Review of the shore-fly genus Oedenopiforma (Diptera: Ephydridae)

81 Review of the shore-fly genus Oedenopiforma (Diptera: Ephydridae) Wayne N. Mathis, 1 Tadeusz Zatwarnicki Abstract The shore-fly genus Oedenopiforma Cogan is reviewed, including new information on Oedenopiforma argentea (distribution), Oedenopiforma javana (comb. nov. and O. orientalis syn. nov.), and Oedenopiforma vockerothi (sp. nov. from United Arab Emirates). Structures of the male terminalia of these three species, a diagnosis of the genus, and a key to the included genera of Dryxini and species of Oedenopiforma are presented. Résumé Nous révisons le genre Oedenopiforma Cogan chez les éphydrides et apportons de nouvelles informations sur Oedenopiforma argentea (répartition), Oedenopiforma javana (nouvelle combinaison et O. orientalis nouvelle synonymie) et Oedenopiforma vockerothi (nouvelle espèce des Émirats arabes unis). Nous présentons des descriptions des structures des derniers segments abdominaux des mâles de ces trois espèces, une diagnose du genre, ainsi qu une clef d identification des genres inclus dans les Dryxini et des espèces d Oedenopiforma. Introduction Publication of a revision frequently spawns further interest in the revised taxon and results in new discoveries that warrant further consideration. This is the case with the shore-fly genus Oedenopiforma Cogan (Diptera: Ephydridae), the subject of this review. Although recently revised (Mathis and Zatwarnicki 2002), we are again reviewing this genus because new and taxonomically valuable information has become available to us. The new information largely stems from study of specimens we recently collected or otherwise became available to us. Field work, including work in collections, continues to be crucial to studies on the systematics of shore flies and ought to be supported. Oedenopiforma was first described as a subgenus of Paralimna Loew and initially included only two species (Cogan 1968): the type species Oedenopiforma madecassa (Giordani Soika) and Oedenopiforma argentea (Cogan). Other than the Afrotropical catalog (Cogan 1980), our world catalog and revision (Mathis and Zatwarnicki 1995, 2002), and a faunistic study from China (Zhang et al. 2009), there are no further references to the genus. Consistent with our previous treatments, we continue to accord generic status to this taxon. Because this is not a comprehensive revision, we have only included species for which new information has become available since the group was last revised (Mathis and Zatwarnicki 2002), such as new species, new combinations, new distributional records, etc. We have included a diagnosis for the tribe and a key to genera to facilitate their identification (many genera of Dryxini are uncommon in collections and are not easily recognized or well known). We dedicate this review to the monumental effort that was initially led by a group of Canadian dipterists to produce the three volumes of the Manual of Nearctic Diptera. That manual set the precedent for a succession of faunistic manuals on Diptera (western Palearctic, Central American, and a forthcoming Afrotropical manual) that have substantially fostered research on this large and economically important order, established a pattern for faunistic manuals, and stabilized morphological Received 17 March 2011. Accepted 21 April 2012. W.N. Mathis, 1 Department of Entomology, PO BOX 37012, MRC 169; Smithsonian Institution, Washington, District of Columbia 20013-7012, United States of America T. Zatwarnicki, Department of Biosystematics, University of Opole, ul. Oleska 22, 45-052 Opole, Poland 1 Corresponding author (e-mail: mathisw@si.edu). doi:10.4039/tce.2012.9 Can. Entomol. 144: 81 92 (2012)

82 Can. Entomol. Vol. 144, 2012 terminology. The morphological terminology is now essentially used universally in papers dealing with Diptera that are written in English. We name the new species described in this paper after J. Richard Vockeroth who coauthored the chapter on shore flies in the Nearctic Manual (Wirth et al. 1987). Materials and methods The descriptive terminology, with the exceptions noted in Mathis (1986) and Mathis and Zatwarnicki (1990a), follows McAlpine (1981). Because specimens are small, usually less than 3.5 mm in length, study and illustration of the male terminalia required use of a compound microscope. We have followed the terminology for most structures of the male terminalia that other workers in Ephydridae have used (references in Mathis 1986; Mathis and Zatwarnicki 1990a, 1990b), such as surstylus. Zatwarnicki (1996) suggested that the pre- and postsurstylus correspond with the pre- and postgonostylus and that the subepandrial sclerite is the same as the medandrium. The terminology for structures of the male terminalia is provided directly in Figures 1 4. Dissections of male and female genitalia and descriptions were performed following Clausen and Cook (1971) and Grimaldi (1987). Abdomens were removed with microforceps and macerated in a potassium hydroxide solution. Cleared genitalia were rinsed in distilled water and 70% ethanol and then transferred to glycerin for observation. If necessary for proper orientation, the specimen was transferred from glycerin to glycerin jelly. The glycerin jelly was heated, and the specimen appropriately oriented. Subsequent cooling immobilized the specimen. An abdomen was placed in a plastic microvial filled with glycerin and attached to the pin supporting the remainder of the insect from which it was removed. The species descriptions are composite and not based solely on the holotypes. Head and two venational ratios used in the descriptions are based on three specimens (largest, smallest, and one other): gena-to-eye ratio genal height (immediately below maximum eye height)/ eye height; costal vein ratio the straight line distance between the apices of R 2 1 3 and R 4 1 5 /distance between the apices of R 1 and R 2 1 3 ; M vein ratio the straight line distance along vein M between crossveins dm cu and r m/distance apicad of dm cu. Many specimens examined for this study are in the National Museum of Natural History, Smithsonian Institution, Washington, District of Columbia (USNM). We also borrowed and studied numerous specimens, especially primary types from the following museums: AM Australian Museum, Sydney, Australia (Drs. Daniel J. Bickel, David K. McAlpine; including collections formerly housed at the School of Public Health and Tropical Medicine, University of Sydney) ANSP Academy of Natural Sciences of Philadelphia, Pennsylvania, United States of America CAU China Agricultural University, Beijing, China CIH Commonwealth Institute of Health, University of Sydney, Sydney, Australia (formerly School of Public Health and Tropical Medicine, University of Sydney). Collections now in AM DEI Deutsches Entomologisches Institut, Müncheberg, Germany (Drs. Frank Menzel) NMW Naturhistorisches Museum, Wien, Austria (Dr. Peter Senhal) PTZ Private collection of Dr. Tadeusz Zatwarnicki, Opole, Poland RNM Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands (presently housed in the Netherlands Centre for Biodiversity; Mr. Rob de Vries) USNM Former United States National Museum, collections in the National Museum of Natural History, Smithsonian Institution, Washington, District of Columbia. Systematics Tribe Dryxini Zatwarnicki Dryxini Zatwarnicki 1992: 85. Type genus: Dryxo Robineau-Desvoidy 1830. Mathis and Zatwarnicki 1995: 115 127; 2002: 1 101. Discussion. In addition to the synapomorphies listed in the tribal diagnosis that indicate a sister-group relationship for Dryxini and Notiphilini (Mathis and Zatwarnicki 1995), Dryxini are further characterized by the following

Mathis and Zatwarnicki 83 autapomorphies that establish the monophyly of the tribe: (1) gena high (secondarily short in some species of Paralimna); (2) face wide, transversely arched, and generally projected anteriorly; (3) male terminalia with surstylus divided into a presurstylus (clasper) and postsurstylus (surstylus); presurstylus with apex angulate and bifurcate; (4) pregonite and postgonite reduced and/or lacking; (5) hypandrium connected basally with postsurstylus, not with epandrium. Key to genera of Dryxini 1 Notopleuron bearing a single large seta; presutural supra-alar seta lacking; mid- and hindfemora moderately long to very long, length subequal to that of abdomen...2 Notopleuron bearing two setae; presutural supra-alar seta usually present; lacking in Papuama Mathis and Zatwarnicki and one species of Oedenops Becker); mid- and hindfemora normally developed, length much shorter than that of abdomen....4 2 Ocellar seta present, although short, inserted slightly in front of anterior ocellus; reclinate fronto-orbital seta present; anepisternum bearing one well-developed seta along posterior margin; vein R 1 bare along dorsum; R stem vein lacking setulae; crossvein dm cu normally developed, nearly straight, forming acute inner angle with vein M (southern Afrotropical)... Corythophora Loew Ocellar seta lacking; reclinate fronto-orbital seta lacking; anepisternum bearing two to three thin, long, hair-like setae along posterior margin; vein R 1 bearing several setulae along dorsum; R stem vein basad of humeral crossvein bearing several pale thin setulae on ventral surface; crossvein dm cu moderately long to long, sinuous......3 3 Arista bearing seven to nine long, dorsal hairs; katepisternum lacking row of slender setae along dorsal margin and katepisternal seta reduced; crossvein dm cu shallowly sinuous, generally forming angle with adjacent margin of wing; mid- and hindfemora normally developed, much shorter than length of abdomen (India, Iran, Oman).... Omyxa Mathis and Zatwarnicki Arista bearing 12 or more long, dorsal hairs; katepisternum bearing a row of slender setae near dorsal margin and katepisternal seta usually well developed (secondarily reduced or absent in some species); crossvein dm cu sinuous, long, generally running parallel with adjacent margin of wing; mid- and hindfemora elongate, subequal to length of abdomen (Afrotropical, Australian, Oriental).. Dryxo Robineau-Desvoidy 4 Katepisternal seta absent or very weakly developed..... 5 Katepisternal seta present, usually well developed (sometimes pale)...... 6 5 Arista bearing three to five dorsal rays (Afrotropical, Australian [Queensland], Nearctic [southern], Neotropical, Oriental, Palearctic [Egypt, Israel, Japan])... Oedenops Becker Arista bearing eight or more dorsal rays (Australasian, Oriental).. Papuama Mathis and Zatwarnicki 6 Two to three long facial setae, length subequal to combined length of pedicel and basal flagellomere, if two setae, these well separated, dorsal seta at about midheight, ventral seta closer to oral margin than to dorsal seta. R stem vein usually bearing one to three setulae on dorsum (usually two; lacking in Oedenopiforma javana (Wulp)) (Afrotropical, Australian, Oriental)...... Oedenopiforma Cogan One long facial seta (if longer setae present they are not as long or as separated as above, usually arranged in a somewhat vertical series of short setulae). R stem vein lacking setulae... 7 7 Forefemur lacking row of closely set, very short, somewhat blunt, toothlike spines along anteroventral surface; anterior proclinate fronto-orbital seta larger than posterior seta; face, gena, anepisternum, anterior surface of tibiae, and basolateral surface of scutellum not silvery microtomentose as in combination below; forefemur lacking long setae as below (pantropical with occasional extensions into temperate)..... Paralimna Loew Forefemur bearing anteroventral row of very short, stout, toothlike setae; proclinate fronto-orbital setae greatly reduced and subequal, setulalike; face, gena, anepisternum, anterior surface of tibiae, and basolateral surface of scutellum densely, silvery microtomentose; forefemur bearing four to five long setae along apical half of posteroventral surface, length approaching twice width of femur (Afrotropical).... Afrolimna Cogan

84 Can. Entomol. Vol. 144, 2012 Oedenopiforma Cogan Oedenopiforma Cogan 1968: 319 (as a subgenus of Paralimna; type species: Paralimna madecassa Giordani Soika 1956, original designation). Cogan 1980: 663 (as a subgenus of Paralimna). Mathis and Zatwarnicki 1995: 117 (accorded genus status); 2002: 42 51. Diagnosis This genus is distinguished from other Dryxini by the following combination of characters: small to moderately small shore flies, body length 1.65 2.85 mm; densely microtomentose, gray to brown dorsally, usually becoming darker ventrally. Description Head. Proclinate fronto-orbital setae greatly reduced and subequal, setulalike. Arista bearing eight or more dorsal rays. Facial setae two to three, if two, these well spaced, dorsal seta at about midheight, ventral closer to oral margin than to dorsal seta. Thorax. Dorsocentral setae 4 (1 1 3), anterior seta well developed; presutural supra-alar seta present; notopleuron bearing two large setae (anterior notopleural seta well developed); anepisternal and katepisternal setae well developed. R stem vein bearing one to three (usually two; lacking in O. javana) setulae on dorsum. Forefemur bearing anteroventral row of very short, stout, spinelike setae (difficult to see) along apical half (lacking in O. javana); mid- and hindfemora normally developed, much shorter than length of abdomen. Abdomen. Lacking pattern of fascia. Male terminalia: epandrium in posterior view, a broad to moderately narrow inverted U, bearing setulae toward posterior margin; cercus ovoid, dorsum slightly narrower in posterior view than venter, bearing numerous setulae, especially on medial portion; presurstylus in posterior view wider than high, bearing few to many short to long setulae; postsurstylus slightly curved anteriorly in lateral view, base wider than apex, especially in lateral view, with preapical notch, bearing setulae in patches along inner curvature; subepandrial sclerite present, sometimes with separate, lateral structures; aedeagus simple, usually wider basally, apex pointed to truncate; phallapodeme with conspicuous, moderately deep to deep keel; gonite apparently lacking; hypandrium wider than long (best seen in posterior view), shallowly and sometimes irregularly curved in lateral view. Distribution Australasian/Oceanian: Australia (New South Wales, Northern Territories, Western Australia). Afrotropical: Kenya, Madagascar, Namibia, Nigeria, Rwanda, South Africa (Transvaal), Sudan, Zaire. Oriental: Bangladesh, Brunei, China (Hainan, Yunnan), India (Haryana, Karnataka, Rajasthan), Indonesia (Java), Sri Lanka, Taiwan (Tainan), Viet Nam. Palearctic: Iran, United Arab Emirates. Discussion Oedenopiforma is characterized by several synapomorphies that confirm the monophyly of the genus. Among derived character states that we have discovered are the following: (1) face bearing two to three long facial setae, length subequal to combined length of pedicel and basal flagellomere; (2) forefemur bearing comblike row of short, stout, spinelike setulae along apical half of anteroventral surface (lacking in O. javana); (3) R stem vein bearing one to three (usually two) setulae (lacking in O. javana); (4) abdomen mostly lacking a distinctive pattern of transverse fascia. Key to species of Oedenopiforma 1 Facial series with three setae; paravertical setae greatly reduced or lacking (Australia)... Oedenopiforma uniseta (Malloch) Facial series with two setae; paravertical setae moderately to well developed... 2 2 Facial coloration sexually dimorphic; face of female whitish gray, that of male yellowish tan to golden or dark brown. Forefemur bearing peg-like row of short setulae along apical half of anteroventral surface (inconspicuous in male of Oedenopiforma vockerothi sp. nov.)...3 Facial coloration the same in males and females, mostly gray but with some brownish areas. Forefemur not bearing peg-like row of short setulae along apical half of anteroventral surface... Oedenopiforma javana (Wulp)

Mathis and Zatwarnicki 85 3 Face and antenna of male appearing velvety, densely microtomentose, dark brown (Madagascar, South Africa, Sudan)...... Oedenopiforma madecassa (Giordani Soika) Face of male yellowish tan to golden but not velvety, dark brown...4 4 Anepisternum gray; postsurstylus in lateral view elbowed medially (Fig. 4), distinctly curved; basal aedeagal opening in lateral view shallow.... Oedenopiforma argentea (Cogan) Anepisternum mostly brown, similar but slightly lighter in color than mesonotal coloration; postsurstylus in lateral view nearly straight (Fig. 15), not distinctly curved; basal aedeagal opening in lateral view distinctly concave, deeply excavated... Oedenopiforma vockerothi sp. nov. Most congeners have the combination of synapomorphies noted previously and their congeneric status is well documented. An exception is O. javana, which lacks synapomorphies two and three. Oedenopiforma argentea (Cogan) (Figs. 1 4) Paralimna (Oedenopiforma) argentea Cogan 1968: 322; 1980: 663. Oedenopiforma argentea. Mathis and Zatwarnicki 1995: 117 (genus recombination); 2002: 45 47. Distribution Afrotropical: Kenya, Namibia, Nigeria, Rwanda, South Africa (Transvaal), Zaire. Remarks The distribution of this species currently includes only localities in the Afrotropics. Previous records from Iran (Mathis and Zatwarnicki 2002) are probably not this species. Based on structures of the male terminalia (Figs. 1 4), especially the shape of the postsurstylus (Fig. 4), this is the sister species of O. vockerothi. Externally, these two species are also similar, especially the sexually dimorphic facial coloration (female whitish gray, male yellowish tan) and the peg-like row of short setulae on the apical half of the anteroventral surface of the forefemur. The reporting here of the sexually dimorphic facial coloration for this species corrects an error in our previous key and description of this species (Mathis and Zatwarnicki 2002), wherein the face of males and females was incorrectly stated to be the same. Oedenopiforma javana (Wulp), comb. nov. (Figs. 5 8) Paralimna javana Wulp 1892: 215. Cogan and Wirth 1977: 334. Bock 1988: 891 893. Mathis and Zatwarnicki 1995: 121. Paralimna biseta Hendel 1914: 105. Hennig 1941: 160 (synonymy). Paralimna atrimana Malloch 1925: 326. Bock 1988: 891 (synonymy). Oedenopiforma orientalis Zhang et al. 2009: 201. syn. nov. Diagnosis This species is distinguished from congeners by the following combination of characters: moderately small shore flies, body length 2.85 3.30 mm, wing length 2.50 3.00 mm. Description Head. Black; frons rusty brown microtomentose except ocellar triangle and fronto-orbits bluish gray; face yellowish gray microtomentose, area between antennae with bluish gray microtomentum, but with a narrow brown median stripe; antenna black with brownish yellow microtomentum. One large lateral vertical seta; one large medial vertical seta; paravertical setae well developed, about 1/2 length of medial vertical seta; one pair of large ocellar setae; one large, reclinate fronto-orbital, proclinate frontoorbital seta greatly reduced and subequal, hair-like; face with two large setae; gena with one large seta. Gena-to-eye ratio 0.23 0.33. Thorax. Black with bluish gray microtomentum, base of setae with brown spots. Legs black with bluish gray microtomentum except tarsomeres one to two brownish yellow and tarsomeres three to five

86 Can. Entomol. Vol. 144, 2012 Figs. 1 4. Male terminalia of Oedinopiforma argentea (Rwanda, Kisenyi): 1. epandrium, cerci, and presurstyli, posterior view; 2. same, lateral view; 3. internal male terminalia (aedeagus, phallapodeme, postsurstylus, subepandrium sclerite, hypandrium), ventral view; 4. same, lateral view. Scale bar 5 0.1 mm. brown. Four strong dorsocentral setae (1 1 3); one large presutural supra-alar seta; one large postpronotal seta; two large notopleural setae; anepisternum bearing one large seta, two slightly small setae and some small setulae along posterior margin; katepisternal seta well developed; two pairs of large scutellar seta. Forefemur with row of long, fine, pale anteroventral setae, these about as long as width of forefemur; midfemur with three strong anteroventral setae at apex. Wing light yellow; R stem vein without setulae dorsally along posterior margin; costal vein ratio 0.33 0.35; M vein ratio 1.00 1.06. Abdomen. Male genitalia (Figs. 5 8): epandrium in posterior view (Fig. 5) wider than high, ventrally extended lateral arms broad, in lateral view (Fig. 6) subrectangular; cercus in posterior view (Fig. 5) elongate, narrowly subtriangular to subrectangular, with dense setae, especially medially, truncate ventrally, in lateral view (Fig. 6) elliptical to lenticular; presurstylus robustly developed, in posterior view (Fig. 5) height slightly

Mathis and Zatwarnicki 87 Figs. 5 8. Male terminalia of Oedinopiforma javana (Taiwan, Tainan): 5. epandrium, cerci, and presurstyli, posterior view; 6. same, lateral view; 7. internal male terminalia, ventral view; 8. same, lateral view. Scale bar 5 0.1 mm. 䉷 2012 Entomological Society of Canada

88 Can. Entomol. Vol. 144, 2012 more than twice width, basal half over twice width of apical half, medial margin shallowly sinuous, basomedial arm of presurstlyus narrow, bearing row of setae along ventral margin, ventral apex of presurstylus pointed, in lateral view (Fig. 6) widest ventrally, about 33 higher than wide, ventral margin almost truncate, very shallowly arched; postsurstylus wider at middle in lateral view (Fig. 8) with a short, anterior, setose projection, apex moderatelly pointed, slightly curved, bifurcate (Fig. 7); aedeagus in lateral view (Fig. 8) with anterior margin rounded, posterior margin nearly straight, posteroventral portion truncate, anteroventral portion round and darkened, in ventral view (Fig. 7) subrectangular, very slightly wider apically, basal margin almost truncate, apical margin rounded; phallapodeme in lateral view (Fig. 8) with keel wide and moderately shallow, truncate, apical process longer than basal process in lateral view; subepandrial sclerite as two separate, lateroventral bars; hypandrium in ventral view (Fig. 7) almost twice as wide as long, anterior margin very broadly V-shaped, without short projection externally toward anterior margin, posterior margin medially concave, nearly flat, both posterior apices oriented medially, in lateral view (Fig. 8) pocket-shaped, rounded. Type material The holotype male of Paralimna javana (Indonesia. Java, in RNM) is apparently lost (Bock 1988) and a neotype should be designated. We are unaware, however, of any good specimens from Indonesia for such a designation. We have examined a headless male (in RNM) from Indonesia that is labeled Semarang Java VIII 05 [August 1905] Jacobson [handwritten]/paralimna javana det. Dr. [perhaps De ] Meijere [handwritten] but because of its condition, we are hesitant to designate it as a neotype. We await the collection of a conspecific male from Indonesia for designation as a neotype. The holotype male of Paralimna atrimana is from Australia, New South Wales: Belaringar, CIH (now in AM). The holotype male of Oedenopiforma orientalis is from China, Hainan, Danzhou, Nadalianyuan (19831 0 N, 109835 0 E),May14,2006,GangYao(in CAU). Six male paratypes bear the same label data as the holotype (CAU, USNM). Other paratypes are as follows: CHINA. Hainan: Haikou, Jiarihaitan (20802 0 N, 110820 0 E), June 9, 2006, Hui Dong (1~; CAU); Jianfengling, Nantianchi (18843 0 N, 108850 0 E), May 22, 2006, Gang Yao (1#; CAU). Yunnan: Jinghong, Jingshatan (22801 0 N, 100848 0 E), April 29, 2005, Xingyue Liu (1#; CAU). Other specimens examined Oriental: INDIA. Karnataka: Madikeri (22 km E; 12826 0 N, 75853.5 0 E), November 26, 2003, I. Yarom (1~; USNM). Rajasthan: Bap Lake, near Phalodi (26843.7 0 N, 73855.3 0 E), November 19, 2002, A. Freidberg (2#, 3~; USNM); Jaisalmer (26854.8 0 N, 70854.8 0 E; lake), November 20, 2002, A. Freidberg (1#; USNM); Nagda Temple (25 km N Udaipur; 26855.5 0 N, 73846.9 0 E; lake), November 22, 2002, A. Freidberg (1~; USNM). SRI LANKA. Anuradhapura: Hunuwilagama, near Wilpattu (8826.5 0 N, 79859.4 0 E; black light; 61 m), October 28 November 3, 1976, G.F. Hevel, S. Karunaratne (2~; USNM). Puttalam: Rajakadaluwa (7838.6 0 N, 79849.5 0 E), August 25, 1953, F. Keiser (USNM). Trincomalee: Kantalai (8822.1 0 N, 8181 0 E), November 12, 1953, F. Keiser (2~; USNM). TAIWAN. Tainan (2380.4 0 N, 120810.9 0 E), November 1909, H. Sauter (2#; ANSP). VIET NAM. Long Binh (10856.5 0 N, 106853.8 0 E; ditch bank), March November 1969, W.H. Pierce (2#, 1~; USNM). Distribution Australasian/Oceanian: Australia (New South Wales, Northern Territory, Queensland, Western Australia). Oriental: Bangladesh, Brunei, China (Hainan, Yunnan), India (Haryana, Karnataka, Rajasthan), Indonesia (Java), Sri Lanka, Taiwan (Tainan), Viet Nam. Remarks This species is somewhat similar and apparently related to O. uniseta (Malloch). Both species, for example, (1) lack a subepandrial sclerite and (2) have an elongate presurstylus that abuts dorsomedially and has a row of setulae basomedially. This species can be distinguished from O. uniseta by the following combination of characters: (1) two large facial setae, (2) paravertical setae well developed, (3) basomedial arm of the presurstylus narrow, (4) apex of presurstylus moderately narrowly pointed, and

Mathis and Zatwarnicki 89 Figs. 9 11. Oedinopiforma vockerothi (United Arab Emirates, Hatta Dam): 9. head, anterior view; 10. same, lateral view; 11. thorax, dorsal view. Scale bar 5 0.5 mm. (5) hypandrium lacking a short projection externally toward the anterior margin. Specimens of O. uniseta (Malloch) have three large facial setae, greatly reduced paravertical setae (or they arelacking),awidebasomedialarmofthepresurstylus, a rounded apex of the presurstylus, and a short projection of the hypandrium that extends externally toward the anterior margin. We also noted that the R stem vein of this species does not bear setulae on the dorsal surface, unlike congeners. The presence of these setulae on the R stem vein has been considered a synapomorphy for the genus (Mathis and Zatwarnicki 2002), and the lack of them in this species is either (1) a character reversal or (2) this species is possibly the sister group to the remaining congeners. The new combination for this species and the new synonymy presented above were initially overlooked (Mathis and Zatwarnicki 1995, 2002; Zhang et al. 2009) in part because our concept of this species was based on Bock s illustration (1988), which differs from that shown by a standard preparation of a typical specimen of O. javana (presurstylus almost parallel sided in illustration). We have now studied specimens, including structures from dissected male terminalia, from throughout the Oriental Region and have found that this is apparently a relatively

90 Can. Entomol. Vol. 144, 2012 Figs. 12 15. Male terminalia of Oedinopiforma vockerothi (United Arab Emirates, Hatta Dam): 12. epandrium, cerci, and presurstyli, posterior view; 13. same, lateral view; 14. internal male terminalia (aedeagus, phallapodeme, postsurstylus, subepandrium sclerite, hypandrium), ventral view; 15. same, lateral view. Scale bar 5 0.1 mm. common, widespread species in the Australasian and Oriental Regions. Oedenopiforma vockerothi sp. nov. (Figs. 9 15) Diagnosis This species is distinguished from congeners by the following combination of characters: small to moderately small shore flies, body length 1.80 2.85 mm, wing length 1.95 2.25 mm. Description Head (Figs. 9 10). Paravertical seta moderately well developed, about 1/3 length of medial vertical seta. Frons brown across vertex and usually extended to ocellar triangle, anterior portion of frons whitish gray to gray. Antenna black; arista with six to nine dorsal rays. Face sexually

Mathis and Zatwarnicki 91 dimorphic; face of male yellowish to brownish gray, especially dorsally, densely microtomentose, but not appearing velvety; face of female whitish gray to gray; two pairs of facial setae. Gena-to-eye ratio 0.29 0.30. Thorax (Fig. 11). Mesonotum broadly brown to rusty brown medially, becoming grayer laterally through postpronotum and notopleural area; anepisternum concolorous with mesonotum or slightly lighter in coloration, otherwise pleural areas gray. Stem vein bearing two (rarely one or three) setulae dorsally along posterior margin, length of each about equal to width of vein at level of insertion; costal vein ratio 0.30 0.36; M vein ratio 0.95 0.97. Legs mostly black; tarsi blackish red ventrally; forefemur of female bearing row of tiny, toothlike setulae along apical half of anteroventral surface, male row inconspicuous, barely evident; forefemur of male lacking row of long, fine, pale, ventrally oriented setulae along anteroventral surface. Abdomen. Male terminalia (Figs. 12 15): epandrium in posterior view (Fig. 12) slightly wider than high, dorsal portion with numerous, evenly scattered setulae, ventrally extended arms broad (Fig. 12), in lateral view (Fig. 13) subrectangular, wider dorsally than ventrally; cercus in posterior view (Fig. 12) semihemispherical to nearly obclavate with rounded corners, narrowly pointed dorsally, ventral margin rounded, densely setulose, especially medially, in lateral view (Fig. 13) semihemispherical; presurstylus in posterior view (Fig. 12) with sclerotized portion small, bar-like, horizontal, in lateral view (Fig. 13) horizontal, bar-like, length twice height, setulose; postsurstylus in lateral view (Fig. 15) large, robust, long, moderately narrow apically, nearly straight, not distinctly curved, apical half nearly parallel sided, digitiform, bearing setulae along lateral margins and especially ventrobasally, base wide; in ventral view (Fig. 14) subrectangular, length 43 width, nearly parallel sided, apex bifurcate; aedeagus relatively simple, in lateral view (Fig. 15) slipper-like with dorsobasal portion broad and deeply excavate, apex broadly pointed, in ventral view (Fig. 14) with apical half parallel sided, apex broadly rounded, basal portion parallel sided; phallapodeme in lateral view (Fig. 15) with keel irregular, pointed, extended processes about equal in length, relatively short; subepandrial sclerite in ventral view (Fig. 14) transversely elongate, very narrow, nearly straight, only lateral apices enlarged, in lateral view (Fig. 15) very narrow and shallowly curved; hypandrium in ventral view (Fig. 14) a moderately wide U-shaped process, narrowly sclerotized around margins only, in lateral view (Fig. 15) shallowly pocket-like, anterior and posterior apices pointed. Type material The holotype male is labeled U. Arab Emirates. Hatta Dam (24847.7 0 N, 56806.3 0 E; 365 m), March 1, 2010[,] Wayne N. Mathis/ USNM ENT 00285980 [plastic bar code label]/ HOLOTYPE # Oedenopiforma vockerothi Mathis & Zatwarnicki, USNM [red]. Two male and one female paratypes bear the same label data as the holotype. Other paratypes are as follows: United Arab Emirates. Wadi Maidaq (25820.8 0 N, 56805.5 0 E; 390 m), March 8, 2010, T. Zatwarnicki (1~; PTZ); Wadi Shawkah (25806.3 0 N, 56802.8 0 E; 300 m; reservoir), February 27, 2010, W.N. Mathis, T. Zatwarnicki (3~; PTZ, USNM). Type locality United Arab Emirates. Hatta Dam (24847.7 0 N, 56806.3 0 E; 365 m). We sampled behind the dam at Fort Hatta, especially where there were some short, emergent plants around isolated pools. Distribution Palearctic: United Arab Emirates. Natural history All specimens from the type series were taken by sweeping close to the surface of muddy to sandy shores associated with small, natural, freshwater oases or reservoirs in otherwise very arid habitats. Etymology The species epithet, vockerothi, is a genitive patronym to honor J. Richard Vockeroth, who contributed to the chapter on Ephydridae (Wirth et al. 1987) in the precedent-setting Manual of Nearctic Diptera. Remarks We anticipate that this species will be found in other countries of the Middle East. For example, we would not be surprised to discover that specimens from Iran that were previously listed

92 Can. Entomol. Vol. 144, 2012 as O. argentea (Mathis and Zatwarnicki 2002) are this species. Acknowledgments We express our sincere thanks to our colleagues and their institutions listed previously who loaned specimens. Without their cooperation this study could not have been completed. For thoroughly reviewing and providing critical and insightful remarks, we thank A.G. Irwin, who also indicated to us the need for a neotype for O. javana. Recent field work in the United Arab Emirates (February March 2010) was done in conjunction with a survey of the arthropods of that country with financial support coming from H.H. Sheikh Tahnoon Bin Zayed Al Nahyan. The arthropod survey is managed by Antonius (Tony) van Harten, who encouraged and facilitated all aspects of our field work and is gratefully acknowledged and thanked. We gratefully thank Karolyn Darrow for expert preparation of the habitus illustrations (Figs. 9 11) of O. vockerothi. References Bock, I.R. 1988. The Australian species of Paralimna and Notiphila (Diptera: Ephydridae). Invertebrate Taxonomy, 2: 885 902. Clausen, P.J., and Cook, E.F. 1971. A revision of the Nearctic species of the tribe Parydrini (Diptera: Ephydridae). Memoirs of the American Entomological Society, 27: 1 150. Cogan, B.H. 1968. A revision of the Ethiopian species of the tribe Notiphilini (Diptera: Ephydridae). Bulletin of the British Museum (Natural History), Entomology, 21: 281 365. Cogan, B.H. 1980. Family Ephydridae. In Catalogue of the Diptera of the Afrotropical Region. Edited by R.W. Crosskey. British Museum (Natural History), London. pp. 655 669. Cogan, B.H., and Wirth, W.W. 1977. Family Ephydridae. In A catalogue of the Diptera of the Oriental Region, vol. III. Suborder Cyclorrhapha (excluding Division Aschiza). Edited by M.D. Delfinado and D.E. Hardy. University Press of Hawaii, Honolulu. pp. 321 339. Giordani Soika, A.G. 1956. Diagnosi preliminari di nuovi Ephydridae e Canaceidae della Regione etiopica e del Madagascar (Diptera). Bollettino del Museo civico di Storia naturale di Venezia, 9: 123 130. Grimaldi, D.A. 1987. Phylogenetics and taxonomy of Zygothrica. Bulletin of the American Museum of Natural History, 186: 103 268. Hendel, F. 1914. Acalyptrate Musciden (Dipt.) III. In H. Sauter s Formosa-Ausbeute. Supplementa Entomologica, 3: 90 117. Hennig, W. 1941. Verzeichnis der Dipteren von Formosa. Entomologische Beihefte aus Berlin- Dahlem, 8: 1 2391iv. Malloch, J.R. 1925. Notes on Australian Diptera. No. vii. Proceedings of the Linnean Society of New South Wales, 50: 311 340. Mathis, W.N. 1986. Studies of Psilopinae (Diptera: Ephydridae), I: a revision of the shore fly genus Placopsidella Kertész. Smithsonian Contributions to Zoology, 430: 1 30. Mathis, W.N., and Zatwarnicki, T. 1990a. A revision of the Western Palearctic species of Athyroglossa (Diptera: Ephydridae). Transactions of the American Entomological Society, 116(1), 103 133. Mathis, W.N., and Zatwarnicki, T. 1990b. Taxonomic notes on Ephydridae (Diptera). Proceedings of the Biological Society of Washington, 103: 891 906. Mathis, W.N., and Zatwarnicki, T. 1995. World catalog of shore flies (Diptera: Ephydridae). Memoirs on Entomology, International, 4: 1 423. Mathis, W.N., and Zatwarnicki, T. 2002. A phylogenetic study of the tribe Dryxini Zatwarnicki (Diptera: Ephydridae). Smithsonian Contributions to Zoology, 617: 1 101. McAlpine, J.F. 1981. Morphology and terminologyadults. In Manual of Nearctic Diptera, vol. 1. Coordinated by J.F. McAlpine, B.V. Peterson, G.E. Shewell, H.J. Teskey, J.R. Vockeroth, and D.M. Wood. Research Branch Agriculture Canada, Monograph No. 27. pp. 9 63. Robineau-Desvoidy, J.B. 1830. Essai sur les Myodaires. Mémoires Preséntes par divers Savans a l Académie Royale des Sciences de l Institut de France, et Imprimés par son Ordre Sciences Mathématiques et Physiques 2: pp. 1 813. Wirth, W.W., Mathis, W.N., and Vockeroth, J.R. 1987. Ephydridae. In Manual of Nearctic Diptera, vol. 2. Coordinated by J.F. McAlpine, B.V. Peterson, G.E.Shewell,H.J.Teskey,J.R.Vockeroth,andD.M. Wood. Research Branch Agriculture Canada, Monograph No. 28. pp. 1027 1047. Wulp, F.M. van der 1892. Eenige Uitlandsche Diptera. Tijdschrift voor Entomologie, 34: 193 218. Zatwarnicki, T. 1992. A new classification of Ephydridae based on phylogenetic reconstruction (Diptera: Cyclorrhapha). Genus, 3: 65 119. Zatwarnicki, T. 1996. A new reconstruction of the origin of eremoneuran hypopygium and its implications for classification (Insecta: Diptera). Genus, 7: 103 175. Zhang, J., Yang, D., and Mathis, W.N. 2009. A new species of the shore-fly genus Oedenopiforma Cogan from the Oriental Region, with an updated key to the species (Diptera: Ephydridae). Proceedings of the Entomological Society of Washington, 111: 199 203.