The F. jezoensis-subgroup was originally established

Ann. soc. entomol. Fr. (n.s.), 2007, 43 (3) : 357-361 ARTICLE Review of the Fannia jezoensis-group (Diptera: Fanniidae) Mingfu Wang, Dong Zhang & Rongrong Wang Institute of Entomology, Shenyang Normal University, Shenyang 110034, China Abstract. The jezoensis species-group is established within the genus Fannia Robineau-Desvoidy, consisting of four species, Fannia curvostylata n. sp., Fannia densa Nishida, Fannia jezoensis Nishida and Fannia zhangi Xue. The new species described here is from the Tibetan Plateau. The known species of this group are all described, illustrated and keyed. Geographic distributions are updated. The systematic position and biogeography of the F. jezoensis-group are also discussed. Résumé. Revue du groupe de Fannia jezoensis (Diptera : Fanniidae). Le groupe d espèce de jezoensis est établi au sein du genre Fannia Robineau-Desvoidy. Ce groupe comprend quatre espèces Fannia curvostylata n. sp., Fannia densa Nishida, Fannia jezoensthe Nishida et Fannia zhangi Xue. La nouvelle espèce décrite ici provient du plateau Tibétain. Les espèces du groupe sont redécrites, illustrées et incluses dans une clef. Leur distribution géographique est mise à jour. La position systématique et la biogéographie du groupe de Fannia jezoensis sont discutées. Keywords: Diptera, Fanniidae, Fannia, F. jezoensis-group, new species. The F. jezoensis-subgroup was originally established in the genus Fannia Robineau-Desvoidy by Nishida (1994), and included two species from Japan and Nepal respectively (Nishida 1976, 1994). While studying specimens of the genus Fannia from the Tibetan Plateau, we found one further undescribed species, closely related to F. jezoensis Nishida, 1976. After a systematic study of related species, the jezoensisgroup is expanded to include two additional species: F. curvostylata n. sp. and F. zhangi Xue, 1996. This has given us an opportunity to provide a definition of this group. As the morphological characters and especially the male genitalic structures of this group are extraordinarily different from other groups in the genus Fannia, we are confident in giving full speciesgroup status to these four species. The primary aims of this article are to review the F. jezoensis-group, to resolve any lingering systematic ambiguity associated with it, to describe one new species, and to provide an identification key to the known species. New data on localities are reported. The systematic position and biogeography of this group are also discussed. Material and methods The specimens examined in the course of this study were collected by sweeping from brushwood in mountainous E-mail: wangmingfu403@163.com Accepté le 28 juin 2007 regions. Specimens were dried and studied, and are deposited in the Institute of Entomology, Shenyang Normal University, Shenyang, Liaoning Province, China. The external morphology was observed under a stereoscopic microscope and metric characters were measured with an ocular micrometer. To observe the detailed characters of the male terminalia, these organs were detached from the body, cleared by warming in a 10% KOH solution (approximately 100 C) for several minutes, placed in a droplet of glycerol, and observed under a compound light microscope. Terminology and measurements The morphological terminology follows McAlpine (1981). Absolute measurements are used for body length in millimetres (mm). Abbreviations used for characters include: a = anterior seta, acr = acrostichal seta, ad = anterodorsal seta, av = anteroventral seta, d = dorsal seta, dc = dorsocentral seta, ia = intra-alar seta, p = posterior seta, pd = posterodorsal seta, pra = prealar seta, and pv = posteroventral seta. Systematic account Fannia jezoensis-group Fannia jezoensis-subgroup Nishida 1994: 93. Diagnosis. Eye with dense long hairs; tibiae usually with more than 1 seta on each surface (including av, ad, pd or pv surface), mid first tarsomere without basal tooth-like spine on ventral surface, hind coxa bare on posterior surface (rarely with 1 or 2 hairs), hind femur with stout pv in distal part; lower calypter slightly projecting beyond upper one; pra 2 4; apex of male tergite 9 with 1 pair of long ventral lobes; male cerci with numerous distal teeth on ventral surface; male bacilliform process absent. Distribution. China, Japan and Nepal. 357

M. Wang, D. Zhang & R. Wang Key to males of the known species of the Fannia jezoensis-group 1. Calypters yellow... 2 Calypters blackish, frontal setae 12 15, hind tibia with 1 av... Fannia jezoensis Nishida 2. Hind tibia with 2 or 3 av, 5 or 6 ad and numerous short p on median part, surstyli straight in dorsal view, frontal setae 8 13... 3 Hind tibia with 2 av and 2 ad, without setae on posterior surface, surstyli sinuous in basal 3/5 in dorsal view, frontal setae 8 or 9... Fannia curvostylata n. sp. 3. Parafacial at middle about 2/3 as wide as width of first flagellomere, frontal setae 10 13, mid tibia with 3 ad, hind tibia with 10 13 fine p, haltere yellow, cercal plate with a median apical tooth in dorsal view... Fannia densa Nishida Parafacial at middle about as wide as width of first flagellomere, frontal setae 8, mid tibia with 2 ad, hind tibia with 5 6 fine p, haltere yellowish-brown, cercal plate without apicaltooth in dorsal view... Fannia zhangi Xue Fannia curvostylata n. sp. (Figs 1 3) Types. Holotype, China, Yunnan: Deqen County: Mt. Baima [28 12 N 99 13 E] (alt. 3800 4200 m), 5.VII.2006, leg. M.F. Wang. Paratypes 14, same data as holotype. Description. Male. Body length 5.5 mm. Eye with long brown hairs, the longest hair about 2/3 as long as width of antennal first flagellomere; fronto-orbital plate and parafacial with silverygrey pruinosity, frons about 2/3 as wide as the distance between two posterior ocelli, frontal vitta as wide as fronto-orbital plate, frontal setae 8 or 9, reaching ocellar triangle, the gaps filled with 5 or 6 fine setae, upper orbital setae absent; parafacial bare, at middle about 2/3 as wide as width of first flagellomere; antenna black, first flagellomere 2.0 times as long as wide, arista pubescent, the longest hair about as long as width of aristal base; epistoma not projecting beyond vibrissal angle, vibrissal angle behind frontal angle in profile; prementum with thin greyish pruinosity, its length 2.0 times as long as its width; palpus black, claviform, slightly longer than prementum. Thorax groundcolour black, notum with brownish-grey pruinosity, without distinct vitta; presutural acr biserial, hair-like, only presutural pairs slightly stout, the distance between the two rows about 1/2 of the distance between acr and dc rows, dc 2+3, ia 0+2, pra 2, the anterior one about 2/3 the length of posterior notopleural seta, notopleuron without setulae; basisternum, proepisternum, 1:1, katepisternum without ventral spines; spiracles brown; calypters yellow, the lower one slightly projecting beyond the upper one. Wing brownish; tegula brown, basicosta yellow, costal spine inconspicuous; node of Rs bare on ventral and dorsal surfaces; vein M straight; crossveins without obvious cloud; haltere yellow. Legs black; fore tibia without median p; mid coxa without any hooked spines or spine-like setae on lower and outer margins; mid femur with a long and sparse av row, becoming gradually denser and shorter towards apex, ad row indistinct, pv row fine and long, becoming biserial or triserial; mid tibia with 2 ad and 2 pd; mid first tarsomere without basal tooth-like spines on ventral surface; hind coxa bare on posterior surface; hind femur with a complete av row, distal 3 or 4 setae stout, pv row stout; hind tibia with 2 av, 2 ad and 1 median d. Abdomen long and flattened, ground-colour black, with dense greyish-brown pollinosity; syntergite 1+2 with a broad median vitta, tergites 3 and 4 each with a median triangular mark, tergite 5 with a median vitta, each tergite with long marginal setae; sternite 1 with hairs, sternite 5 with sparse setae, apex indented, its width in basal 1/4 about 0.65 times as wide as its width at middle; cercal plate short and broad, its length about 1.5 times as long as its width in dorsal view; surstyli sinuous in basal 3/5 in dorsal view, distal 1/3 subulate in lateral view. Etymology. This specific name is from the Latin and refers to the shape of the male surstyli. Remarks. This new species resembles F. jezoensis but differs from it in having frontal setae 8 or 9; calypters and haltere yellow; hind tibia with 2 av; sternite 5 broader in basal part and with sparse hairs; length of cercal plate about 1.5 times as long as its width in dorsal view; surstyli sinuous in basal 3/5 in dorsal view, distal 1/3 subulate in lateral view. Distribution. China (Yunnan). Fannia densa Nishida 1994 (Figs 4 6) Fannia densa Nishida 1994: 89-92. Specimens examined. 2, China, Yunnan: Deqen County: Mt. Baima [28 12 N 99 13 E] (alt. 3800 4200 m), 5.VII.2006, leg. M.F. Wang. Figures 1 3 Fannia curvostylata n. sp. (male holotype). 1, terminalia, dorsal view; 2, terminalia, lateral view; 3, sternite 5. Scale 0.25 mm. Figures 4 6 Fannia densa Nishida (male). 4, terminalia, dorsal view; 5, terminalia, lateral view; 6, sternite 5. Scale 0.25 mm. 358

Review of Fannia jezoensis-group Description. Male. Body length 5.5 mm. Eye with long brown hairs, the longest hair about 3/4 as long as width of antennal first flagellomere; fronto-orbital plate and parafacial with silvery-grey pruinosity, frons about 2/3 as wide as the distance between two posterior ocelli, frontal vitta as wide as frontoorbital plate, frontal setae 10-13, reaching ocellar triangle, the gaps filled with 3 or 4 fine setae, upper orbital setae absent; parafacial bare, at middle about 2/3 as wide as width of first flagellomere; antenna black, first flagellomere 2.0 times as long as wide, arista pubescent, the longest hair about as long as basal aristal width; epistoma not projecting beyond vibrissal angle, vibrissal angle behind frontal angle in profile; prementum with thin greyish pruinosity, its length 2.5 times as long as its width; palpus black, claviform, slightly longer than prementum. Thorax ground-colour black, notum with brownish-grey pruinosity, without distinct vitta; presutural acr biserial, hair-like, only presutural pairs slightly stout, the distance between acr rows about 1/2 of the distance between acr and dc rows, dc 2+3, ia 0+2, pra 3 or 4, the anterior one about 2/3 as long as posterior notopleural seta, notopleuron without setulae; basisternum, proepisternum, anepimeron, meron and katepimeron bare; katepisternal setae 1:1, katepisternum without ventral spines; spiracles brown; calypters yellow, the lower one slightly projecting beyond upper one. Wing brownish; tegula brown, basicosta yellow, costal spine inconspicuous; node of Rs bare on ventral and dorsal surfaces; vein M straight; crossveins without obvious cloud; haltere yellow. Legs black; fore tibia without setae on lower and outer margins; mid femur with av row, stout and long in basal half, becoming gradually denser and shorter towards apex, ad row indistinct, pv and p rows stout, the longest seta 1.5 times as long as femoral width; mid tibia slightly swollen in distal half, with numerous ventral hairs, the longest hair slightly shorter than tibial width, with 3 ad and 2 pd; mid first tarsomere without basal tooth-like spines on ventral surface; hind coxa bare on posterior surface; hind femur with an incomplete av row, only distal 3 or 4 setae stout, 3 or 4 stout a and 4 or 5 ad in distal half, pv row stout; hind tibia with 2 av, 6 ad, 1 median d and 10-13 short p around middle. Abdomen long and flattened, ground-colour black, with dense greyish-brown pollinosity; tergites 2 to 4 each with a median triangular mark, tergite 5 with a median vitta, each tergite with long marginal setae; sternite 1 with hairs, sternite 5 distinctly broadened apically, apex indented, the setae scattered evenly over the surface; cercal plate rounded, pointed apically, with a median setulae dense on dorsal surface; surstyli straight in dorsal view, distal 1/3 subulate in lateral view. Distribution. China (Yunnan), Nepal (Thudam). Fannia jezoensis Nishida 1976 (Figs 7 9) Fannia jezoensis Nishida 1976: 137-138. Specimens examined. 1, China, Yunnan: Deqen County: Mt. Baima [28 12 N 99 13 E] (alt. 3800 4200 m), 5.VII.2006, leg. M.F. Wang. Description. Male. Body length 6.0 mm. Eye with long brown hairs, the longest hair about as long as frontal width; frontoorbital plate and parafacial with whitish-grey pruinosity, frons 1.5 times as wide as width of anterior ocellus, about 2/3 as wide as the distance between two posterior ocelli, frontal vitta Figures 7 9 Fannia jezoensis Nishida (male). 7, terminalia, dorsal view; 8, terminalia, lateral view; 9, sternite 5. Scale 0.25 mm. linear, frontal setae 12 or 13, nearly reaching ocellar triangle, the gaps filled with 5 or 6 fine setae, upper orbital setae absent; parafacial bare, at middle about 2/3-3/4 as wide as width of first flagellomere; antenna black, first flagellomere 2.0 times as long as wide, arista pubescent, the longest hair about as long as basal aristal width; epistoma not projecting beyond vibrissal angle, vibrissal angle behind frontal angle in profile; prementum with thin greyish pruinosity, its length 2.0 times as long as its width; palpus black, about as long as prementum. Thorax groundcolour black, notum with brown pruinosity, without distinct vitta; presutural acr biserial, stout, dc 2+3, ia 0+2, pra 3, the longest one about 2/3 as long as posterior notopleural seta, notopleuron without setulae; basisternum, proepisternum, 1:1, katepisternum without ventral spines; spiracles brown; calypters blackish, the lower one slightly projecting beyond upper one. Wing brownish; tegula black, basicosta brown, costal spine inconspicuous; node of Rs bare on ventral and dorsal surfaces; vein M straight; crossveins without obvious cloud; haltere brown. Legs entirely black; fore tibia without setae on lower and outer margins; mid femur with av row stout and long in basal half, becoming gradually denser and shorter towards apex, a row complete and stout, the longest seta as long as femoral width, pv and p rows stout, the longest seta 1.5 times as long as femoral width; mid tibia slightly swollen in distal half, with numerous ventral hairs, the longest hair about 4/5 of tibial width, with 2 ad and 2 pd; mid first tarsomere without basal tooth-like spines on ventral surface; hind coxa bare on posterior surface, other parts lost. Abdomen long and flattened, ground-colour black, with dense greyish-brown pollinosity; Figures 10 12 Fannia zhangi Xue (male holotype). 10, terminalia, dorsal view; 11, terminalia, lateral view; 12, sternite 5. Scale 0.25 mm. 359

M. Wang, D. Zhang & R. Wang tergites 2 to 4 each with a median triangular mark; sternite 1 with hairs, sternite 5 with dense setae, apex indented, its width in basal 1/4 about 0.8 as wide as its width at middle; cercal plate short and broad, its length as long as its width in dorsal view; surstyli straight in dorsal view. Distribution. China (Yunnan), Japan (Hokkaido). Fannia zhangi Xue 1996 (Figs 10 12) Fannia zhangi Xue in Xue & Wang 1996: 830. Type examined. Holotype, CHINA, Tibet: Zogang County [29 41 N 97 54 E] (alt. 3800 m), 30.VI.1976, leg. X.Z. Zhang. Paratype 1, same data as holotype. Description. Male. Body length 4.8 mm. Eye with long brown hairs, the longest hair about as long as width of antennal first flagellomere; fronto-orbital plate and parafacial with whitishgrey pruinosity, frons about as wide as width of antennal first flagellomere, frontal vitta linear in upper half, frontal setae 8, reaching ocellar triangle, the gaps filled with 3 or 4 fine setae, upper orbital setae absent; parafacial bare, at middle about as wide width of first flagellomere; antenna black, first flagellomere 2.5 times as long as wide, arista pubescent, the longest hair about as long as basal aristal width; epistoma not projecting beyond vibrissal angle, vibrissal angle behind frontal angle in profile; prementum with thin pruinosity, its length 2.5 times as long as its width; palpus black, about as long as prementum. Thorax ground-colour black, notum with thin brownish-grey pruinosity, without distinct vitta; presutural acr biserial, slightly stout, the distance between acr rows slightly narrower than the distance between acr and dc rows, dc 2+3, ia 0+2, pra 2 or 3, the anterior one about 2/3 of the length of anterior notopleural seta, notopleuron without setulae; basisternum, proepisternum, 1:1, katepisternum without ventral spines; spiracles yellow; calypters yellow, lower one slightly projecting beyond upper one. Wing brownish; tegula brown, basicosta brownish, costal spine inconspicuous; node of Rs bare on ventral and dorsal surfaces; vein M straight; crossveins without obvious cloud; haltere yellowish-brown. Legs entirely black; fore tibia without Figure 13 Distribution of Fannia jezoensis-group. setae on lower and outer margins; mid femur with complete av and pv rows; mid tibia with 2 ad and 2 pd; mid first tarsomere without basal tooth-like spines on ventral surface; hind coxa bare on posterior surface; hind femur only with 3 or 4 distal av, pv row complete; hind tibia with 2 or 3 av, 5 or 6 ad, 1 median d and 5 or 6 short p in middle part. Abdomen long and flattened, ground-colour black, with greyish-brown pollinosity; tergites 2 to 4 each with a median triangular mark, tergite 5 with a median vitta; sternite 1 with hairs, sternite 5 gradually broadened apically, apex indented, the setae scattered evenly over the surface; cercal plate rounded, pointed at apex, without distal dense setulae on dorsal surface; surstyli straight in dorsal view, distal 1/3 subulate in lateral view. Distribution. China (Tibet). Discussion Systematic considerations Nishida (1976) recognized that the morphological characters and especially the male genitalic structures of F. jezoensis are extraordinarily different from other species in the genus Fannia. Almost twenty years later, he (1994) described F. densa from Nepal as a new species, which is closely related to F. jezoensis. Based on Chillcott (1961), he (1994) assigned these two species to the Fannia carbonaria-group and erected a new subgroup for them, the F. jezoensissubgroup. Amongst fanniid speciemens collected by the first author, as part of an ongoing survey of flies in the south-eastern Tibetan Plateau, we found one undescribed species (F. curvostylata n. sp.) and two species (F. jezoensis and F. densa) that are new records for this region. When we consulted the descriptions and figures by Xue & Wang (1996), the morphological characters and especially the male genitalic structures of F. zhangi showed that this species can also be assigned to this group. The monographs of Palaearctic, Nearctic, Australian and European species, with comprehensive descriptions and keys by Hennig (1955), Chillcott (1961), Pont (1977) and Rozkošný et al. (1997), are very useful for taxonomic studies of the Fanniidae. They provide a sound basis for a comprehensive study of this family at the species-group level. A systematic study of these four closely related species showed that both the external pattern elements and the genitalia of this group differ substantially from those of the currently-recognised species-groups in Fannia. The group can be distinguished from other Fannia groups by the male cercal plate with numerous distal teeth on the ventral surface and the apex of male tergite 9 with a pair of long ventral lobes. A number of other 360

Review of Fannia jezoensis-group reliable characters, such as the eye with dense long hairs, pra 2-4 and male bacilliform process absent, are also used as species-group diagnostic characters. So we can confirm that the F. jezoensis-group is an independent species-group in Fannia. Biogeographic considerations The distribution pattern of this group is worthy of special attention. The four known species have been found mainly in the Tibetan Plateau, and three of them are endemic and occur sympatrically on Mt. Baima, generally restricted to altitudes between approximately 3800 and 4700 m. Only F. jezoensis has a wider range which also includes Hokkaido, Japan (Fig. 13). Fanniidae most probable existed in the Cretaceous (Hennig 1965), and the great diversity of species in the Palaearctic region indicates that this area may well have been the point of origin of the family (Chillcott 1961). Judging by the distribution pattern of this group, its precursor was probably first introduced to the Tibetan Plateau by the way of Eurasia during the Miocene, after the Indian Plate collided with the Eurasian Plate. With the uplifting of the Tibetan Plateau, the complex topography resulted in a wide range of climatic conditions and vegetation types which in turn gave rise to diverse niches for insect speciation. This region thus became the centre of origin for this group. Most Japanese islands were once connected to the Asian mainland during the Late Pleistocene (Lawlor 1986; Virginie & Jaeger 1999), offering some direct pathways for the migration of species from the mainland into these new territories. The cold and warm cycles of the glaciations during the Pleistocene may have led the ranges of some jezoensis-group populations to expand and contract. F. jezoensis may have been introduced into Japan by then. It also resulted in the range of this group being disjunct between the Tibetan Plateau and Japan. Acknowledgments. Sincere thanks to Dr Adrian C. Pont (Oxford University Museum of Natural History, Oxford, U.K.), Dr Rudolf Rozkošný (Faculty of Science, Masaryk University, Brno, Czech Republic), and Dr Kazumi Nishida (Kobe Institute of Health, Kobe, Japan) for giving us invaluable help over many years. We are grateful to Prof. Wanqi Xue, Prof. Chuntian Zhang and Ms Lin Liu (Institute of Entomology, Shenyang Normal University), who have given us invaluable help with this study. This study was supported by the Director Foundation of the Experimental Centre of Shenyang Normal University (No.SY200404) and the Scientific Research for College Foundation of the Educational Department of Liaoning Province (No.2005386). References Chillcott J. G. 1961. A revision of the Nearctic species of Fanniinae (Diptera: Muscidae). The Canadian Entomologist Supplement 14 (1960): 1-295. Hennig W. 1955. Muscidae [part], p. 1-99 in: Lindner, E. (ed.). Die Fliegen der palaearktischen Region 63b. Schweizerbart, Stuttgart. Hennig W. 1965. Vorarbeiten zu einem phylogenetischen System der Muscidae (Diptera: Cyclorrhapha). Stuttgarter Beiträge zur Naturkunde 141: 1-100. Lawlor T. E. 1986. Comparative biogeography of mammals on islands. Biological Journal of the Linnean Society 28: 99-105. McAlpine J. F. 1981. Morphology and terminology - adults, p. 9-63 in: McAlpine J. F., Peterson B. V., Shewell G. E., Teskey H. J., Vockwroth J. R., Wood D. M. (coords.). Manual of Nearctic Diptera. Volume 1. Research Branch, Agriculture Canada Monograph 27. Canadian Government Publishing Center, Hull. Nishida K. 1976. Studies on the species of Fanniinae (Diptera: Muscidae) from Japan IV. Five new and two newly recorded species of genus Fannia from Japan. Japanese Journal of Sanitary Zoology 27: 133-143. Nishida K. 1994. The Fanniidae from Nepal (Diptera). Japanese Journal of Sanitary Zoology 45 (Supplement): 81-97. Pont A.C. 1977. A revision of Australian Fanniidae (Diptera: Calyptrata). Australian Journal of Zoology, Supplement Series 51: 1-60. Rozkošný R., Gregor F., Pont A. C. 1997. The European Fanniidae (Diptera). Acta Scientiarum Naturalium Academiae Scientiarum Bohemicae Brno (n.s.) 31(2): 1-80. Virginie M. P., Jaeger J. J. 1999. Island biogeography of the Japanese terrestrial mammal assemblages: an example of a relict fauna. Journal of Biogeography 26: 959-972. Xue W. Q., Wang M. F. 1996. Family Fanniidae [In Chinese.], p. 809-835 in: Xue W. Q., Chao C. M. (eds.). Flies of China, Volume I. Liaoning Science and Technology Press, Shenyang. 361