Tijdschrift voor Entomologie 156 (2013) 127–139 brill.com/tve

Hybotidae of the Galápagos Islands (Diptera: : Tachydromiinae) Bradley J. Sinclair & Jeffrey M. Cumming

Three genera (Isodrapetis Collin, Walker, Elaphropeza Macquart) and six species of Tachydromiinae () are recorded from the Galápagos Islands. One new species of Isodrapetis (I. meridionalis sp. n.)andthreenewspeciesof Chersodromia (C. floreana sp. n., C. galapagensis sp. n., C. isabela sp. n.)are described. Elaphropeza zonalis (Curran) is redescribed and a fourth species of Chersodromia is identified based on a single female specimen. A key to all Galápagos species is provided. Isodrapetis is recorded for the first time outside of New Zealand. The affinities of the Galápagos Tachydromiinae to the New World fauna are discussed. Keywords: Galápagos Islands, Neotropics, Hybotidae, Empidoidea, Elaphropeza, Chersodromia, Isodrapetis, new species. Bradley J. Sinclair*, Canadian National Collection of and Ottawa Plant Laboratory – Entomology, CFIA, K.W. Neatby Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, K1A OC6, Canada. [email protected] Jeffrey M. Cumming, Invertebrate Biodiversity, Agriculture and Agri-Food Canada, K.W. Neatby Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, K1A OC6, Canada. [email protected]

Introduction datazone.darwinfoundation.org/terrestrial-inverte Knowledge of the Diptera of the Galápagos Islands brates/diptera/. has progressed tremendously over the past 20 years, The empidoid flies that are known to occur on facilitated primarily on the basis of specimens from the Galápagos Islands include the , the surveys of 1985–1996 organized and previously treated by Bickel & Sinclair (1997), and directed by Stewart Peck (Peck 2001, 2006; Sin- the Hybotidae. Among the hybotid flies (Tachy- clair 2009). Modern revisions have been published dromiinae), one species was previously recorded from for many families based on this material, includ- the Galápagos, Elaphropeza zonalis (Curran). This ing Agromyzidae (Sasakawa 2007), Asteiidae (For- species was collected during the Norwegian Zoologi- rest & Wheeler 2002), Canacidae: Tethininae (Foster cal Expedition of 1925 (Curran 1932). Based on ex- & Mathis 2008), Carnidae (Wheeler 2000), Cerato- tensive recent collections, four new species are de- pogonidae (Borkent 1991), Chloropidae (Wheeler scribed in this study and inter-island distributions & Forrest 2003), Chyromyidae (Wheeler & Sin- of all species are listed. All the Galápagos Hybotidae clair 1994), Ephydridae (Mathis 1995), and Heleo- reported in this paper belong to the tachydromiine myzidae: Trixoscelidinae (Foster & Mathis 2011). tribe Drapetini (= Drapetidini) as defined by Sin- A checklist of the species of Diptera of the clair & Cumming (2006: 79, but see correction in Galápagos islands is available at: http://checklists. Sinclair & Cumming 2007: 37). The affinities of the

Tijdschrift voor Entomologie 156: 127–139, Figs 1–18. [ISSN 0040-7496]. brill.com/tve © 2013 Government of Canada. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommercial 3.0 Unported (CC BY-NC 3.0) License, http://creativecommons.org/licenses/by-nc/3.0/. Published by Koninklijke Brill NV, Leiden. Published 20 December 2013. DOI 10.1163/22119434-00002026

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Galápagos Drapetini with the New World fauna are Cruz)(Fig.11)...... also discussed...... Chersodromia galapagensis sp. n. – Palpus, halter and legs (at least ventrally) yellowish brown. Thoracic setae pale, espe- Material and methods cially when viewed anteriorly. Right sursty- This study is based on specimens borrowed or de- lus slender, apex not forked (Figs 8, 9, 12, posited in the following collections: American Mu- 13)...... 5 seum of Natural History, New York, USA (AMNH); 5. Right epandrial lamella with dense row of California Academy of Sciences, San Francisco, USA long stout setae (Isla Floreana) (Fig. 8) . . . . (CAS); Canadian National Collection of Insects, Ot- ...... Chersodromia floreana sp. n. tawa, Canada (CNC). – Right epandrial lamella with sparse row of Terms used for adult structures primarily follow long slender setae (Isla Isabela) (Fig. 12). . . those of Cumming & Wood (2009), except for the ...... Chersodromia isabela sp. n. antenna and wing venation where the terms of Stuck- enberg (1999) and Saigusa (2006) are used respec- tively. In the system outlined by Saigusa (2006), the Chersodromia Walker dipteran wing vein A1 (as used in McAlpine 1981) is Chersodromia Walker, 1849: 1157. Type species: Ta c hy pez a homologized with the mecopteran CuP, and conse- brevipennis Zetterstedt, 1838, by subsequent designation of Rondani (1856: 147) [= arenaria (Haliday, 1833)]. quently CuA1 (of McAlpine) is termed M4,whereas Coloboneura Melander, 1902: 229. Type species: Coloboneura CuA2 is CuA, the anal cell is cell cua and the anal vein (A + CuA )isCuP+ CuA. Homologies of inusitata Melander, 1902, by monotypy. 1 2 Halsanalotes Becker, 1902: 41. Type species: Halsanalotes the male terminalia follow those of Sinclair & Cum- amaurus Becker, 1902, by monotypy. ming (2006). Thinodromia Melander, 1906: 370. Type species: Thinodromia Label data for primary types are cited from the inchoata Melander, 1906, by monotypy. top downward, with the data from each label in quotation marks. Labels are cited in full, including Recognition original spelling, punctuation, and date, with label The genus is distinguished from other tachydromiine lines delimited by a slash (/). Additional information genera by its nearly equal length basal cells (br and is included in square [ ] brackets. The repository bm), widely separated eyes on face and frons, broad of each type is given in parentheses. Secondary type gena, and bristly legs (particularly tibiae). data are abridged and listed alphabetically.

Remarks Key to genera and species of Galápagos Chersodromia is primarily confined to marine shore- Hybotidae lines with more than 55 described species worldwide 1. Cell br equal to or longer than cell bm at (Grootaert et al. 2007). Four species are currently apex(Figs4,6)...... 2 recognized from the Galápagos, readily divided into – Cell br much shorter than cell bm at apex two groups on the basis of the shape of the postpedi- (Figs2,5)...... Elaphropeza zonalis (Curran) cel and position of the stylus. The first group (Cherso- 2. Eyes widely separated on face and frons; dromia undescribed sp.), referred to here as the elon- gena distinctly extended below eye; legs gate postpedicel group, also occurs in Central Amer- bristly (Fig. 1) ...... Chersodromia Walker...3 ica, with two described species (Arnaud 1975). The – Eyes nearly contiguous on face; gena scarce- second group with short postpedicel and subdorsal ly extended below eye; leg clothed in long stylus, comprises three species (C. floreana sp. n., C. slender setae, lacking distinct isolated bris- galapagensis sp. n., C. isabela sp. n.)eachconfined tles(IslaEspañola)(Fig.3)...... to a single main island. An additional dark coloured ...... Isodrapetis meridionalis sp. n. specimen (female only) that belongs to this second 3. Postpedicel more than 2 times longer than group is known from the northernmost isolated is- basal width; stylus arising apically (Isla Is- land of Darwin, but a male specimen is required to abela) ...... Chersodromia undescribed species. assess its relationship with the other three short post- – Postpedicel less than 2 times longer than pedicel species. basal width; stylus arising subdorsally The presence of these two species groups indicates (Fig.1)...... 4 that there was a minimum of two colonization or dis- 4. Palpus, halter and legs brown (Fig. 1). Tho- persal events from the mainland. The short postpedi- racic setae dark. Right surstylus somewhat cel species appear to be closely related and possibly a broad and weakly forked apically (Isla Santa monophyletic group on the basis of the male termi-

Downloaded from Brill.com10/04/2021 07:59:01AM via free access Sinclair & Cumming: Hybotidae of the Galápagos Islands (Diptera: Empidoidea: Tachydromiinae) 129 nalia, particularly the shape of the left cercus and left with sparse row of posteroventral setae with long surstylus. basal seta; 1 strong anterodorsal and anteroventral preapical seta. Mid tibia with 2 anterodorsal and Chersodromia floreana sp. n. 1 anterior preapical setae; some weak anteroventral spiny-setae on apical half. Hind femur similar in Figs 8, 9 thickness to other femora, posterior face somewhat Type material. Holotype,  labelled: “ECU: Galap: flattened and arched medially. Hind tibia with stout Floreana/Pt. Cormorant, littoral/white sand beach setae on apical half: 3 anterodorsals, 4 anteriors, 3 26.iii.1989/BJ Sinclair”; “HOLOTYPE/Cherso- anteroventrals. dromia/floreana/Sinclair & Cumming [red label]” Wing (length 1.5–1.6 mm) infuscate, with brown (CNC). Paratypes: Ecuador. Galápagos. Floreana: veins. Basal costal seta shorter than postalar bris- same data as holotype (2,CNC). tle, brown. Vein Rs nearly 4 times longer than crossvein bm-m (bm-cu). R2+3 joining costa be- Recognition yond mid-length. M1+2 and R4+5 parallel at wing This species is distinguished by the short postpedicel tip. Crossvein bm-m (bm-cu) strongly oblique. and subdorsal stylus; pale whitish palpi and halter Crossvein r-m slightly beyond cell bm. No anal vein. and generally pale legs, especially fore coxa and Squama minute. Halter with pale whitish knob. ventral faces of femora; right epandrial lamella with Abdomen brown, subshining, paler than thorax. row of long stout setae. Terminalia (Figs 8, 9): Left cercus mitten-shaped, arched and well sclerotized. Right cercus not differ- entiated. Left epandrial lamella smaller than right Description lamella; fused basally with hypandrium. Right epan- Male: Head brown in ground-colour, clothed in thin drial lamella with dense row of long stout setae; pos- greyish pruinescence; palpi yellowish white; eyes red- terior margin produced into slender finger-like pro- dish brown. Frons slightly divergent towards ocellar jection bearing pair of short setulae. Left surstylus triangle; ventrally as wide as pedicel. Eyes separated divided into short finger-like setose inner lobe and on face, slightly narrower than ventral margin of face; broad outer lobe; outer lobe with rounded apical divergent towards mouthparts. Ommatidia enlarged margin, tapered to sharp posterior apex. Right sursty- on lower anterior half of eye. Anterior and poste- lus flattened, thumb-like in shape. Hypandrium with rior ocellars long, brown, subequal in length. One narrow, posteroapical finger-like prolongation. pair of convergent vertical bristles, slightly longer Female: Unknown. than ocellars. Postoculars very short, pale, uniserial. Gena narrow, one-quarter height of eye. Occiput somewhat flattened above neck. Antennal segments Etymology brown; pedicel with several light brown setae ven- The species is named after the island where the trally; postpedicel slightly longer than pedicel; post- species was collected. pedicel with dorsoapical extension; stylus more than 2 times as long as pedicel and postpedicel combined. Distribution Palpus broad, tapered apically, half length of pro- This species is endemic to the Galápagos Archipela- boscis, with 1 pale terminal seta and some setulae. go, recorded from the island of Floreana. Thorax dark brown, concolorous with head, with dark brown bristles and golden setae. Scutum Remarks clothed in thin pruinescence, lacking vittae; katepis- This species was collected on white sand beaches. ternum with small shiny spot without pruinescence. Postpronotal lobe with 1 long inclinate bristle and 1–2 anterior setae. Mesonotum with numerous short Chersodromia galapagensis sp. n. setae amongst outstanding bristles; 2 notopleurals; 1 Figs 1, 4, 10, 11 postsutural supra-alar; 1 postalar; 2 pairs of scutel- Type material. Holotype,  labelled: “ECU: Galap: lars; acrostichals biserial; 3 long dorsocentrals. St. Cruz/CDRS [Charles Darwin Research Sta- Legs generally pale with fore coxa and ventral faces tion], littoral, 5.ii.1989/white sand beach, BJ Sin- of femora yellowish brown; tarsi increasingly darker clair”; “HOLOTYPE/Chersodromia/galapagensis/ apically. Fore femur thickened, slighter broader than Sinclair & Cumming [red label]” (CNC). Pa- mid femur; with sparse row of short antero- and ratypes: Ecuador. Galápagos. Caamaño: littoral posteroventral setae and 1 long preapical antero- zone, 3–10.iv.1989, yellow pan trap, B.J. Sinclair and posteroventral seta. Fore tibia with 1 distinct (1,1,CNC).Santa Cruz:CharlesDarwinRe- antero- and posteroventral preapical seta. Mid femur search Station (CDRS), littoral zone, 5.ii.1989, 10–

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Figs 1–3. –1,Chersodromia galapagensis sp. n.; 2, Elaphropeza zonalis (Curran); 3, Isodrapetis meridionalis sp. n. Scale bars = 1.0 mm. 17.xi.2001, B.J. Sinclair (24,19,CNC);same Description locality, 19.i–9.ii.1989, 12–13.xi.2001, yellow pan Male: Head brown in ground-colour, clothed in thin traps (18,24, CNC); same locality, 4–9.ii.1989, greyish pruinescence; palpi pale brown; eyes reddish dung trap (2, CNC); same locality, 27.i.1989, brown. Frons slightly divergent towards ocellar tri- beach at high tide line (4,CNC). angle; ventrally as wide as pedicel. Eyes separated on face, slightly narrower than ventral margin of face; Recognition divergent towards mouthparts. Ommatidia enlarged This species is distinguished by the short postpedicel on lower anterior half of eye. Anterior and poste- and subdorsal stylus; brown coloured legs and palpi; rior ocellars long, brown, subequal in length. One and shape of the left cercus and left surstylus. pair of convergent vertical bristles, slightly longer

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terior setae. Mesonotum with numerous short setae amongst outstanding bristles; 2 notopleurals; 1 post- sutural supra-alar; 1 postalar; 2 pairs of scutellars; acrostichals biserial; 3 long dorsocentrals. Legs dark brown, concolorous with thorax. Fore femur thickened, slighter broader than mid femur; with sparse row of short antero- and posteroven- tral setae and 1 long preapical antero- and pos- teroventral seta. Fore tibia with 1 distinct antero- and posteroventral preapical seta. Mid femur with sparse row of posteroventral setae with long basal seta; 1 strong anterodorsal and anteroventral preapi- cal seta. Mid tibia with 2 anterodorsal and 1 anterior preapical setae; some weak anteroventral spiny-setae on apical half. Hind femur similar in thickness to other femora, posterior face somewhat flattened and arched medially. Hind tibia with stout setae on apical half: 4 anterodorsals, 2 anteriors, 2 anteroventrals. Wing (length 1.2–1.4 mm) infuscate, with brown veins (Fig. 4). Basal costal seta shorter than postalar bristle, brown. Vein Rs nearly 3 times longer than crossvein bm-m (bm-cu). R2+3 joining costa beyond mid-length. M1+2 and R4+5 parallel at wing tip. Crossvein bm-m (bm-cu) slightly oblique. Crossvein r-m slightly beyond cell bm. No anal vein. Squama minute. Halter with pale brown knob. Abdomen brown with pale pruinescence. Termi- nalia (Figs 10, 11): Left cercus elongate and arched; sclerotized with inner margin with pair of short, rounded subbasal projections. Right cercus very short, setose and membranous. Left epandrial lamella smaller than right lamella; fused basally with hypan- drium. Right epandrial lamella with row of short dorsal setae; posterior margin produced with subapi- cal, short finger-like projection bearing pair of short Figs 4–7. Wings and foreleg tarsomeres. – 4, Chersodro- setae. Left surstylus divided into short finger-like se- mia galapagensis sp. n.; 5, Elaphropeza zonalis (Curran); tose inner lobe and broad outer lobe; outer lobe 6, Isodrapetis meridionalis sp. n.; 7, I. meridionalis sp. with rounded apical margin and inner margin with n., foreleg terminal tarsomeres. Scale bars = 0.5 mm, jagged projections. Right surstylus somewhat broad except Fig. 7 = 0.25 mm. and weakly forked apically. Hypandrium with pos- teroapical sinuous, setose finger-like prolongation. Female: Similar to male, except as follows: Mid than ocellars. Postoculars very short, brown, unis- tibia lacking anteroventral spiny-setae on apical half. erial. Gena narrow, one-seventh height of eye. Oc- Segment 8 1.5 times longer than segment 7; cercus ciput somewhat flattened above neck. Antennal seg- brown, with several brown setulae. ments brown; pedicel with 2 long brown setae ven- trally; pedicel and postpedicel subequal in length; Etymology postpedicel with dorsoapical extension; stylus nearly Named after the archipelago where this species oc- 3 times as long as pedicel and postpedicel combined. curs. Palpus broad, half length of proboscis, with 1 dark terminal seta and some setulae. Thorax dark brown, concolorous with head, with Distribution dark brown bristles and setae. Scutum clothed in This species is endemic to the Galápagos Archipel- thin pruinescence, lacking vittae; katepisternum with ago, known from the following islands: Caamaño large shiny spot without pruinescence. Postprono- (small island in Academy Bay), Santa Cruz. tal lobe with 1 long inclinate bristle and 1–2 an-

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Figs 8–13. Male terminalia. – 8, Chersodromia floreana sp. n.; 9, C. floreana, right surstylus and epandrium; 10, C. galapagensis sp. n., right surstylus and epandrium; 11, C. galapagensis sp. n.; 12, C. isabela sp. n.; 13, C. isabela sp. n., right surstylus and epandrium. Scale bars = 0.1 mm. Abbreviations: lft cerc – left cercus; lft epand – left epandrium; lft sur – left surstylus; rt epand – right epandrium; rt sur – right surstylus.

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Remarks with sparse row of posteroventral setae with long This species was collected on white sand beaches. basal seta; 1 strong anterodorsal and anteroventral preapical seta. Mid tibia with 2 anterodorsal and Chersodromia isabela sp. n. 1 anterior preapical setae; some weak anteroventral spiny-setae on apical half. Hind femur similar in Figs 12, 13 thickness to other femora, posterior face somewhat Type material. Holotype,  labelled: “ECU: Galap: flattened and arched medially. Hind tibia with stout Isabela/Pto. [Puerto] Villamil, littoral/7.iii.1989, setae on apical half: 3 anterodorsals, 4 anteriors, 3 sweeping sand/beach, BJ Sinclair”; “HOLOTYPE/ anteroventrals. Chersodromia/isabela/Sinclair & Cumming [red la- Wing (length 1.4–1.5 mm) lightly infuscate, with bel]” (CNC). Paratypes: Ecuador. Galápagos. Is- pale veins. Basal costal seta shorter than posta- abela: same data as holotype (1,2,CNC);same lar bristle, golden. Vein Rs nearly 3 times longer   locality, yellow pan traps, 6.iii.1989 (1 ,5 ,CNC). than crossvein bm-m (bm-cu). R2+3 joining costa beyond mid-length. M1+2 and R4+5 parallel at Recognition wing tip. Crossvein bm-m (bm-cu) strongly oblique. This species is distinguished by the short postpedicel Crossvein r-m slightly beyond cell bm. No anal vein. and subdorsal stylus; pale whitish palpi and halter Squama minute. Halter with pale whitish knob. and generally pale legs, especially fore coxa and Abdomen brown, subshining, paler than thorax. ventral faces of femora; right epandrial lamella with Terminalia (Figs 12, 13): Left cercus with three row of slender setae. rounded lobes, arched and well sclerotized. Right cercus not differentiated. Left epandrial lamella Description smaller than right lamella; fused basally with hypan- Male: Head brown in ground-colour, clothed in thin drium. Right epandrial lamella with sparse row of greyish pruinescence; palpi yellowish white; eyes red- long slender setae; posterior margin produced into dish brown. Frons slightly divergent towards ocellar slender finger-like projection bearing pair of short triangle; ventrally as wide as pedicel. Eyes separated setulae. Left surstylus divided into short finger-like on face, slightly narrower than ventral margin of face; setose inner lobe and broad outer lobe; outer lobe divergent towards mouthparts. Ommatidia enlarged with rounded apical margin. Right surstylus flat- on lower anterior half of eye. Anterior and poste- tened, thumb-like in shape. Hypandrium with nar- rior ocellars long, brown, subequal in length. One row, posteroapical finger-like prolongation. pair of convergent vertical bristles, slightly longer Female: Similar to male, except as follows: Mid than ocellars. Postoculars very short, pale, uniserial. tibia lacking anteroventral spiny-setae on apical half. Gena narrow, one-quarter height of eye. Occiput Segment 8 subequal in length to segment 7; cercus somewhat flattened above neck. Antennal segments brown, with several brown setulae. brown; pedicel with several light brown setae ven- trally; postpedicel slightly longer than pedicel; post- Etymology pedicel with dorsoapical extension; stylus more than The species is named after the island where the 2 times as long as pedicel and postpedicel combined. species was collected. Palpus broad, tapered apically, half length of pro- boscis, with 1 pale terminal seta and some setulae. Distribution Thorax dark brown, concolorous with head, with This species is endemic to the Galápagos Archipela- golden bristles and setae. Scutum clothed in thin go, recorded from the island of Isabela. pruinescence, lacking vittae; katepisternum with small shiny spot without pruinescence. Postprono- Remarks tal lobe with 1 long inclinate bristle and 1–2 an- This species was collected on white sand beaches. terior setae. Mesonotum with numerous short setae amongst outstanding bristles; 2 notopleurals; 1 post- Chersodromia undescribed species sutural supra-alar; 1 postalar; 2 pairs of scutellars; acrostichals biserial; 3 long dorsocentrals. Material examined. Ecuador. Galápagos. Isabela: Legs generally pale with fore coxa and ventral faces Puerto Villamil, intertidal rocks, 7.iii.1989, B.J. Sin- of femora yellowish brown; tarsi increasingly darker clair (1,CNC). apically. Fore femur thickened, slighter broader than mid femur; with sparse row of short antero- and Recognition posteroventral setae and 1 long preapical antero- This species is distinguished from the other Galapa- and posteroventral seta. Fore tibia with 1 distinct gos Chersodromia by the elongate postpedicel, apical antero- and posteroventral preapical seta. Mid femur stylus and brown coloration.

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Distribution 1997, yellow pan traps, L. Rogue (1,CNC).San This undescribed species is known from the island of Cristóbal: Cerro Tijeretas, S00.53358 W089.36421, Isabela (Galápagos). 8.xii.2004, J. Loayza (1,CNC).Santa Cruz: 5.v.1932, M. Willows, Jr., Templeton Crocker Ex-  Remarks pedition 1933 [Indefatigable Island] (1 ,CAS); This species was collected on black intertidal lava CDRS, tortoise pool, 21.i.1989, alt. 5 m, B.J. Sinclair (1, CNC); Tortuga Bay, littoral zone, rocks. The two very similar species from the Pacific  coast of Central America were also collected on sweeping beach morning glories, B.J. Sinclair (1 , 1, CNC); CDRS, 20–30.i.1989, alt. 10 m, MT, intertidal rocks (Arnaud 1975; Cumming & Sinclair  2009). This species is known from a single female S. Peck, 89-209b (1 ,CNC);CDRS,arid/littoral zone, 10–13.xi.2001, yellow pan traps, B.J. Sin- specimen and is not described here, because males  are required for accurate species diagnosis. clair (2 , CNC); CDRS, Academy Bay, littoral zone, 17.xi.2001, B.J. Sinclair (7,5,CNC);Pto.Ay- ora, Las Ninfas, S0.74592, W90.31886, 20.iii.2006, Elaphropeza Maquart A. Ulloa, R. Miranda. McPhail 38. Muyuyo (1, Elaphropeza Macquart, 1827: 86. Type species: Ta c hyd ro m i a CNC). ephippiata Fallén, 1815, by monotypy. Ctenodrapetis Bezzi, 1904: 351 (as subgenus of Drapetis Recognition (following descriptive format of Meigen). Type species: Drapetis (Ctenodrapetis) ciliato- costa Bezzi, 1904, by subsequent designation of Melander Shamshev & Grootaert 2007) (1928: 309). Species with dark thorax; dark antenna, postpedicel about 3 times as long as wide, stylus nearly 1.5 times as long as postpedicel; palpus white; hind tibia with Recognition 1 anterodorsal seta; halter pale; abdominal tergite 4 The genus is distinguished from other tachydromi- with lateral squamiform setae. ine genera by its short basal radial (br) cell com- pared with basal medial (bm) cell, anal vein (CuA + CuP) lacking, occiput convex, eyes nearly contigu- Re-Description (following format of Shamshev & ous on face, frons divergent, gena narrow, antenna Grootaert 2007) with postpedicel conical to lanceolate and stylus ter- Male: Occiput entirely dark brown, subshining, with minal, anepisternum bare without scattered setulae, yellowish setation. Anterior ocellars long, proclinate abdomen with intersegmental gland-like modifica- and cruciate; posterior ocellars slightly longer than tions between tergites 3–5 or 4–5. anterior ocellars. Inner verticals two-thirds length of long outer verticals. Frons narrow, somewhat broad- ened toward ocellar tubercle, subshining. Antenna Remarks entirely brown. Pedicel with circlet of subequally Freitas-Silva & Ale-Rocha (2011) indicate that there short setulae. Postpedicel short, about 2 times longer are 36 described species of Elaphropeza currently rec- than wide. Stylus normally pubescent, brownish, ognized from the Neotropical Region. However, this nearly 2 times as long as postpedicel. Proboscis yel- is probably just a small portion of the actual diver- low. Palpus whitish, small, rounded; with scattered sity of the region, because the genus is known to be pale setulae, bearing several longer subapical setae. exceedingly diverse throughout the tropics (Sham- Thorax (including postnotum) entirely brown, shev & Grootaert 2007). Freitas-Silva & Ale-Rocha shining, with brownish yellow setation (stronger se- (2009, 2011) provide keys to the seven species from tae darker); anterior spiracle whitish. Proepisternum the Amazon Basin, but the taxonomy of Neotropi- bare, lacking setae. Postpronotum with numerous se- cal Elaphropeza as a whole is mostly obsolete making tulae, lacking prominent seta. Mesonotum with 2 species identification difficult. notopleural, 1 long postsutural supra-alar, 1 posta- lar and 4 scutellar setae (outer scutellars very short); Elaphropeza zonalis (Curran) additionally, some setulae present on notopleural and Figs 2, 5, 14, 15, 16 postsutural supra-alar faces. Acrostichal setae multi- serial, lacking on prescutellar depression. Dorsocen- Drapetis zonalis Curran, 1932: 351; Curran, 1934: 151. Elaphropeza zonalis: Rogers, 1983: 450. tral setae multiserial anteriorly, uniform, ending at Type material examined. Paratype,  labelled: “Wo- pair of long prescutellars. llebæk Galapagos-/5 Eksp. 1925” [Floreana]; “Dra- Legs gradually darkening from whitish coxae to petis/PARATYPE/zonalis /Curran/No.” (AMNH). brownish on tarsi; tarsomere 5 of all legs brown. Additional material examined. Ecuador. Galápagos. Coxae and trochanters with unmodified pale seta- Isabela: Playa Tortuga negra, Mongrove, 6–12.ii. tion. Fore femur with row of long, slender antero-

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Figs 14–18. Male terminalia. – 14, Elaphropeza zonalis (Curran); 15, E. zonalis (Curran); left surstylus; 16, E. zonalis (Curran); right surstylus; 17, Isodrapetis meridionalis sp. n., left surstylus; 18, I. meridionalis sp.n.Scale bars = 0.1 mm. Abbreviations: lft cerc – left cercus; lft sur – left surstylus; rt cerc – right cercus; rt sur – right surstylus. and posteroventral setae; increasing in length prox- Wing (length: 1.7–1.9 mm) normally developed, imally; basal seta subequal to basal width of femur. faintly uniformly infuscate, covered with uniform Fore tibia lacking prominent setae (except subapi- microtrichia; veins brownish yellow (Fig. 5). Costal cals). Mid femur with 2 multi-rows of short ven- vein with moderately long setulae along anterior tral setulae; 1 long thin seta near base and 1 ante- margin. Basal costal seta long, brownish yellow. Vein rior preapical seta. Mid tibia with single row of ven- Rs longer than crossvein bm-m (bm-cu). Veins R4+5 tral spinules, lacking prominent setae (except subapi- and M1+2 somewhat divergent near wing apex. Vein cals). Hind femur with row of short anteroventral se- M4 reaching wing margin as fold. Crossvein r-m tae and row of darkish posteroventral setae on basal beyond middle of cell bm. Halter whitish. third. Hind tibia with anterodorsal seta near middle Abdomen with all tergites (except tergite 4) and slightly shorter anterodorsal seta beyond mid- brownish. Tergite 1 entirely lacking sclerotisation. dle; apical projection small, pointed, clothed in dense Tergites 2 and 3 triangular laterally, broadly con- whitish setulae. Hind tarsomere 1 slightly thickened, cave dorsally and divided along midline, with un- with some short, ventral spinule-like setulae. modified setation. Tergite 4 broadest, brown, with lateral squamiform setae. Tergite 5 narrow, lacking

Downloaded from Brill.com10/04/2021 07:59:01AM via free access 136 Tijdschrift voor Entomologie, volume 156, 2013 squamiform setae. Tergites 6–7 nearly as broad as hair-like leg bristles, and male genitalia with left tergite 4, with long posteromarginal setae. Tergite epandrial lamella fused to hypandrium. 8 very slender and well sclerotized. Paired, spheri- cal gland-like structures present between tergites 4– Remarks 5. Terminalia (Figs 14, 15, 16): Cerci separate, dig- Prior to this study, Isodrapetis was known exclu- itiform, covered with moderately long, unmodified sively from New Zealand with eight described species setae; left cercus slightly longer than right. Epan- (Plant 1999). This is the first record of Isodrapetis drium completely divided. Right epandrial lamella outside of New Zealand. We expand the concept subtriangular, deeply excised, with pair of setae an- of the genus here to include the new Neotropi- terior to base of surstylus. Right surstylus not dif- cal species described below. A second undescribed ferentiated from epandrium, digitiform; right epan- Chilean species of Isodrapetis was also sorted from drial lamella narrow, gradually tapered. Left epan- CNC material during this study, but is not de- drial lamella fused to hypandrium, with 2 short setae scribed here. In Cumming & Sinclair’s (2009) key apically. Left surstylus digitiform, bent at mid-length to the Central American genera of Empidoidea (ex- and tapered to rounded apex; inner face slightly pro- clusive of Dolichopodidae sensu stricto), specimens jected at mid-length; numerous unmodified setae of of Isodrapetis run to couplet 11, which separates different lengths along dorsal margin. Hypandrium Chaetodromia Chillcott from Austrodromia Collin. with truncate apex and 1 subapical, spine-like seta. Isodrapetis can be most easily distinguished from Phallus very short. these genera by inconspicuous short hair-like bristles Female: Similar to male except as follows: Seg- on the legs and lack of prominent dorsocentral bris- ment 8 yellow, somewhat elongate, with sclerites sep- tles on the scutum. Males of Isodrapetis also signifi- arated; tergite 8 with several setae longer posteriorly; cantly differ from those of Austrodromia in the fusion sternite 8 folded apically. Cercus short, rounded, of the left epandrial lamella with the hypandrium. pale. Isodrapetis meridionalis sp. n. Distribution Figs 3, 6, 7, 17, 18 This species is considered endemic to the Galápagos (but see Remarks) and recorded from the following Type material. Holotype  labelled: “CHILE: Re- islands: Floreana, Isabela, San Cristóbal and Santa gion X, Antillanca/road, 700 m, 40°45 S 72°09 W/ Cruz. 8–10.xii.2002, Nothofagus/forest, L. Masner, YPT”; “HOLOTYPE/Isodraptis/meridionalis/Sinclair & Remarks Cumming [red label]” (CNC). Paratypes: Chile: Región X – Los Lagos: same data as holotype (39, The type specimens were collected on Floreana, Post  Office Bay, November 1, 1925 (Curran 1932) and 21 ,CNC).Ecuador. Galápagos: Española:Bahia Manzanilla, littoral Cryptocarpus & Prosopis, FIT, additional specimens have been collected in littoral   and arid zones. Elaphropeza zonalis was re-described MT, 5–10.vi.1985, S. & J. Peck (1 ,1 ,CNC). Additional material examined. Chile: Región by Rogers (1983) who illustrated the male antenna IX – La Araucanía: PN Nahuelbuta, 39°48 10 S and wing. Currently it is difficult to determine if E. zonalis occurs more widely in the Neotropics, 73°01 27 W, alt. 1327 m, 8–10.ii.2005, yellow pan traps, Araucaria/Chusquea forest, UCR AToL C05- because of the incomplete taxonomy for the genus  in the region. 004 (1 , CNC); Cautin, alt. 1150 m, Conguillio Nat. Park, Araucaria/Nothofagus, 4.ii.1988, L. Mas- ner (20,17,CNC).Región X – Los Lagos: Isodrapetis Collin Chiloé Is., Ahoni, 22.ii., iv.1988, primary forest, L.  Isodrapetis Collin, 1928: 6. Type species: Isodrapetis nitidula Pena (3 , CNC); Chiloé Is., Lago Huinchyillo, alt. Collin, 1928. 10 m, yellow pan traps, 11–12.xii.2002, L. Mas- ner (2,3, CNC); Chiloé Is., Chiloé Nat. Park, 42°20 S 74°07 W, sea level, yellow pan traps, L. Mas- Recognition   The genus is distinguished from other tachydromiine ner (7 ,9, CNC); Chiloé Is., alt. 50 m, Terao nr. Chonchi, 19–21.ii.1988, L. Masner (1,CNC); genera by its nearly equal length basal cells (br and  bm), relatively short radial sector (Rs) arising beyond Llanquihue, Ensenada, 13–15.i.1962, L. Pena (1 , midpoint of R ,analvein(CuA+ CuP) weak but CNC); Llanquihue, alt. 200 m, ca. Lago Chapo, 1 Nothofagus forest, 18–19.ii.1988, L. Masner (5, still visible near posterior margin of wing, short fine  acrostichal and dorsocentral bristles, inconspicuous 1 , CNC); Osorno, alt. 250 m, Puyehue Nat. Park, ca. Anticura, Nothofagus forest, ii.1988, L. Masner

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(1,2, CNC); P.N. Vincente Rossales Perez, Mi- ventral pale setae. Hind tibia clothed in long silky rador El Bosque, 41°08.31 S 72°32.2 W, alt. 900 m, brown setae. dung pans, 2.xii.2008, J. Kits, S. Marshall (1,1, Wing (length: 2.4–3.3 mm) normally developed, CNC). Región XIV – Los Ríos: Valdivia, alt. 500 m, faintly uniformly infuscate, covered with uniform 30 km W La Union, Las Truncas, Nothofagus,9– microtrichia; veins brownish yellow (Fig. 6). Costal 12.ii.1988, L. Masner (11,4,CNC). vein with moderately long setulae along anterior margin. Basal costal seta long, brownish. Veins R4+5 Recognition and M1+2 distinctly curved at midlength, divergent Males of this species are distinguished by the comb of at wing margin. Halter yellowish brown. dark apical setae on foreleg tarsomere 3 and flattened Abdomen with all sclerites pale and weakly scle- tarsomere 4, and left surstylus divided in broad U- rotized; anterior margin of tergites brownish. Ter- shaped lobe. Female abdomen mostly membranous, gite 8 very slender and well sclerotized. Terminalia except for shiny, brownish tergites 7 and 8. (Figs 17, 18): Left cercus digitiform, well sclerotized with short basal lobe bearing crown of long setae. Right cercus, elongate and digitiform, nearly sube- Description qual in length to left cercus. Left epandrial lamella Male: Occiput entirely dark brown, clothed in grey broadly fused with hypandrium. Left surstylus di- pruinescence, with dark setation. Anterior ocellars vided into deeply U-shaped process; anterior branch long, proclinate and parallel; posterior ocellars two- clothed in dense setae, gradually tapered to narrow thirds as long as anterior ocellars. Inner verticals apex; posterior branch with expanded and truncate one-third longer than outer verticals. Frons nar- apex bearing long stout apical setae, inner margin row, broadened toward ocellar tubercle, clothed in lined with slender setae. Right surstylus finger-like grey pruinescence. Antenna mostly brown, scape and in shape, elongate, subequal in length to anterior pedicel yellowish brown. Pedicel with circlet of sube- branch of left surstylus. qually short setulae, longer than scape. Postpedicel Female: Similar to male except as follows: Seg- elongate, about 6 times longer than basal width. Sty- ments 7 and 8 well sclerotized, tergites shiny brown; lus pubescent, brownish, slightly more than two- clothed in long pale brown setae. Cercus 2 times thirds length of postpedicel. Proboscis yellow. Pal- longer than wide, with long pale brown setae. pus yellowish, tapered apically to rounded apex; one- half length of proboscis, with scattered pale setu- lae. Etymology Thorax (including postnotum) entirely clothed From the Latin adjective meridionalis, meaning in whitish pruinescence, mesonotum brown, except southern, in reference to the Neotropical distribution postpronotal area yellow and scutellum paler; post- of this species, which is known from Chile and the notum pale brown dorsally, darker on ventral half. Galápagos Islands. Pleura yellowish brown, darker towards legs; spira- cles yellowish. Postpronotum with 1 prominent seta. Distribution Mesonotum with 1 notopleural and 2 short setae, This species is known from Chile and the island of 1 postalar and 4 scutellar setae (outer scutellars half Española in the Galápagos Archipelago. length of inner). Acrostichal setae biserial, lacking on prescutellar depression. Dorsocentral setae multise- Remarks rial anteriorly, uniform, ending at pair of long pres- This species has been collected from the littoral zone cutellars. on Española. In Chile, this species is recorded in Legs gradually darkening from whitish coxae to Araucaria and Nothofagus forests and at altitudes brownish on tarsi; tarsomere 5 of all legs brown. from sea level to 1300 m. Coxae and trochanters with unmodified pale or dark Isodrapetis meridionalis and other identified Neo- setation. Fore femur and tibia lacking prominent tropical species can tentatively be assigned to species setae. Fore tarsomere 3 with dorsoapical row of 5– group III as defined by Plant (1999). The head is 6 black setae, subequal in length to tarsomere 4; spherical, lacking an apparent flattened upper oc- tarsomere 4 with inner face flattened, bearing long ciput and the postpedicel is greatly elongate, longer silky setae (Fig. 7). Mid femur with 2 long, basal the stylus. Although the cerci and right surstylus posteroventral setae; apical two-thirds with row of (= right epandrial lamella sensu Plant 1999) are posteroventral setae, shorter than basal setae. Mid elongate, the abdomen lacks sclerotized bands and tibia with single row of anteroventral spinules. Hind hence can not be assigned to Group I. femur narrowed on basal third; with long dorsal and

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Discussion References The origins and affinities of the Galápagos hybotid Arnaud, P.H., 1975. A new species of Chersodromia from fauna described in this study require interpretation. Mexico. – The Pan-Pacific Entomologist 51: 152–158. The Galápagos record of the New Zealand genus Becker, T., 1902. Aegyptische Dipteren. – Mitteilungen Isodrapetis at first glance might suggest colonization aus dem Zoologischen Museum zu Berlin 2(2): 1–66 + from the western Pacific; however I. meridionalis is 1pl. primarily a Chilean species that probably represents a Bezzi, M., 1904. Indo-australiani raccolti dal recent dispersal event from continental South Amer- Signor L. Biro. – Annales Historico-Naturales Musei ica. Discovery of a second undescribed species of Nationalis Hungarici 2: 320–361. Bickel, D.J. & B.J. Sinclair, 1997. The Dolichopodidae Isodrapetis from Chile indicates that this genus is (Diptera) of the Galápagos Islands, with notes on the not endemic to New Zealand, but that it has been New World fauna. – Entomologica Scandinavica 28: overlooked as a consequence of inadequate study of 241–270. small tachydromiine flies throughout the Neotropi- Borkent, A., 1991. The Ceratopogonidae (Diptera) of the cal Region. Similarly, interpretation of the origin of Galapagos Islands, Ecuador with a discussion of their Elaphropeza zonalis is hampered by our poor knowl- phylogenetic relationships and zoogeographic origins. edge of the genus throughout the Neotropics. Al- – Entomologica Scandinavica 22: 97–122. though apparently a Galápagos endemic, this species Collin, J.E., 1928. New Zealand Empididae Based on may yet be discovered among the many undescribed Material in the British Museum (Natural History). – species known from the Central and South American British Museum (Natural History), London, 110 pp. mainland (Cumming & Sinclair 2009; Freitas-Silva Cumming, J.M. & B.J. Sinclair, 2009. Empididae (dance & Ale-Rocha 2011). flies, balloon flies, predaceous flies). – In: B.V. Brown, The species of Chersodromia from the Galápagos A. Borkent, J.M. Cumming, D.M. Wood, N.E. Wood- are likely due to two colonization events. The un- ley & M.A. Zumbado (eds), Manual of Central Amer- described female with a long postpedicel from Is- ican Diptera, Vol. 1: 653–670. NRC Research Press, abela is very similar to the two described species of Ottawa. Cumming, J.M. & D.M. Wood, 2009. Adult morphology Chersodromia known from the Pacific coast of Cen- and terminology [Chapter] 2. – In: B.V. Brown, A. tral America. Discovery of a male of this species from Borkent, J.M. Cumming, D.M. Wood, N.E. Woodley Isabela might reveal that it is conspecific with one of & M.A. Zumbado (eds), Manual of Central American these Central American species and therefore the re- Diptera, Vol. 1: 9–50. NRC Research Press, Ottawa. sult of a more recent dispersal event. The three short Curran, C.H., 1932. [The Norwegian Zoological Expedi- postpedicel species, however, appear closely related to tion to the Galapagos Islands 1925, conducted by Alf each other on the basis of similar male genitalia, with Wollebæk. IV.] Diptera (excl. of Tipulidae and Culi- each species confined to a single main island (i.e., Is- cidae). – Nyt Magazin Naturvidenskaberne 71: 347– abela, Floreana, Santa Cruz). This suggests an earlier 366. colonization and subsequent radiation on the Galá- Curran, C.H., 1934. The Templeton Crocker Expedition pagos Islands, with a possible fourth species confined of the California Academy of Sciences, 1932. No. 13. to the island of Darwin (known only from a single Diptera. – Proceedings of the California Academy of female). Unfortunately the relationships of this short Sciences, ser. 4, 21: 147–172. postpedicel group to other species of Chersodromia Forrest, J. & T.A. Wheeler, 2002. Asteiidae (Diptera) of found throughout the Pacific are presently unknown. the Galápagos Islands, Ecuador. – Studia dipterologica 9: 307–317. Foster, G.A. & W.N. Mathis, 2008. A review of the Acknowledgements Tethininae (Diptera: Canacidae) from the Galápagos Islands. – Proceedings of the Entomological Society of We are indebted to the following curators and their Washington 110: 743–752. respective institutions for the loan of specimens used Foster, G.A. & W.N. Mathis, 2011. A revision of the in this study: David Grimaldi (AMNH) and Charles Nearctic species of the genus Trixoscelis Rondani Griswold and Norm Penny (CAS). Field work by BJS (Diptera: Heleomyzidae: Trixoscelidinae). – Smithso- in 1989 was partially supported by an NSERC grant nian Contributions to Zoology 637: 1–128. to S.B. Peck (Carleton University, Ottawa). Collect- Freitas-Silva, R.A.P.& R. Ale-Rocha, 2009. Revision of the ing permits were provided by the Galápagos National species of Elaphropeza Macquart, 1827 (Diptera: Hy- Park. BJS thanks Stewart Peck for his invitation to botidae, Tachydromiinae) from Amazon Basin region join his 1989 expedition and continued support over and some remarks about E. ciliatocosta (Bezzi, 1904). – the years. We also thank an anonymous reviewer for Zootaxa 2245: 32–46. offering valuable comments on the manuscript. Scott Freitas-Silva, R.A.P. & R. Ale-Rocha, 2011. Two new Brooks (CNC) kindly produced the photographs. species of Elaphropeza Macquart, 1827 (Diptera, Hy-

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botidae, Tachydromiinae) from Bolivia, Brazil and Sasakawa, M., 2007. The Neotropical Agromyzidae (In- Peru. – Acta Amazonica 41: 567–574. secta: Diptera) Part 7. Leafminers from the Galápagos Grootaert, P., J. Cumming & I. Shamshev, 2007. Non- Islands. – Species Diversity 12: 193–198. marine Chersodromia Walker from Southeast Asia and Shamshev, I.V. & P.Grootaert, 2007. Revision of the genus Papua New Guinea (Diptera, Empidoidea, Hybotidae). Elaphropeza Macquart (Diptera: Hybotidae) from the – Zootaxa 1661: 55–61. Oriental Region, with a special attention to the fauna Macquart, J., 1827. Insectes diptères du nord de la France. of Singapore. – Zootaxa 1488: 1–164. Platypézines, dolichopodes, empides, hybotides. – Lille. Sinclair, B.J., 2009. Dipteran biodiversity of the Galápagos. 159 pp. – In: T. Pape, D. Bickel & R. Meier (eds), Dipteran Di- Mathis, W.N., 1995. Shore flies of the Galápagos Islands versity: Status, Challenges and Tools: 97–118. Konin- (Diptera: Ephydridae). – Annals of Entomological So- klijke Brill NV, Leiden, xx + 459 pp. ciety of America 88: 627–640. Sinclair, B.J. & J.M. Cumming, 2006. The morphology, McAlpine, J.F., 1981. Morphology and terminology – higher-level phylogeny and classification of the Empi- Adults [Chapter] 2. – In: J.F. McAlpine, B.V. Peter- doidea (Diptera). – Zootaxa 1180: 1–172. son, G.E. Shewell, H.J. Teskey, J.R. Vockeroth, & Sinclair, B.J. & J.M. Cumming, 2007. Leptopezella,a D.M. Wood (eds), Manual of Nearctic Diptera, Vol. 1: new Southern Hemisphere genus of Ocydromiinae 9–63. Ottawa, Agriculture Canada Monograph 27. (Diptera: Empidoidea: Hybotidae). – Zootaxa 1629: Melander, A.L., 1902. A monograph of the North Ameri- 27–37. can Empididae. Part 1. – Transactions of the American Stuckenberg, B.R., 1999. Antennal evolution in the Entomological Society 28: 195–367. (Diptera), with a reassessment of terminol- Melander, A.L., 1906. Some new or little-known genera of ogy relating to the flagellum. – Studia dipterologica 6: Empididae. – Entomological News 17: 370–379. 33–48. Melander, A.L., 1928. Diptera, Fam. Empididae. – In: Walker, F., 1849. List of the specimens of dipterous insects O. Wytsman (ed.), Genera Insectorum Fasc. 185: 1– in the collection of the British Museum. Part IV. – 434. Louis Desmet-Verteneuil, Bruxelles. London, pp. 689–1172. Peck, S.B., 2001. Smaller Orders of Insects of the Galápa- Wheeler, T.A., 2000. Carnidae of the Galapagos Islands, gos Islands, Ecuador: Evolution, Ecology and Diversity. Ecuador: description and phylogenetic relationships of – NRC Research Press, Ottawa, 278 pp. a new species of Neotropical Meonura Rondani, 1856 Peck, S.B., 2006. The Beetles of the Galápagos Islands, (Diptera: Carnidae). – Studia dipterologica 7: 115– Ecuador: Evolution, Ecology and Diversity (Insecta: 120. Coleoptera). – NRC Research Press, Ottawa, 313 pp. Wheeler, T.A. & J. Forrest, 2003. The Chloropidae Plant, A.R., 1999. New species of Isodrapetis Collin, 1928 (Diptera) of the Galápagos Islands, Ecuador. – (Diptera, Empidoidea, Hybotidae) with a detailed di- Systematics & Evolution 34: 265–280. agnosis of the genus. – Studia dipterologica 6(2): 279– Wheeler, T.A. & B.J. Sinclair, 1994. Chyromyidae 294. (Diptera) from the Galápagos Islands, Ecuador: Three Rogers, E., 1983. The Neotropical species of Drapetis new species of Aphaniosoma Becker. – Proceedings of Meigen (Diptera: Empididae). – Systematic Entomo- the Entomological Society of Washington 96: 440– logy 8: 431–452. 453. Rondani, C., 1856. Dipterologiae Italicae prodromus. Vol. 1. Genera Italica ordinis Dipterorum ordinatim disposita et distincta et in familias et stirpes aggregata – A. Storschi, Parmae [= Parma], 226 + [2] pp. Saigusa, T., 2006. Homology of wing venation of Diptera. Unpublished handout distributed at the 6th Interna- Received: March 13, 2013 tional Congress of Dipterology, Fukuoka, Japan, 26 pp. Accepted: August 19, 2013

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