The misplaced genus Trischidocera Villeneuve (Diptera ... - Biotaxa

Zootaxa 3926 (2): 279–286 www.mapress.com /zootaxa / Copyright © 2015 Magnolia Press

Article

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3926.2.8 http://zoobank.org/urn:lsid:zoobank.org:pub:B3F0E470-5984-4450-B363-1EE99B080CA8

The misplaced genus Trischidocera Villeneuve (Diptera, Tachinidae) SILVIO S. NIHEI Department of Zoology, Institute of Biosciences, University of São Paulo, Rua do Matão, Travessa 14, n.101, São Paulo, SP, 05508090, Brasil. E-mail: [email protected]

Abstract Trischidocera Villeneuve, 1915 includes two species, T. sauteri Villeneuve, 1915 (Taiwan and Malaysia) and T. yunnanensis Chao & Zhou, 1987 (China). The systematic placement of Trischidocera has been controversial. It was originally placed within the “Thryptoceratidae” (= “Actiidae”), then moved to Germariini, then considered an unplaced Tachinidae, and more recently placed in Ormiini. Here, the genus is revised, the type-species is redescribed and illustrated, and its systematic placement is discussed. The genus is removed from Ormiini and considered incertae sedis. Key words: Germariini, Ormiini, revision, Siphonini, systematics, taxonomy

Introduction Trischidocera was erected by Villeneuve (1915) for the new species T. sauteri based on three males from Taiwan. Villeneuve originally placed the new genus in “Thryptoceratidae”, a family-group name proposed by RobineauDesvoidy (1851) for a group of genera, which would be equivalent to the “Actiidae” of Bigot (1882). Townsend (1932) provided some notes on T. sauteri and placed the genus in “Actiini”. Thereafter, Townsend (1936, 1939) transferred Trischidocera to Germariini with the following note (1939: 358): “While this form exhibits strong actiine characters, the exposed sternites, bare prosternum and undeveloped IPAL [intrapostalar seta] combine to bring it here in the keys.” Townsend (1939) also synonymized Orectocerina Malloch, 1924 with Trischidocera. Crosskey (1976) listed Trischidocera in the final section of his Oriental conspectus as an unplaced Tachinidae, and keyed it in the key to tribes but put it apart from any tribe. Crosskey (1976) explicitly doubted the synonymy of Orectocerina atratula Malloch (Malaysia) with T. sauteri (Taiwan) proposed by Townsend (1939). He wrote “It appears possible that distinct species are involved.” But his doubt was probably due to the strong dimorphism between the male holotype of T. sauteri and the female holotype of O. atratula. Chao & Zhou (1987) described a second species, T. yunnanensis, from China (Yunnan). While the male postpedicel of T. sauteri is trilobed and notably elongate, in T. yunnanensis it is simple (not-trilobed), although the distally inserted arista is shared by both species. The terminal arista in T. yunnanensis results in an aberrant form resembling a stylate antenna. In Flies of China, Chao et al. (1998: 1957) curiously included Trischidocera within Ormiini. They did not provide any argument supporting this new placement. More recently, the Chinese catalogue by O’Hara et al. (2009: 161) only maintained this last placement. For this study, I examined one paralectotype male of T. sauteri Villeneuve, photographs of the holotype female and paratype male of Orectocerina atratula Malloch (= T. sauteri), and holotype male of T. yunnanensis Chao & Zhou. Based on the available evidence, I consider the genus Trischidocera as incertae sedis, remove it from the tribe Ormiini, and relegate it to the unplaced Tachinidae. Despite this placement, a discussion is provided below considering the possible systematic placement of Trischidocera in the tribe Siphonini or near it.

Accepted by J. O'Hara: 5 Feb. 2015; published: 5 Mar. 2015

279

Material and methods The examined material is deposited at the following institutions: The Natural History Museum, London, United Kingdom (BMNH) and National Museum of Natural History, Washington DC, USA (USNM). Other repositories cited in the text: Hungarian Natural History Museum, Budapest, Hungary (HNHM); National Zoological Museum of China, Institute of Zoology, Chinese Academy of Sciences, Beijing, China (NZMC); and Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany (SDEI). The paralectotype male of T. sauteri in USNM was photographed using a stereomicroscope Leica Z16 and a digital camera JVC (KY-F75U). Images were captured, combined and scaled using the software Cartograph 7.2.5 and Archimed 6.1.4 (Microvision Inc.). The holotype male of T. yunnanensis in NZMC was kindly photographed by Lili Zhang and Xiao-lin Chen (curators). The holotype female and paratype male of Orectocerina atratula in BMNH was kindly photographed by Nigel Wyatt (curator). Photographs were edited using the software Adobe Photoshop CS5, and line drawings were prepared and edited in Adobe Illustrator CS5. All plates were prepared in Adobe Illustrator CS5. The morphological terminology follows Cumming & Wood (2009) and Wood & Zumbado (2010), but for antennal morphology follows Stuckenberg (1999). Label data of type specimens are quoted within double quotation marks, and different lines, if present, are separated by a slash. Geographical records were gathered based on label data and then georeferenced by searching in Google Maps. Distributional maps were prepared using software QGIS 2.4.

Results Trischidocera Villeneuve Trischidocera Villeneuve, 1915: 93. Type species: Trischidocera sauteri Villeneuve, 1915, by monotypy. Orectocerina Malloch, 1924: 521. Type species: Orectocerina atratula Malloch, 1924 [= Trischidocera sauteri Villeneuve, 1915], by original designation. (Synonymy by Townsend, 1939: 357, and confirmed here). References. Villeneuve, 1915: 93 (described Trischidocera and the type species, in “Thryptoceratidae”); Malloch, 1924: 521 (described Orectocerina and the type species); Townsend, 1932: 57 (notes on syntype of T. sauteri, in “Actiini”); Townsend, 1936: 224 (in key to Germariini genera); Townsend, 1939: 357 (synonymy of Orectocerina, in Germariini); Crosskey, 1967: 21, 30 (list); Crosskey, 1976: 54 (cat., in key to tribes, but as unplaced Tachinidae); Chao & Zhou, 1987: 208 (described T. yunnanensis from China); Chao et al., 1998: 1957 (cat., in key to Ormiini genera); O’Hara et al., 2009: 161 (cat., in Ormiini).

Trischidocera sauteri Villeneuve (Figs 1–3, 8) sauteri Villeneuve, 1915: 94. Lectotype male lost (formerly in HNHM), one paralectotype male remaining in USNM (lectotypification by Townsend, 1932: 57 with the mention of “Male Ht”). Type-locality: Taiwan, Mt. Hoozan [Kaohsiung Hsien, Fengshan]. atratula Malloch, 1924: 521 (Orectocerina). Holotype female (BMNH). Type locality: “Malaya”. (Synonymy by Townsend, 1939: 357, and confirmed here).

Redescription. Male. Body length: 4.3 mm, wing length: 4.1 mm. Coloration: Head (Figs 2–3) blackish with weak pale golden pruinosity on frons, fronto-orbital plate, parafacial and gena; face and occiput with silver pruinosity. Antenna blackish. Proboscis blackish. Thorax (Figs 1, 3) blackish with silver pruinosity on pleura and very pale golden pruinosity on scutum, with four non-pruinose longitudinal stripes; scutellum dark brown. Wing (Fig. 1) hyaline, but the membrane tinged with light brown along the veins; calypters white; halter yellowish. Legs (Fig. 1) blackish with some silver pruinosity. Abdomen (Fig. 1) blackish with silver pruinosity on basal half of tergites. Head (Figs 2–3): Eye bare. Frons with medioclinate (the lower ones) and apparently medioreclinate (the upper ones) setae. Fronto-orbital plate and parafacials with a continuous row of tiny setulae from level of anterior ocellus

280 · Zootaxa 3926 (2) © 2015 Magnolia Press

NIHEI

ventrally to the lower margin of eye, and one short lateroreclinate fronto-orbital seta at level of posterior ocelli. Pair of ocellar setae proclinate and moderately developed. Facial ridge with tiny setulae on lower 2/5. Antenna inserted above the upper margin of eye and at the same level of ocelli. Pedicel reduced, much shorter than scape, with short but stout setulae; postpedicel trilobed, anterior branch bearing stout arista at the tip, and median and posterior branches deriving from a common base; postpedicel notably elongate, its length longer than head height. Lower facial margin moderately warped forwards. Vibrissa moderate, inserted at level of lower facial margin. Oral cavity reduced (narrow) and deep (Fig. 1), so that proboscis and palpus are not exposed and visible as in other tachinids. Palpus, if present, not visible. Proboscis not entirely visible, but retractile and apparently short. Antennal axis slightly shorter than oral axis, giving the head a flat appearance. Gena high. Thorax (Figs 1, 3): Acrostichal setae 2+3, presutural setae weak. Dorsocentral setae 2+3. Posthumeral seta 1, aligned with intra-alar row. Presutural seta 1, aligned with supra-alar row. Postpronotal setae 2. Notopleural setae 2. Intra-alar setae 0+2; intra-postalar seta absent. Postsutural supra-alar seta 1, no prealar seta. Prosternum bare, narrow and flat. Proepisternum bare. Proepisternal and proepimeral setae short and directed upward. Six anepisternal setae, with some few tiny setulae upcurved at upper anterior corner. Katepisternal setae 2+1, upper setae longer than lower seta. Katepimeron bare. Scutellum with one basal, one lateral, one subapical (parallel to each other), and one discal pairs of setae. Wing (Fig. 1): Costal spine undeveloped; base of R4+5 setulose dorsally and ventrally; M vein bent forward to R4+5 without angle (rounded) and ending at wing margin; anal vein not reaching wing margin. Legs: Fore tibia with row of anterodorsal setae and two posterior setae. Mid tibia with one submedian anterodorsal, two short posterodorsal, two posterior and one submedian ventral seta. Hind tibia with four anterodorsal setae, the two basalmost weaker; three posterodorsal setae and two anteroventral setae.

FIGURES 1–3. Trischidocera sauteri Villeneuve, paralectotype male. 1. Lateral habitus. 2. Head, frontal view. 3. Head, lateral view, and thorax, dorsal view. (Scale bars 1 mm.)

MISPLACED GENUS TRISCHIDOCERA

Zootaxa 3926 (2) © 2015 Magnolia Press ·

281

Abdomen (Fig. 1): Syntergite 1+2 with weak lateral marginal setae, excavation not reaching posterior margin. Tergite 3 with one median marginal, one discal and one lateral marginal pair of setae. Tergite 4 with a marginal row of setae and one discal pair of setae. Tergite 5 with discal and marginal rows. Terminalia: Specimen not dissected. Type-material examined. Paralectotype male (USNM) labeled “Formosa / Sauter”; “Mt. Hoozan / 1910.V.”; “Townsend / Genotype / Collection”; “Type” [red label]; “Trischidocera / sauteri / Villen.” [handwritten]; “SYNTYPE [male symbol] / Trischidocera / sauteri / Villeneuve, 1915: 94 / labeled 1991 NEW”. Remarks. Villeneuve (1915: 94) did not designate a holotype from among the three males in the type series, and the subsequent treatment of a syntype as “Ht” [holotype] by Townsend (1932: 57) is accepted as a lectotype fixation (see discussion in O’Hara et al. 2009: 10). With regard to the type-specimens location, Villeneuve (1915: 94) listed his three male syntypes as deposited at Budapest (HNHM). Then, Townsend (1932: 57) examined the “Male Ht” [lectotype] of T. sauteri but cited the depository as “Berlin-Dahlem” [now SDEI]. Later, Townsend (1939: 357) cited Budapest as the “Ht male” [lectotype] location, with “Pts male” in “Rambouillet and Washington”. Townsend (1932) apparently erred in citing the “Male Ht” as deposited in Senckenberg collection because the type catalogue by Rohlfien & Ewald (1974) lists instead three other “sauteri” in Tachinidae (Metopomintho sauteri Townsend, 1927, Chaetanicia sauteri Townsend, 1932, and one later species, Trichoformosomyia sauteri Baranoff, 1934). Perhaps, Townsend (1932) confused the location of Trischidocera sauteri with one of the other “sauteri” deposited at SDEI. The mention of “Ht male ... location, Budapest” by Townsend (1939: 357) is interpreted as a correction to the depository cited in Townsend’s earlier work. In HNHM, all Tachinidae types (and most of the Diptera collection) formerly deposited in the collection were destroyed in 1956 when Budapest was under attack from Soviet troops (Boros 1957, Földvári & Papp 2007). The paralectotype mentioned to be in Rambouillet [Villeneuve’s residence] may be anywhere, in France, back to Budapest, etc. Therefore, the information available is that the last remaining paralectotype is the one deposited in the USNM and studied herein. The holotype female and paratype male of Orectocerina atratula Malloch were kindly photographed by Nigel Wyatt (BMNH). The female has a simple postpedicel, not trilobed as in male, although equally elongate and slender and bearing a terminal arista. All features observed are similar to those of T. sauteri and, based on that, I confirm and keep them as conspecific. Distribution (Fig. 8). Taiwan and Malaysia.

Trischidocera yunnanensis Chao & Zhou (Figs 4–7) yunnanensis Chao & Zhou, 1987: 208. Holotype male (NZMC). Type locality: China, Yunnan, Weixi [as “Weisi”], 2500 m.

Remarks. No material was available for study, but the holotype male deposited at the NZMC (Beijing) was kindly photographed by the curators Lili Zhang and Xiao-lin Chen. The morphological characters of T. yunnanensis discussed below were based on those photographs (Figs 4–7) and on the original description and illustrations by Chao & Zhou (1987). Distribution (Fig. 8). China (provinces of Sichuan, Shanxi, Tibet and Yunnan).

Systematic placement of Trischidocera There are some striking differences between T. sauteri and T. yunnanensis. The head of T. sauteri is clearly wider than high when seen in frontal view (Fig. 2), while slightly wider than high in T. yunnanensis (Fig. 6). The frons is broader and shorter in T. sauteri (Figs 2, 6), and the face, although long in both species, is remarkably broad in T. sauteri. The male postpedicel is the most striking feature notable in both species: it is trilobed and slender in T. sauteri, with a terminal arista on the anterior branch (Figs 2–3), while in T. yunnanensis the postpedicel is simple (not branched), slender and tapering to the apex, and also bears a terminal arista (Figs 6–7). The male postpedicel of T. yunnanensis resembles the female postpedicel of T. sauteri rather than that of the male.


NIHEI

FIGURES 4–7. Trischidocera yunnanensis Chao & Zhou, holotype male. 4. Dorsal habitus. 5. Lateral habitus. 6. Head, frontal view. 7. Head, lateral view. (Scale bars 1 mm, except Fig. 7 in 0.5 mm.)

Despite the differences, these two species share some features that apparently indicate a congeneric placement, namely, the terminal arista (distally inserted on the postpedicel); the general chaetotaxy on thorax; setulose parafacial (in T. sauteri the setulae are more numerous and in a row continuous with the fronto-orbital setulae) (Figs 2–3 and 6–7); wing (Figs 1, 5) shape and venation (M bend and the dm-cu nearly straight), and chaetotaxy (all veins similar in both species); and the abdominal chaetotaxy. Perhaps further examination of male and female terminalia will show more definitively whether T. sauteri and T. yunnanensis are congeneric, or whether they deserve to be separated in distinct genera. The systematic placement of Trischidocera had been changed from Siphonini (Villeneuve 1915) to Germariini (Townsend 1936, 1939), but it was thereafter considered an unplaced Tachinidae (Crosskey 1976), and only recently retrieved and transferred to Ormiini (Chao et al. 1998, O’Hara et al. 2009). Definitely it does not belong to Ormiini, as external morphological features do not support the placement of Trischidocera in Ormiini. The Ormiini members show several characters contrasting to those observed in Trischidocera: the arista is anterobasally inserted in the postpedicel; the facial ridge is flattened and broadened to a greater or lesser degree (from the oestridlike forms such as Aulacephala and Therobia to the more discrete forms as in all other ormiines); inflated and welldeveloped prosternum; crossveins dm-cu and r-m not close to each other (their distance conspicuously longer than the length of dm-cu); M vein with a sharply angled bend; and abdomen globose. Unfortunately, male and female terminalia of Trischidocera were not dissected and examined due to the scarcity of material available for study. To the best of my knowledge, this genus is represented in collections by a few specimens, including the four type specimens: male paralectotype of T. sauteri, female holotype and male paratype of O. atratula (=T. sauteri), and male holotype of T. yunnanensis. Chao & Zhou (1987) provided illustrations of the male terminalia in lateral and dorsal views (Figures 2b–c of Chao & Zhou 1987), but without details of the phallus and associated structures (pregonite, postgonite, etc.), which would provide important evidence for placement of Trischidocera. However, what is possible to infer based on Chao & Zhou’s (1987) description and illustrations is that the epandrium, MISPLACED GENUS TRISCHIDOCERA


283

surstylus, distiphallus and pregonite of Trischidocera do not resemble those observed in Ormiini. In this tribe, for example, the epandrium is strongly emarginate at the innerlower margin, where the surstyli have enlarged basal processes; the apex of the surstylus surpasses the apices of the cerci; the distiphallus is strongly folded downwards; and the pregonite is not visible with a free distal end, it is articulated and connected basally to the base of the phallus and distally to the lower posterior margin of the hypandrium. That Trischidocera is not an Ormiini seems to be supported by the arguments above, but what is its proper tribal placement? Based on the available evidence (i.e., without examining the male and female terminalia in detail), it is not possible to put this genus into any tribal placement with confidence. However, there are some features observed in Trischidocera that could indicate a possible inclusion within the tribe Siphonini. Based on external characters, Trischidocera fits most of the diagnostic characters of Siphonini discussed by O’Hara (1989) and Andersen (1996), with special reference to: i) setulose parafacial; ii) bare eyes; iii) shape of aristomeres; iv) intra-alars 0+2, with the postsutural ia placed posteriorly, not close to the suture; v) katepisternals 2+1; vi) the excavation on syntergite 1+2 short, not reaching the posterior margin; vii) median discals on tergites 3 and 4 (which occurs in some Siphonini); viii) the anal vein fading out before wing margin; ix) the smooth (not angulate) bend of M; and x) the nearly straight dm-cu.

FIGURE 8. Geographical distribution of the genus Trischidocera (circles: T. sauteri; squares: T. yunnanensis) [empty symbols denote unexact locality records, where the country or province capitals were used to plot the occurrence].

But while converging in some features, Trischidocera also shows some characters contrasting with the common characters observed in the Siphonini: i) there are no reclinate frontal setae (coded as “present” by O’Hara 1989 and coded as having “2-3 setae” by Cerretti et al. 2014); ii) the prosternum is bare (“normally” setulose according to Andersen 1996 and indicated as a homoplastic character state shared by the Siphonini clade in Cerretti et al. 2014); iii) its wing shape is slightly more elongate than the short and broad wing of Siphonini (sec Andersen 1996); iv) R4+5 is setulose only at the base (but Andersen 1996: 29 describes it as widely polymorphic as “dorsally is either bare, setulose distally or setulose along its entire length” and ventrally “either bare or setulose distally”); and v) section of M between r-m and dm-cu crossveins is conspicuously shorter than the following section of M between dm-cu and the bend (this was indicated as a homoplastic character shared by the Siphonini clade in Cerretti et al. 2014, but this is probably a result of low taxonomic sampling as this character is polymorphic within


NIHEI

the group, for many Siphonini species present the condition observed in Trischidocera). Additionally, the two proclinate fronto-orbital setae common in many Siphonini are apparently absent in Trischidocera, which has a row of short setulae on the entire extent of the fronto-orbital plate. On the scutellum, the discal setae are absent, and subapical setae are parallel to each other, not crossed. Cerretti et al. (2014) performed the first cladistic analysis of the family Tachinidae, although the taxonomic sampling was nearly exclusively Palaearctic taxa (only 1 Oriental, 1 Nearctic and 8 Afrotropical tachinid representatives were included in their analysis along with 482 Palaearctic species). According to the available cladograms so far for Palaearctic Tachinidae, the Siphonini clade was supported by only two synapomorphies, namely, the anterior portion of the pregonite membranous and the number of spermathecae reduced to two. These characters were already discussed by Andersen (1996), but unfortunately I could not examine the male and female terminalia of Trischidocera. However, as illustrated by Chao and Zhou (1987), the enlarged surstylus in lateral view (figs 2b–c of Chao & Zhou 1987) and the enlarged, stout and curved pregonite (fig. 2c of Chao & Zhou 1987) seem to be similar to those of Siphonini. Collecting efforts are required in China, Taiwan and southeastern Asia to obtain additional specimens of Trischidocera. The study and dissection of additional males and females of both T. sauteri and T. yunnanensis will certainly enhance its morphological characterization and will subsequently allow a more conclusive decision about the systematic placement of Trischidocera. In Siphonini, there are genera in which modified forms of postpedicels (bilobed, trilobed or pectinate) have independently appeared (O’Hara 1989), like Ceromya Robineau-Desvoidy and Actia Robineau-Desvoidy in which some species have a bilobed postpedicel, Peribaea Robineau-Desvoidy which can have a bilobed, trilobed, or pectinate postpedicel, and Borgmeiermyia Townsend in which all species have a pectinate postpedicel (O’Hara 1989; Andersen 1996; Nihei & Toma 2010).

Acknowledgements Thanks to Nigel Wyatt (BMNH) and Norman Woodley (USNM) for the loan of material. I am indebted to Erica McAllister and Nigel Wyatt (BMNH) and to Norman Woodley and Torsten Dikow (USNM) for facilities and kind assistance during my visits, and to Matthew Buffington (USNM) for kindly allowing me to use his image capturing system for taking photographs. I am especially grateful to Lili Zhang and Xiao-lin Chen (NZMC) and Nigel Wyatt (BMNH) for kindly taking the photographs of the holotype of T. yunnanensis and of the holotype Orectocerina atratula, respectively, and to Dong Zhang (Beijing Forestry University) for kindly translating parts of Flies of China. Thanks to Chun-tian Zhang (Shenyang Normal University) for kindly providing useful information about Chinese Ormiini, and to Joachim Ziegler (Museum für Naturkunde, Berlin) for helping me during the search for the T. sauteri type. Thanks to Rodrigo Dios, Priscylla Moll, Filipe Gudin, Pedro Dias, Norman Woodley and two anonymous reviewers for suggestions on an earlier version of this manuscript. Financial support from FAPESP (proc. n. 2007/50836-7, and 2013/05131-6).

References Andersen, S. (1996) The Siphonini (Diptera: Tachinidae) of Europe. Fauna Entomologica Scandinavica, 33, 1–146. Bigot, J.M.F. (1882) Diptères nouveaux ou peu connus. 19e partie. XXIX. Genres Roeselia, Actia, Melia, Phytomyptera et tribu des Anthomyzidae (Schiner, Rondani, Meade). XXX. Genre Ctenostylum. Annales de la Société Entomologique de France, Series 6, 2, 5–21, 21–22. Boros, I. (1957) The tragedy of the Hungarian Natural History Museum. Annales Historico-Naturales Musei Nationalis Hungarici, 8, 491–504. Cerretti, P., O’Hara, J.E., Wood, D.M., Shima, H., Inclan, D.J. & Stireman, J.O. (2014) Signal through the noise? Phylogeny of the Tachinidae (Diptera) as inferred from morphological evidence. Systematic Entomology, 39, 335–353. http://dx.doi.org/10.1111/syen.12062 Chao, C. & Zhou, S. (1987) New species of tachinid flies from Hengduan Mountains of China (Diptera: Tachinidae). Sinozoologia, 5, 207–215. [in Chinese] Chao, C.-M., Liang, E.-Y., Shi, Y.-S., Zhou, S.-X., Sun, X.-K. & Chen, R.-J. (1998) Tachinidae. In: Xue, W.-Q. & Chao, C.-M. (Eds.), Flies of China. Vol. 2. Liaoning Science and Technology Press, Shenyang, pp. 1661–2206 + pls. 1–30. [in Chinese] Crosskey, R.W. (1967) An index-catalogue of the genus names of Oriental and Australasian Tachinidae (Diptera) and their

MISPLACED GENUS TRISCHIDOCERA


285

type-species. Bulletin of the British Museum (Natural History), 20 (1), 1–39. Crosskey, R.W. (1976) A taxonomic conspectus of the Tachinidae (Diptera) of the Oriental Region. Bulletin of the British Museum (Natural History), 26 (Entomology Supplement), 1–357. Cumming, J.M. & Wood, D.M. (2009) Adult morphology and terminology. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E. & Zumbado, M.A. (Eds.), Manual of Central American Diptera. Vol. 1. NRC Research Press, Ottawa, pp. 9–50. Földvári, M. & Papp, L. (2007) Damage in the Diptera Collection of the HNHM, Budapest in the year of 1956. Studia dipterologica, 14, 25–26. Malloch, J.R. (1924) Exotic Muscaridae (Diptera). – XIV. Annals and Magazine of Natural History, Series 9, 14, 513–522. http://dx.doi.org/10.1080/00222932408633150 Nihei, S.S. & Toma, R. (2010) Taxonomic notes on Borgmeiermyia Townsend (Diptera, Tachinidae) with the first host record for the genus. ZooKeys, 42, 101–110. http://dx.doi.org/10.3897/zookeys.42.190 O’Hara, J.E. (1989) Systematics of the genus group taxa of the Siphonini (Diptera: Tachinidae). Quaestiones Entomologicae, 25, 1–229. O’Hara, J.E., Shima, H. & Zhang, C.-T. (2009) Annotated catalogue of the Tachinidae (Insecta: Diptera) of China. Zootaxa, 2190, 1–236. Robineau-Desvoidy, J.B. (1851) Myodaires des environs de Paris (Suite). Annales de la Société Entomologique de France, Series 2, 9, 177–190. Rohlfien, K. & Ewald, B. (1974) Katalog der in den Sammlungen des ehemaligen Deutschen Entomologischen Institutes aufbewahrten Typen – XI. (Diptera: Cyclorrhapha: Schizophora: Calyptratae). Beiträge zur Entomologie, 24, 107–147. Stuckenberg, B.R. (1999) Antennal evolution in the Brachycera (Diptera), with a reassessment of terminology relating to the flagellum. Studia dipterologica, 6, 33–48. Townsend, C.H.T. (1932) Notes on Old-World oestromuscoid types. – Part II. Annals and Magazine of Natural History, Series 10, 9, 33–57. http://dx.doi.org/10.1080/00222933208673463 Townsend, C.H.T. (1936) Manual of Myiology in twelve parts. Part III. Oestroid classification and habits. Gymnosomatidae to Tachinidae.Townsend & Filhos, Itaquaquecetuba, 249 pp. Townsend, C.H.T. (1939) Manual of Myiology in twelve parts. Part VIII. Oestroid generic diagnoses and data. Microtropesini to Voriini. Townsend & Filhos, Itaquaquecetuba, 405 pp. Villeneuve, J. (1915) Nouveaux myodaires supérieurs de Formose. Annales Musei Nationalis Hungarici, 13, 90–94. Wood, D.M. & Zumbado, M.A. (2010) Tachinidae. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E. & Zumbado, M.A. (Eds.), Manual of Central American Diptera. Vol. 2. NRC Research Press, Ottawa, pp. 1343–1417.


NIHEI