopatric distribution and diversification without niche shift in a bryophyte-feeding basal moth lineage (Lepidoptera: Micropterigidae). Proceedings of the Royal Society B, 278, 3026 - 3033. http: // dx. doi. org / 10.1098 / rspb. 2011.0134"]}

Litoleptis asterellaphile Imada & Kato sp. n. [Japanese name: saihai-shitone-abu] (Figs. 3 j, 3 k, 4 f, 10) Description. Body length: 2.3 mm (n= 5) in male, 2.4 mm (n= 1) in female. Wing length: 2.3 mm (n= 6) in male, 2.6 mm (n= 2) in female. Head Frons bare in both sexes. Antenna 0.31 mm (n= 2). Antenna 0.8 times in male and 1.2 times in female as long as diameter of eyes. First flagellomere oval and enlarged near base, elongate as long tapering segment, sharpened like a straight long hair at tip; in male, flagellomere sometimes sharply curved at middle (Fig. 10 a). Male (Fig. 10 b) Gonostylus slightly tapering toward apex (basal width at most 1.8 times as wide as width of narrowest part), posterior surface of apex hollowed and posterior margin projected as truncated, upward apex inwardly. Sperm sac bulbous not visible in ventral view. Lateral ejaculatory process baculiform. Ejaculatory apodeme moderately long but not reaching anterior margin of gonocoxite. Posteromedial margin in ventral surface of gonocoxite bear one bristle. Aedeagus smoothly connected with paramere at wide joint. Female (Fig. 10 c) Spermatheca membranous, long and narrow, less than half as wide as long. Spermathecal duct accessory gland with short, sclerotized columnar duct curves middle at right angle, arising at approximately three-fourth distal length from genital chamber to spermathecal ducts. Common spermathecal duct absent. Genital chamber oval, very narrow, occupying fraction of sternite 9 area. Type material. Holotype. JAPAN [HONSHU] 1 ♂, emerged on 22.IV. 2011 from the larva collected by MK on 14.XI. 2010 at Nakatsugawa-keikoku, Saitama Pref (Fig. 1: 24), ���Rh 0204���, NMNS. Paratype. 5 ♂, 2 ♀ (Rh 0205���0211), emerged on 2���22.IV. 2011 from the larvae collected by MK on 14.XI. 2010 at the same place (Fig. 1: 24). Etymology. The specific epithet is a noun in apposition, formed by the combination of ��� asterella ���, which being named after the genus of their host-plant, and ���phile���, a suffix that derived from Latin and means ���lover of���. Distribution. Japan (Honshu: Saitama Prefecture) (Fig. 1: 24). Natural history. Habitat of this species is rocky cliff along streams and roads in deciduous forests. Larvae of this species are thallus miners of Asterella odora (Marchantiales: Aytoniaceae). Adults emerged in spring (April, in laboratory condition). Diagnosis. Litoleptis asterellaphile is smaller in body size than the other species of Litoleptis known in Japan. This species can be distinguished from the other Litoleptis species in Japan by the following characters: gonostylus slightly tapering toward apex (basal width at most 1.8 times as wide as width of narrowest part), posterior surface of apex hollowed and posterior margin projected as truncated, upward apex inwardly; posteromedial margin in ventral surface of the gonocoxite bear one bristle; lateral ejaculatory process baculiform., Published as part of Imada, Yume & Kato, Makoto, 2016, Bryophyte-feeding of Litoleptis (Diptera: Rhagionidae) with descriptions of new species from Japan, pp. 41-58 in Zootaxa 4097 (1) on pages 52-53, DOI: 10.11646/zootaxa.4097.1.2, http://zenodo.org/record/271005

Imada, Yume, Kato, Makoto (2016): Bryophyte-feeding of Litoleptis (Diptera: Rhagionidae) with descriptions of new species from Japan. Zootaxa 4097 (1): 41-58, DOI: http://dx.doi.org/10.11646/zootaxa.4097.1.2

Litoleptis izuensis Imada & Kato sp. n. [Japanese name: izu-jingasa-shitone-abu] (Figs. 3 i, 4 e, 9) Description. Body length: 3.4 mm (n= 2) in male. Wing length: 4.3 mm (n= 2) in male. Head Frons bare in male, with sparse hairs in female. Antenna 0.32 mm (n= 2). Antenna 1.1 times as long as diameter of eyes. First flagellomere straight and gradually tapering towards apex, slightly bulbous dorsally near apex; densely covered with comparatively long hairs (Fig. 9 a). Male (Fig. 9 b) Gonostylus with triangle-shaped extension posteriorly, tapering toward apex, widest point at base twice as wide as apex. Sperm sac posteriorly cylindrical in internal aedeagual process, and widen and forked toward anterior end. Lateral ejaculatory process small and round-shaped without elongation, weakly sclerotized. Ejaculatory apodeme short, posterior end broad and tapering toward anterior margin of gonocoxite. Posteromedial margin in ventral surface of gonocoxite bear less than four bristles. Aedeagus tapering toward apex; smoothly connected with paramere. Paramere consistently slender, approximately six times as narrow as width of gonostylus at basal widest point. Female. Unknown. Type material. Holotype. JAPAN [HONSHU] 1 ♂, holotype, emerged on 7.IV. 2009 from larva collected by MK on 22.III. 2009 at Izu Oshima, Tokyo Metropolitan, Japan (Fig. 1: 23), ���Rh 0202���, NMNS. Paratype. 1 ♂ (Rh 0203), emerged on 2.IV. 2009 from larva collected by MK on 22.III. 2009 at same locality as holotype (Fig. 1: 23), KUHE. Etymology. The specific epithet is a noun in apposition, taken from the Izu Oshima Island where this species was found. Distribution. Japan (Honshu: Tokyo Metropolitan) (Fig. 1). Natural history. Habitat of this species is shaded clayey slope along streams and roads in evergreen Castanopsis forests. Larvae of this species are thallus miners of Reboulia hemisphaerica. Adults emerged in spring (April, in laboratory condition). Diagnosis. Litoleptis izuensis can be distinguished from the congeners by the following characters: gonostylus greatly wide at base with triangle-shaped extension posteriorly, tapering toward apex, widest point at base twice as wide as apex; paramere narrow, approximately six times as narrow as width of gonostylus at basal widest point; lateral ejaculatory process small and round-shaped without elongation; sperm sac developed and forked at anterior end., Published as part of Imada, Yume & Kato, Makoto, 2016, Bryophyte-feeding of Litoleptis (Diptera: Rhagionidae) with descriptions of new species from Japan, pp. 41-58 in Zootaxa 4097 (1) on pages 50-51, DOI: 10.11646/zootaxa.4097.1.2, http://zenodo.org/record/271005

Litoleptis japonica Imada & Kato sp. n. [Japanese name: jagoke-shitone-abu] (Figs. 3 a, 3 b, 4 a, 5) Description. Body length: 3.0 mm (n= 3) in male, 3.0 mm (n= 12) in female. Wing length: 3.2 mm (n= 3) in male, 3.4 mm (n= 9) in female. Head (Fig. 3 a, b) Background color dark brown, clypeus in male generally darker than face. Frons bare in male, generally covered with sparse setae in female. Antennal length 0.5 mm (n= 3). Antenna 0.75 times as long as diameter of eye, covered with dense, appressed microsetae, consistently except distal half of first flagellomere (Fig. 5 a). First flagellomere laterally compressed, oval and enlarged at basal half, sharply tapering toward apex, sharpened to be needle-like at tip and covered with sparse and rough microsetae. Male (Fig. 5 b) Gonostylus wide and thick at basal main component and tapering steeply toward tip (apex approximately 3 times narrower than width at base), with forceps-like extension hollowed dorsally and bifid unevenly at apex. Sperm sac barely visible. Lateral ejaculatory process absent. Ejaculatory apodeme relatively long, reaching anterior margin of gonocoxite. Dorsal surface of gonocoxite with clear medial chevron in anterior margin. Posteromedial margin in ventral surface of gonocoxite without bristles. Aedeagus deeply notched at tip, smoothly connected with paramere at base. Paramere widest at connection with aedeagus, approximately three times wider than narrowest point. Female (Fig. 5 c) Spermatheca spherical, not sclerotized. Spermathecal duct accessory gland with short, sclerotized columnar duct, curved at middle to make a right angle. Spermathecal duct accessory glands arising at base of spermatheca. Ring at base of spermathecal duct lightly sclerotized. Common spermathecal duct nearly absent. Genital chamber moderately sized, oval. Type material. Holotype. JAPAN [HONSHU] 1 ♂, emerged on 20.IV. 2011 from larva collected by MK on 2.IV. 2011 at Haruno-cho, Shizuoka Pref (Fig. 1: 20), “Rh 0001”, NMNS. Paratype. 2 ♂, 5 ♀(Rh 0002–0008), emerged 20–22.IV. 2011 from larvae collected by MK on 2.IV. 2011, same locality as holotype, Shizuoka Pref (Fig. 1: 20), KUHE. Additional materials. In total, 53 specimens were collected. All following materials were obtained as larvae. All specimens are stored in Kyoto University (KUHE). JAPAN [HOKKAIDO] 1 ♀(Rh 0009), emerged 1.V. 2010 from larva collected by MK on 19.X. 2009 at Aizan-kei, Hokkaido Pref (Fig. 1: 1); 2 ♀(Rh 0 0 10, 0011), emerged 16.VI. 2012 from larvae collected by MK on 11.VI. 2012 at Mt. Daisengen, Hokkaido Pref (Fig. 1: 2). [HONSHU] 1 ♀(Rh 0012), emerged 29.IV. 2010 from larva collected by MK on 24.XI. 2009 at Sekikawa-mura, Niigata Pref (Fig. 1: 3); 1 ♀(Rh 0013), emerged 18.V. 2010 from larva collected by MK on 26.IV. 2010 at Hakusan, Fukui Pref (Fig. 1: 5). 25 ♀(Rh 0014–0038), emerged 21.IV– 4.V. 2010 from larvae collected by MK on 6.IV. 2010 at Kibune, Kyoto Pref (Fig. 1: 6). 5 ♀(Rh 0039–0043), emerged 30.III– 2.IV. 2011 from larvae collected by MK on 20.II.. 2011 at Akame-no-taki, Mie Pref (Fig. 1: 7). 2 ♂, 3 ♀(Rh 0044–0048), emerged 26.III– 1.IV. 2012 from larvae collected by MK on 4.III. 2012 at Koto-no-taki, Wakayama Pref (Fig. 1: 14). 2 ♀(Rh 0 0 49, 0050), emerged 14.IV. 2009 from larvae collected by MK on 1.IV. 2009 at Kuki, Wakayama Pref (Fig. 1: 17). 1 ♀(Rh 0051), emerged 20.IV. 2011 from larva collected by MK on 2.IV. 2011 at Wayama-touge, Shizuoka Pref (Fig. 1: 20). 1 ♀(Rh 0052), emerged 26.IV. 2008 from larva collected by MK on 20.IV. 2008 at Shirokura-kyo, Shizuoka Pref (Fig. 1: 22). [SHIKOKU] 3 ♀(Rh 0053–0055), emerged 1–8.IV. 2011 from larvae collected by MK on 27. II. 2011 at Yasui-keikoku, Kouchi Pref (Fig. 1: 11). [KYUSHU] 2 ♀ (Rh 0 0 56, 0057), emerged 20–25.IV. 2010 from larvae collected by MK on 12.IV. 2010 at Gokanosho, Kumamoto Pref (Fig. 1: 10). 4 ♀(Rh 0058–0061), emerged 18–22.IV. 2010 from larvae collected by MK on 11.IV. 2010 at Mt. Kosho, Fukuoka Pref (Fig. 1: 8). Etymology. The specific epithet is a noun in apposition, derived from the distribution of this species. Distribution. Japan (Hokkaido, Honshu, Shikoku, Kyushu) (Fig. 1). Natural history. Adults of this species emerged from thalli of the Conocephalum conicum species complex (Marchantiales: Conocephalaceae) growing on moist rocky or clay slopes along streams in both evergreen and deciduous forests (Fig. 11 a). Larvae of this species are thallus miners of C. conicum species complex (Fig. 11 d). Adults emerged in spring (March–May, in laboratory conditions). Diagnosis. Litolepis japonica is easily separated from all other congeners by having a long stout setae at the tip of first flagellomere. This species can also be easily distinguished from the other species by the form of apical extension of gonostylus that is unevenly bifurcated in males, and spermathecal duct accessory glands arising at base of spermatheca in females. Remarks. It is noteworthy that Litoleptis japonica has extremely female-biased sex ratios: only seven males of 61 adult flies of L. japonica were obtained, even though the collecting method and timing (collecting as larvae mining in the liverwort mats) did not appear to produce the sampling biases.

Litoleptis niyodoensis Imada & Kato sp. n. [Japanese name: niyodo-jingasa-shitone-abu] (Figs. 3 e, 3 f, 4 c, 7) Description. Body length: 2.9 mm (n= 7) in male, 3.1 mm (n= 3) in female. Wing length: 2.7 mm (n= 7) in male, 3.0 mm (n= 3) in female. Head Frons bare in male, with sparse setae in female. Antenna 0.47 mm (n= 3). Antenna 1.1 times as long as diameter of eye, densely covered with short setae consistently. First flagellomere elongate as long tapering segment or oval and enlarged near base, concave ventrally at basal one-third to half; distal end not enlarged (Fig. 7 a). Male (Fig. 7 b) Gonostylus with flat, oblong-shaped extension apically, bluntly pointed inwardly at apex; undulating in posterior edge of basal part. Sperm sac bulbous hardly visible. Lateral ejaculatory process integrated into sperm sac, elongate transversely accompanying with distal end of sperm sac. Ejaculatory apodeme short, round at anterior margin of gonocoxite. Posteromedial margin in ventral surface of gonocoxite bear 5���7 bristles. Aedeagus tapering toward apex, connected with gonocoxite at paramere at thick and wide joint. Paramere swollen dorsally near distal end, widest at connection with aedeagus, approximately three times as wide as distal end. Female (Fig. 7 c) Spermatheca membranous, rounded at end. Spermathecal duct accessory gland prominent, with short and sclerotized columnar duct, arising at approximately two-thirds the distal length between genital chamber and spermatheca, with saccate structure visible. Common spermathecal duct absent. Genital chamber small. Type material. Holotype. JAPAN [HONSHU] 1 ♂, emerged 29.III. 2011 from larva collected by MK on 27. II. 2011 at Yasuikeikoku, Kouchi Pref (Fig. 1: 11), ���Rh 0184���, NMNS. Paratypes. 7 ♂, 3 ♀ (Rh 0185���0194) emerged during 27���29.III. 2011 from larvae collected by MK on 27. II. 2011, same locality as holotype, KUHE. Etymology. The specific epithet is a noun in apposition, taken from Niyodo river, the main stream of Yasui river flowing near the type locality. Distribution. Japan (Honshu: Kouchi Prefecture) (Fig. 1). Natural history. Habitat of this species is similar to that of L. kiiensis. Larvae of this species are miners of thallus of Reboulia hemisphaerica. Adults emerged in spring (March, in laboratory condition). Diagnosis. Litoleptis niyodoensis resembles L. kiiensis and himukaensis, and be distinguished from them by the following characters: first flagellomere tapering toward apex with a slight depression at basal one-third to half without enlarged apex; gonostylus with oblong-shaped extension, undulating in posterior edge of basal part; posteromedial margin in ventral surface of gonocoxite bear 5���7 bristles; paramere swollen posteriorly near distal end (without swollen in other two species); spermatheca rounded at end in female (as opposed to conical-shaped in L. kiiensis)., Published as part of Imada, Yume & Kato, Makoto, 2016, Bryophyte-feeding of Litoleptis (Diptera: Rhagionidae) with descriptions of new species from Japan, pp. 41-58 in Zootaxa 4097 (1) on pages 49-50, DOI: 10.11646/zootaxa.4097.1.2, http://zenodo.org/record/271005, {"references":["Friis, E. M., Crane, P. R. & Pedersen, K. R. (2011) Early flowers and angiosperm evolution. Cambridge University Press, Cambridge, 596 pp."]}

Litoleptis Chillcott Type species: Litoleptis alaskensis Chillcott 1963: 1187, by original designation. L. chilensis Hennig 1972: 6. L. orientalis as Hilarimorpha; Frey 1954: 25. L. fossilis Arillo et al. 2009. Description. Head (Fig. 2 b, c) dark brown, glossy. Vertex, occiput, postgena with sparse setae, gena bare. Eyes inconspicuously setulose; slightly dichoptic or holoptic in male, dichoptic in female; in male, upper part orange, lower part black. Mandibles absent. Antenna black or pale grey. Pedicel clearly larger than scape. Scape minute, pedicel as broad as long and weakly bristled around distal edge. Clypeus slightly bulbous. Palpus one-segmented, with black setae. Labellum developed, without pseudotracheae. Thorax (Fig. 2 a) brownish black with pruinosity and with three dark brown vittae. Postpronotal lobe, proepimeron, bare. Scutellum with sparse setae on posterior margin. Mesonotum without vittae. Proscutellum absent. Subscutellum inconspicuous. Proepimeron developed. Postscutellum bare. Anatergite and katatergite indistinguishable. Wing (Fig. 4) infuscate, without any dark markings, despite having intraspecific variation in color darkness. Pterostigma inconspicuous. Lower calypter reduced. Upper calypter triangular in form, underdeveloped; with broad curvature. Costa ending before wing tip. Humeral crossvein weakly developed. Crossvein sc-r present, positioned proximal of humeral crossvein. R 1 dorsal and ventral surface bare. R 2 + 3 sinuous, apical third of R 2 + 3 ultimately bends anteriorly slightly, toward leading edge of wing margin; longer than R 5, but less than twice as long. Fork of R 4 + 5 distal of distal end of fork of M 1 + 2. R 4 at base mostly straight, ending at wing tip entire length. R 5 ending at wing tip. M 3 absent. Medial crossvein absent. CuA 2 generally join with A 1. CuA 2 +A 1 reaching or sometimes not reaching wing margin. Anal cell closed usually before and sometimes at wing margin. Alula reduced with narrow curvature; rounded evenly. Legs dark brown to black; coxae and femora more or less pollinose with weak setae. Empodium pulvilliform. Tibial spur formula 0:0:0. Macrotracheae absent. Postmetacoxal bridge absent. Hind coxae bare behind. Abdomen dark brown to black with pruinosity. Male Cercus displaced away from epandrium, widely displaced from one another, separation distance about 3 / 4 width of cercus; held vertically over abdomen; flat dorso-ventrally. Tergite 10 absent. Hypandrium fused entirely to gonocoxites. Gonocoxal ridge absent. Gonocoxal apodeme absent. Gonostylus with developed extension apically from apex of main component. Aedeagus slender and conical-shaped; apical surface covered with tiny pustules; connected with paramere at base. Lateral ejaculatory process either present or absent. Aedeagal tines absent. Endoaedeagal process absent. Sperm sac bulbous, generally without distinct lobes, envelope aedeagus ventrally. Female (for L. japonica, kiiensis, niyodoensis, asterellaphile; Fig. 2 d) Tergite 7 wider than long. Tergite 9 bare, inconspicuous and largely retracted within tergite 8; having ventrolateral projections, extending posteriorly, surrounding and fusing sternite 9 laterally. Intersegmental membrane clearly separating tergites 7 and 8. Spermathecae three, membranous. Spermathecal duct short, less than length of sternite 9. Spermathecal duct accessory glands with short, sclerotized columnar duct. Circular ridge of ejection apparatus absent. Ring at base of spermathecal ducts lightly or not sclerotized. Spermathecal duct ejection apparatus absent. Common spermathecal duct absent or inconspicuous. Surrounding area of genital chamber membranous, contained sclerotization of sternite 9 laterally. Diagnosis. Litoleptis can immediately be distinguished from the other genera of Rhagionidae. Major autapomorphies are as follows: medial crossvein (and discal cell) absent; all tibiae without spurs; gonocoxal apodeme absent. Litoleptis specifically mostly resembles Spania in general appearance and stylus-shaped antenna, although they can easily be distinguished by the following character states besides the above-mentioned traits; male gonostylus with an extension from apex of main component expanding into an irregular form (as opposed to constantly slender or club-like form in Spania); gonocoxal apodeme absent. Also, the Japanese Litoleptis shows inconsistency in having the following traits unlike other congeners (e.g. L. alaskensis): male eyes dichoptic; lateral ejaculatory process present (except L. japonica); ejaculatory apodeme present. The following traits are uniquely found in the Japanese Litoleptis: apex of aedeagus covered with numerous tiny pustules; spermathecal duct accessory glands connected by short, sclerotized columnar duct arising from spermathecal duct., Published as part of Imada, Yume & Kato, Makoto, 2016, Bryophyte-feeding of Litoleptis (Diptera: Rhagionidae) with descriptions of new species from Japan, pp. 41-58 in Zootaxa 4097 (1) on pages 43-44, DOI: 10.11646/zootaxa.4097.1.2, http://zenodo.org/record/271005, {"references":["Chillcott, J. G. (1963) A new genus of Rhagionidae (Diptera) with notes and descriptions of Bolbomyia Loew. The Canadian Entomologist, 95, 1185 - 1190. http: // dx. doi. org / 10.4039 / Ent 951185 - 11","Hennig, W. (1972) Eine neue Art der Rhagionidengattung Litoleptis aus Chile, mit Bemerkungen uber Fuhlerbildung und Verwandtschaftsbexiebungen einiger Brachycerenfamilien (Diptera: Brachycera). Stuttgarter Beitraege zur Naturkunde. Serie A. Biologie, 174, 1 - 51.","Arillo, A., Penalver, E. & Garcia-Gimeno, V. (2009) First fossil Litoleptis (Diptera: Spaniidae) from the Lower Cretaceous amber of San Just (Teruel Province, Spain). Zootaxa, 2026, 33 - 39."]}

Litoleptis himukaensis Imada & Kato sp. n. [Japanese name: himuka-shitone-abu] (Figs. 3 g, 3 h, 4 d, 8) Description. Body length: 2.8 mm (n= 3) in male, 2.9 mm (n= 2) in female. Wing length: 2.4 mm (n= 4) in male, 2.6 mm (n= 2) in female. Head Frons bare in both sexes. Antenna 0.53 mm (n= 3). Antenna 1.0 times as long as diameter of eye. First flagellomere straight and gradually tapering towards apex; covered with comparatively rough and long setae (Fig. 8 a). Male (Fig. 8 b) Gonostylus with oblong-shaped extension apically; posterior surface of apex hollowed and weakly pointed inwardly; undulating in posterior edge of basal part. Sperm sac bulbous and hardly visible. Lateral ejaculatory process apparently not integrated into sperm sac, freely elongate transversely. Ejaculatory apodeme moderately long but not reaching anterior margin of gonocoxite. Posteromedial margin in ventral surface of gonocoxite bear less than four bristles. Aedeagus smoothly connected with paramere at thick and wide joint. Paramere slightly swollen posteriorly near distal end; become widest at connection with aedeagus, less than two times as wide as distal end. Type material. Holotype. JAPAN [KYUSHU] 1 ♂, emerged 14. II 2009 from larvae collected by MK on 17.I. 2009 at Udo Jingū, Miyazaki Pref, Japan (Fig. 1: 9), ���Rh 0195���, NMNS. Paratypes. 3 ♂, 3 ♀ (Rh 0196���0201), emerged 10���17.II. 2009 from larvae collected by MK on 17.I. 2009 at same locality as holotype (Fig. 1: 9), KUHE. Etymology. The specific epithet is taken from classical name of the type locality, ���Himuka���, southern part of Miyazaki Prefecture. Distribution. Japan (Honshu: Miyazaki Prefecture) (Fig. 1). Natural history. Habitat of this species is rocky slope along streams in coastal evergreen Castanopsis forests. Larvae of this species are thallus miner of Reboulia hemisphaerica (Fig. 11 g). Adults emerged in spring (March- July, in laboratory condition). Diagnosis. Litoleptis himukaensis resembles L. kiiensis and L. niyodoensis, and can be distinguished from them by the gonostylus with oblong-shaped extension apically (as opposed to L. kiiensis), of which posterior surface of apex is hollowed (as opposed to flat at apex in L. niyodoensis)., Published as part of Imada, Yume & Kato, Makoto, 2016, Bryophyte-feeding of Litoleptis (Diptera: Rhagionidae) with descriptions of new species from Japan, pp. 41-58 in Zootaxa 4097 (1) on page 50, DOI: 10.11646/zootaxa.4097.1.2, http://zenodo.org/record/271005