Review of the Genus Tethina Haliday (Diptera: Canacidae: Tethininae) From Western North America

George A. Foster; Wayne N. Mathis

All genera and species of the family Canacidae as well as all synonyms and the world distribution for each species are listed to form an updated world catalog. Since McAlpine’s (2007) placement of the families Canacidae sensu stricto and Tethinidae into a single, inclusive family (Canacidae sensu lato, i.e. the older family-group name), a comprehensive world catalog has been needed to include the new taxonomic arrangement and the corpus of new entries published over the last fifteen years, that is since the preceding catalogs (Mathis, 1992; Mathis and Munari, 1996). Identification keys to all supraspecific taxa are also given for each taxonomic section.

Review of the Genus Tethina Haliday (Diptera: Canacidae: Tethininae) From Western North America Author(s): George A. Foster and Wayne N. Mathis Source: Proceedings of the Entomological Society of Washington, 110(2):300-330. 2008. Published By: Entomological Society of Washington DOI: http://dx.doi.org/10.4289/07-059.1 URL: http://www.bioone.org/doi/full/10.4289/07-059.1 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. PROC. ENTOMOL. SOC. WASH. 110(2), 2008, pp. 300–330 REVIEW OF THE GENUS TETHINA HALIDAY (DIPTERA: CANACIDAE: TETHININAE) FROM WESTERN NORTH AMERICA GEORGE A. FOSTER AND WAYNE N. MATHIS Department of Entomology, P.O. Box 37012, MRC 169, Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A. (e-mail: foster@alpha-mos.com and mathisw@si.edu) Abstract.—Ten species of the genus Tethina Haliday from western North America are reviewed with an emphasis on structures of the male terminalia, which are illustrated. One new species (type locality in parenthesis) is described: Tethina sasakawai (California. San Diego: Camp Pendleton; 33u13.39N, 117u24.59W, beach). Tethina thula Sasakawa, which was described from specimens collected in Japan, is reported from Alaska. Four new synonymies are proposed: Rhicnoessa denudata Melander and Rhicnoessa lavendula Melander 5 Rhicnoessa angustipennis Melander; Tethina woodi Foster and Tethina steyskali Foster 5 Rhicnoessa milichioides Melander. Lectotypes are designated for the following five species’ names: Tethina angustifrons Melander, Rhicnoessa angustipennis Melander, R. denudata Melander, R. lavendula Melander, and R. horripilans Melander. All species for which males are known are illustrated and distribution maps are provided. Key Words: Tethina Haliday, Canacidae (Tethininae), beach flies, western North America Recent collecting in western North America has resulted in the discovery of an undescribed species of Tethina from southern California and in the first records of T. thula Sasakawa from the Nearctic Region (Alaska). These discoveries prompted this review of all western Nearctic species of the genus, especially since the last study of this fauna was published over fifty years ago (Melander 1952) and is now woefully out of date and inadequate, given numerous taxonomic changes in the interim. In addition, we are providing new data, synonymies, comments, and illustrations for several species that Melander described. Many of Melander’s species are difficult to identify because his descriptions are vague and no study, description, or illustrations of structures of the male terminalia were included. A revised key to species is also provided. For completeness and uniformity, we are including a revised diagnosis for all species being treated. The new species from southern California is fully described and illustrated. MATERIALS AND METHODS The descriptive terminology for external structures and many internal structures follows that published in the Manual of Nearctic Diptera (McAlpine 1981). For structures of the male terminalia, however, we have adopted the terminology that Cumming et al. (1995) VOLUME 110, NUMBER 2 have suggested. Because specimens are small, usually less than 4 mm in length, study and illustration required use of dissecting and compound microscopes. The terminology for structures of the male terminalia is labeled on Figs. 3–4 and is not repeated on comparable illustrations of other species. The description of the new species is based primarily on its holotype with variation being accounted for in the remarks section. Other species’ descriptions are composite, not based solely on holotypes, and paired structures are described in the singular except where the context makes this inappropriate. The gena-toeye ratio (ratio is the average of three specimens (the largest, smallest, and one other) is the genal height measured at the maximum eye height divided by the eye height. Label data from each specimen were recorded and listed alphabetically according to country, state or province, county, and specific locality, such as city. As available, date of collection, collector, sex, and specimen location were listed. Label data from holotype specimens were recorded exactly, and clarifying information, such as script style and label color, is enclosed within brackets. Distribution maps were made using ESRI ArcViewE GIS 3.2. Longitude and latitude coordinates were obtained for the locality where each specimen was collected and entered into a Microsoft ExcelE spreadsheet. If unavailable directly from specimen labels, longitude and latitude were estimated using gazetteers and maps to determine the geographical coordinates. Dissections of male and female terminalia and descriptions were performed using the methods of Clausen and Cook (1971) and Grimaldi (1987). Microforceps were used to remove abdomens, which were macerated in a potassium or sodium hydroxide solution. Cleared 301 structures were rinsed in 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. After cooling, the embedded specimen in glycerin jelly became immobilized. Abdomens were placed in an attached plastic microvial filled with glycerin and attached to the pin supporting the remainder of the insect from which it was removed. For freshly caught specimens, we recommend that the epandrium and associated structures of the male terminalia be teased open, thus allowing examination of these structures and identification of the species without need of dissection. Although many specimens examined for this study are in the National Museum of Natural History, Smithsonian Institution, Washington, D. C. (USNM), we also borrowed or studied numerous specimens that are deposited in the following museums: AMNH ANSP CAS CDFA CNC MCZ MZSP OMNH American Museum of Natural History, New York, New York Academy of Natural Sciences of Philadelphia, Pennsylvania California Academy of Sciences, San Francisco, California California Department of Food and Agriculture, Sacramento, California Canadian National Collection, Ottawa, Canada Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts Museu de Zoologia de São Paulo, Universidade de São Paulo, São Paulo, Brazil Osaka Museum of Natural History, Osaka, Japan 302 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SYSTEMATICS Genus Tethina Haliday Tethina Haliday, in Curtis 1837: 293 (as a subgenus of Opomyza; published in synonymy; first made available by use in Haliday 1838: 188). Type species: Opomyza (Tethina) illota Haliday 1838, by subsequent monotypy (Haliday 1838: 188).—Sturtevant 1923: 5–7 [discussion of synonymy, listing of Nearctic species].—Czerny 1928: 3 [revision].— Malloch 1934: 453 [revision Chilean species, discussion, key].—Melander 1952: 199 [revision Nearctic species].— Ardö 1957: 131 [citation, fauna of northern Europe].—Vockeroth 1965: 727–728 [Nearctic catalog]; 1987: 1075 [key].—Prado and Tavares 1966: 429– 431 [review, Brazilian species].—Foster 1976b: 2–3 [Neotropical catalog].— Thompson and Mathis 1981: 86 [citation, nomenclature].—Mathis and Munari 1996: 13–19 [world catalog].— Foster and Mathis 1998: 608–630 [revision of Caribbean and Gulf of Mexico species].—Sabrosky 1999: 32, 304 [citations, nomenclature]. Rhicnoessa Loew 1862: 174. Type species: Rhicnoessa cinerea Loew, by monotypy.—Loew 1865: 34–39 [revision].—Hendel 1902: 261–264 [systematics].—Williston 1908: 292, 296 [Fig. of head, key].—Collin 1911: 234 [probable synonymy with Tethina].—Malloch 1913: 147 [discussion, Fig. of head].—Melander 1913: 298 [key to Nearctic species]; 1952: 200 [revision of Nearctic species].—Hendel 1911: 41 [generic remarks]; 1917: 46 [synonymy in key]; 1934: 46 [references].—de Meijere 1928: 78 [discussion].—Curran 1934: 331 [key].—Munari 1990: 60–61 [status as a subgenus of Tethina].— Beschovski 1993: 104 [diagnosis, as a genus]; 1994: 18 [diagnosis, as a genus]. Phycomyza Melander 1952: 198. Type species: Rhicnoessa milichioides Melander, by original designation.—Vocker- oth 1965: 727 [Nearctic catalog].— Foster 1976a: 338 [synonymy]. Diagnosis.—Tethina is distinguished from other genera of the subfamily Tethininae by the following combination of characters: Head: Frons bearing some setulae in addition to larger setae; frontoorbital and orbital setae usually with similar orientation, mostly reclinate or lateroclinate; fronto-orbital setae 3–4; paravertical setae more or less convergent. Face with shiny tubercle above vibrissal pore present. Eye appearing bare, interfacetal setulae very sparse or lacking. Gena bare except for a ventral or nearly ventral row of setulae; gena high in many species gena-to-eye ratio 0.5–0.75. Palpus and proboscis usually normally developed; clypeus small, if exposed not protruding anteriad beyond oral margin. Thorax: Scutum with more or less numerous rows of coarse setulae arising from punctures; scutellar disc bare; postpronotum with 3 or more setae, ventral seta curved upward; acrostichal setulae in two or more complete or nearly complete rows; prescutellar acrostichal setae present. Wing with costa not spinose; vein A1 + CuA2 short, much shorter than discal cell; wing usually shorter, about twice as long as wide (less often 2.5–3.0 times); cell bm and discal cell distinct. Mid- and hindtibiae evenly setulose, lacking anterodorsal or posterodorsal setae. Abdomen: Tergites wider than long; tergite 6 well differentiated from short syntergosternite 7 + 8, the latter forming a dorsal pregenital sclerite. Male terminalia: Surstylus positioned at ventral margin of epandrium, usually broadly articulated externally with epandrium, internally with subepandrial sclerite; aedeagus usually very long and sinuous, either thick and straplike or narrow and ribbonlike; aedeagus micropubescent dorsally. Discussion.—Among genera of Tethininae, Tethina has by far the greatest number of species worldwide, with more VOLUME 110, NUMBER 2 than 60% of the described species in the subfamily (67 of 106; Mathis and Munari 1996, Munari 2002). Ten species are included in this study from western North America. The key, however, includes two additional species (T. pallipes (Loew) and T. xanthopoda (Williston)) that eventually may be found in the western Nearctic Region. KEY TO SPECIES OF TETHINA FROM THE WESTERN NEARCTIC REGION 1. Gena with a shiny band extended from face nearly to occiput . . . . . . T. pallipes (Loew) – Gena uniformly colored . . . . . . . . . . . . . 2 2. Apex of scutellum with distinct yellowish to reddish spot . . . . T. xanthopoda (Williston) – Scutellum uniformly gray or whitish microtomentose . . . . . . . . . . . . . . . . . . . . . . . 3 3. Gena less than 0.30 eye height, body rather slender, frail . . . . . . . . . . . . . . . . . . . . . . 4 – Gena 0.35 to 0.66 eye height, if frail bodied, gena is at least 0.35 eye height . . . . . . . . . 5 4. Head distinctly prognathous, labrum and labellum extended . . T. angustifrons Melander – Head normal, not prognathous . . . . . . . . . . . . . . . . . . . T. angustipennis (Melander) 5. Head distinctly prognathous, labrum and labellum extended, face short . . . . . . . . . . 6 – Head as high as or higher than long, mouthparts and face normal . . . . . . . . . . 7 6. Foretibia yellowish, female cerci with short, thick setae . . . . . . T. prognatha (Melander) – Foretibia gray, female cerci with only weak setulae . . . . . . . . T. milichioides (Melander) 7. Body light gray to whitish, most setae and setulae white, wings may also be whitish with brown veins . . . . . . . . . . . . . . . . . . 8 – Body darker gray microtomentose, all setae and setulae black, wings hyaline . . . . . . . 9 8. Surstylus curved in lateral view, somewhat rounded distally . . . . . . . . T. albula (Loew) – Surstylus straight in lateral view, pointed distally . . . . . . . . . T. willistoni (Melander) 9. Gena at least 0.66 eye height, body dark gray, robust, heavily setose . . . . . . . . . . . . . . . . . . . . . . . . T. horripilans (Melander) – Gena less than 0.50 (usually 0.40–0.44) eye height, body of medium weight . . . . . . . . 9 10. Tibiae uniformly yellow T. spinulosa (Cole) – Tibiae uniformly gray . . . . . . . . . . . . . . 10 11. Posterior margin of surstylus broadly rounded as in Fig. 24–25 (California) . . . . . . . . . . . . . . . . . T. sasakawai, new species – Posterior margin of surstylus as in Fig. 31– 32 (Alaska) . . . . . . . . . . T. thula Sasakawa 303 Tethina albula (Loew) (Figs. 1–4, 7) Rhicnoessa albula Loew 1869: 44 [United States. Rhode Island. Newport: Newport (41u29.49N, 71u18.89W); ST male/ female, MCZ].—Johnson 1910: 812 [citation]; 1913: 89 [citation]; 1930: 156 [citation].—Malloch 1913: 147 [citation].—Melander 1913: 298 [key]; 1952: 201–202 [key, citation].—Frey 1919: 15 [citation].—Hendel 1934: 43 [key], 46–47 [citation].—Hennig 1937: 140 [citation]. Tethina albula: Sturtevant 1923: 6 [generic combination].—Johnson 1925: 286 [citation].—Curran 1934: 330 [citation].—Vockeroth 1965: 727 [Nearctic catalog].—Prado and Tavares 1966: 431–432 [revision, Figs. of - terminalia].—Foster 1976b: 2 [Neotropical catalog].—Mathis and Munari 1996: 14 [world catalog].—Foster and Mathis 1998: 609–611 [revision; Caribbean and Gulf of Mexico, Figs. of head and - terminalia].—Mathis and Foster 2007: 415–518 [review of species from Delmarva States]. Rhicnoessa sonorensis Melander 1952: 207 [Mexico. Baja California and Sonora: Rocky Point Marsh; LT (designated by Foster 1976b: 2), USNM].—Cole 1969: 387 [distribution, diagnosis]; Foster and Mathis 2000: 544 [synonymy, citation]. Tethina sonorensis.—Foster 1976b: 2 [generic combination, lectotype designation, Neotropical catalog].—Mathis and Munari 1996: 18 [world catalog]. Diagnosis.—This species is distinguished from congeners by the following combination of characters: Body length 1.60–3.15 mm; body with gray to whitish gray microtomentum; setae may be all white to some black. Head (Fig. 1): Antenna and frons yellow; gena variable but high, greater than 0.40 eye height, often at least 0.50. Thorax: Uniformly 304 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1–4. Tethina albula. 1, Head, lateral view. 2, Right surstylus, posterior view. 3, External male terminalia (epandrium, cerci, and surstyli), posterior view (surstylus more from a posterolateral view). 4, Same, lateral view. gray microtomentose, sometimes with a light brownish or golden cast; 4 irregular rows of acrostichal setulae; scutellum uniformly gray. Wing whitish with veins yellowish to brownish. Femora mostly gray, midfemur often more yellowish gray; hindfemur of male slightly more swollen than fore- and midfemur; tibiae mostly yellow, sometimes slightly grayish in middle third; foretarsus all yellow except apical tarsomere may be dark, midtarsus also may have slightly dark- ened apical tarsomere, apical tarsomere of hindtarsus brown to black. Abdomen: Gray microtomentose with extreme apices of sternites yellowish. Male terminalia (Figs. 2–4): epandrium concolorous with rest of abdomen, surstylus articulated with and broadly attached to epandrium, narrowly spatulate in posterior view (Fig. 2), length 2.53 width, apex rounded; medial margin bearing irregular row of sparse setulae along entire length, setulae moderately well VOLUME 110, NUMBER 2 developed; surstylus in lateral view (Fig. 4) narrow, curved anteroventrally; aedeagus thick, straplike, bearing dense velvety hairlike pubescence on dorsal surface. Female terminalia: cerci mostly yellow, brownish to gray apically, bearing only weak setulae. Type material.—The syntype series of Rhicnoessa albula Loew, labeled ‘‘Loew Collection’’ and comprising one male (only two legs and a left wing remain) and four females (one bearing a red ‘‘Type’’ label (13444); MCZ), does not allow for accurate and reliable identification of this species. Osten Sacken, however, collected and retained a male (head missing) from the type locality (Newport, Rhode Island) when he collected the type series. Osten Sacken’s practice was to retain a few specimens of species represented by a long series, sending the majority to Loew for description. That retained, headless male, which is presumably conspecific with the type series, was identified, dissected, and is the basis for our diagnosis of this species. The lectotype male of Rhicnoessa sonorensis Melander (designated by Foster 1976b: 2) is labeled ‘‘SonoraMEXICO RockyP[oin]tMarsh 21 April 947 [1947] A.L. Melander/ALMelander Collection 1961 [stippled green band on right side of label]/Lectotype Tethina sonorensis (Melander) George A. Foster 1974 [handwritten; black submarginal border].’’ The lectotype is double mounted (minuten in a rectangular card), is in excellent condition (abdomen removed and dissected, the structures in an attached microvial), and is deposited in the USNM. Specimens examined from the western Nearctic Region (Fig. 7).—MEXICO. BAJA CALIFORNIA SUR: Loreto (26u019N, 111u219W; Sefton Orca Expedition to Gulf of California), 29 Mar 1953, P. H. Arnaud (7-, 3U; CAS). SONORA: Puerto Peñasco (31u18.29N, 113u22.99W), 21 Apr 1948, A. L. Melan- 305 der (1-, 3U; USNM); Rocky Point (Puerto Peñasco; 31u199N, 113u329W; marsh), 21 Apr 1947, A. L. Melander (7-, 10U; CAS (1- labeled as a paratype of T. sonorensis Melander), USNM). Distribution.—Australasian/Oceanian: Hawaii (Hawaii, Kahoolawe, Kauai, Maui, Oahu). Nearctic: United States (California, Delaware, Florida, Massachusetts, Maryland, North Carolina, New York, South Carolina, Rhode Island, Virginia). Neotropical: Bahamas, Belize, Brazil (Rio de Janeiro), Costa Rica, Curaçao, Ecuador, Guyana, Mexico (Baja California Sur, Chiapas, Quintana Roo, Sonora), Panama, Peru, Trinidad, Tobago, Turks and Caicos, West Indies (Anguilla, Antigua, Barbados, Barbuda, Dominica, Dominican Republic, Grand Cayman, Grenada, Jamaica, Montserrat, Puerto Rico, St. Croix, St. Lucia, St. Vincent). Remarks.—Specimens of T. albula exhibit variation in setal color. In many specimens, particularly those from the eastern Nearctic Region, all setae and setulae are white. Specimens from the southern United States, the Caribbean and the specimens we’ve examined here from Mexico, however, vary in having one or two of the larger setae black. We have also observed variation in the coloration of the legs between specimens from northeastern United States and the Caribbean. The femora of Caribbean specimens may be almost entirely yellow while specimens from the Delmarva states and northward have completely gray femora. We also observed variation in the color of the basal flagellomere and the mesonotum in specimens from the Delmarva states. These external characters, therefore, like setal color, are not completely reliable. Tethina angustifrons Melander (Figs. 5–7) Tethina angustifrons Melander 1952: 199.—Vockeroth 1965: 727 [generic 306 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5–6. Tethina angustifrons. 5, External male terminalia (epandrium, cerci, and surstyli), posterior view (surstylus more from a posterolateral view). 6, Same, lateral view. combination; Nearctic catalog].—Cole 1969: 386 [distribution, diagnosis].— Mathis and Munari 1996: 14 [world catalog]. Diagnosis.—This species is distinguished from congeners by the following combination of characters: Body length 1.83–2.56 mm; body with light gray microtomentum; all setae and setulae black. Head: Antenna yellow; frons orange yellow; gena narrow, gena-toeye ratio about 0.25; labrum and labellum quite long, yellow; palpus yellow. Thorax: 2 irregular rows of acrostichal setulae; scutellum uniformly gray. Wings hyaline with yellowish veins. Legs entirely yellow. Abdomen: Gray to brown, subshiny to microtomentose, apices of sternites white. Male terminalia (Figs. 5– 6): epandrium concolorous with rest of abdomen, surstylus somewhat elongate and spatulate in lateral view, armed with short, peglike spicules on medial surface of apex, aedeagus thin ribbonlike (Figs. 5–6). Female terminalia: cerci light brown, rather narrow and strong, studded with short, thick setae and in dried material appearing to be fused into one cercus (when cleared in a solution of NaOH, however, the cerci are obviously not fused). Type material.—Melander’s (1952) type series included 17 specimens from Morro Dunes, Asilomar, and Pismo Beach, California, and were collected from July to October. We have examined all 17 specimens and have determined that one specimen from Morro Dunes is T. angustipennis not T. angustifrons. Since Melander did not designate a holotype, we herein designate as lectotype a - labeled ‘‘Morro Dunes [35u20.89N, 120u50.29W], CAL[IFORNIA. San Luis Obispo:] 6/9/45 [9 Jul 1945] ALMelander/ALMelander Collection 1961[right third of label with green stippling]/HOLOTYPE Tethina angustifrons Melander [red label]/LECTOTYPE - Tethina angustifrons Melander by Foster & Mathis [handwritten except for ‘‘LECTOTYPE’’ and ‘‘By’’; black VOLUME 110, NUMBER 2 Fig. 7. Region. 307 Distribution map of Tethina albula (squares) and T. angustifrons (dots) in the western Nearctic submarginal border]. The specimen is double mounted (minuten in a cardboard rectangle), is in excellent condition, and is deposited in the USNM. Paralectotypes are as follows: UNITED STATES: CALIFORNIA. Monterey: Asilomar (36u37.29N, 121u56.19W), 1 Sep 1945, A. L. Melander (1-, 1U; USNM). San Luis Obispo: Morro Dunes (35u20.89N, 120u50.29W), 6 Sep 1945, A. L. Melander (2-, 1U; USNM); Pismo Beach (35u08.69N, 120u38.59W), 29 Aug 1945, A. L. Melander (5-, 3U; USNM). Additional specimens examined from the western Nearctic Region (Fig. 7).— MEXICO. BAJA CALIFORNIA: Ensenada (31u51.59N, 116u389W), 24 Jun 1950, A. L. Melander (4U; USNM). UNITED STATES. CALIFORNIA. Monterey: Asilomar (36u37.29N, 121u 56.19W), 11 Jul 1957, A. L. Melander (3-; USNM). Orange: Pacific Grove (36u379N, 121u54.69W), 28 Jul 1940, A. L. Melander (1-; USNM). San Luis Obispo: Morro Bay (35u20.19N, 120u51.19W), 7 Oct 1946, A. L. Melander (1U; USNM); Pismo Beach (35u08.69N, 120u38.59W), 25 Sep 1946, A. L. Melander (1-; USNM). Distribution.—Nearctic: United States (California). Neotropical: Mexico (Baja California). Remarks.—Specimens of this species are small, rather delicate, and are easily recognized by their extended labrum and labellum and their moderately prognathous head. The legs are entirely yellow and the thorax is light gray microtomentose. There are only two sparse rows of acrostichal setulae. The abdomen is semi-shiny dark gray or brownish with the posterior margins of the sternites 308 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON white. Females have long, strong cerci that sometimes appear fused and are studded with short, thick setae. A similar species, T. angustipennis, is also small and delicate, but the head is not prognathous, the labrum and labellum are not elongated, and the thorax and abdomen are somewhat reddish instead of gray. Tethina angustifrons is apparently related to T. milichioides and is also similar to T. insulans Curran. Tethina angustipennis (Melander) (Figs. 8–10) Rhicnoessa angustipennis Melander 1952: 199.—Cole 1969: 387 [distribution, diagnosis]. Tethina angustipennis: Vockeroth 1965: 727 [generic combination; Nearctic catalog].—Mathis and Munari 1996: 14 [world catalog]. Rhicnoessa denudata Melander 1952: 204.—Cole 1969: 387 [distribution, diagnosis]. New synonym. Tethina denudata: Vockeroth 1965: 727 [generic combination; Nearctic catalog].—Mathis and Munari 1996: 15 [world catalog]. Rhicnoessa lavendula Melander 1952: 204.—Cole 1969: 387 [distribution, diagnosis]. New synonym. Tethina lavendula: Vockeroth 1965: 727 [generic combination, Nearctic catalog].—Mathis and Munari 1996: 17 [catalog]. Diagnosis.—This species is distinguished from congeners by the following combination of characters: Body length 1.94–2.49 mm; body brownish with light brown microtomentum; thorax light brown, abdomen darker. Head: Antenna yellow; anterior half of frons yellow remainder dark brown to black; three pairs of fronto-orbitals; gena white, narrow, gena-to-eye ratio approximately 0.28. Thorax: Color light brown to yellowish; 2 irregular and sparse rows of acrostichal setulae; scutellum uni- formly light brown. Legs entirely yellow. Abdomen: Dark brown microtomentose. Male terminalia (Figs. 8–9): epandrium shiny dark brown; male surstylus in lateral view widely spatulate, posteriorly curved , armed with short peglike spicules on medial surface, aedeagus thin ribbonlike (Figs. 8–9). Female terminalia: cerci brown, studded with short, but strong, sharp setae. Type material.—Melander (1952) reported two male and two female specimens of Rhicnoessa angustipennis from Morro Bay, California. The USNM collection only has one male and one female so labeled. Because Melander did not designate a holotype, we herein designate as lectotype the - labeled ‘‘Morro Dunes [35u20.89N, 120u50.29W], CAL[IFORNIA. San Luis Obispo:] 17/6/ 47 ALMelander/ALMelander Collection 1961[right third of label with green stippling]/HOLOTYPE Tethina angustipennis Melander [red label]/LECTOTYPE - Tethina angustipennis Melander by Foster and Mathis [handwritten except for ‘‘LECTOTYPE’’ and ‘‘By’’; black submarginal border]. The specimen is double mounted (minuten in a cardboard rectangle), is in excellent condition, and is deposited in the USNM. Paralectotypes are as follows: United States: California. San Luis Obispo: Morro Dunes (35u20.89N, 120u50.29W), 17 Jun 1947, A. L. Melander (1U; USNM). We have examined the type series of R. denudata Melander and have concluded that it is conspecific with, and thus a synonym of T. angustipennis based on both external characters and structures of the male and female terminalia. Melander (1952) stated that he had a series of 8 males and 18 females in his type series. We have found the original 8 males (plus another that is deposited in the CAS) but only 17 females. Melander must have mistaken one of the males for a female. He also wrote that the collection dates of his syntypes are ‘‘August VOLUME 110, NUMBER 2 309 Figs. 8–9. Tethina angustipennis. 8, External male terminalia (epandrium, cerci, and surstyli), posterior view (surstylus more from a posterolateral view). 9, Same, lateral view. 28, 1945, October 8, 1946, and June 15, 1947.’’ We have noted, however, that part of the series was actually collected on October 9, 1946, in addition to October 8, 1946. Since Melander did not designate a holotype, we herein designate as lectotype a male labeled ‘‘Carpenteria [34u23.99N, 119u31.19W], CAL[IFORNIA] 9/X/46 ALMelander/ ALMelander Collection 1961[right third of label with green stippling]/HOLOTYPE Tethina denudata Melander [red label]/LECTOTYPE - Tethina denudata Melander by Foster and Mathis [handwritten except for ‘‘LECTOTYPE’’ and ‘‘By’’; black submarginal border]. The specimen is double mounted (minuten in a cardboard rectangle), is in excellent condition, and is deposited in the USNM. Paralectotypes are as follows: UNITED STATES. CALIFORNIA. Santa Barbara: Carpinteria (34u23.99N, 119u31.19W), 15 Jun–9 Oct 1946, 1947, A. L. Melander (7-, 17U; USNM), 8 Oct 1946 (1-; CAS). Study of the Melander’s three syntypes of R. lavendula reveals that they are also conspecific with T. angustipennis. We dissected the male terminalia of one syntype and they are identical to those of T. angustipennis. In external characters, including the thick setulae of the female cerci, they are also identical. As Melander did not designate a holotype, we herein designate as lectotype a - labeled ‘‘Huntington B[ea]ch, CAL[IFORNIA] 4/6/45 ALMelander/ALMelander Collection 1961[right third of label with green stippling]/HOLOTYPE Tethina lavendula Melander [red label]/LECTOTYPE - Tethina lavendula Melander by Foster and Mathis [handwritten except for ‘‘LECTOTYPE’’ and ‘‘By’’; black submarginal border].’’ The specimen is double mounted (minuten in a cardboard rectangle), is in excellent condition, and is deposited in the USNM. Paralectotypes are as follows: UNITED STATES. CALIFORNIA. Orange: Balboa (33u35.29N, 117u549W), 15 Jul 1940, A. L. Melander 310 Fig. 10. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Distribution map of Tethina angustipennis in the western Nearctic Region. (1U; USNM); Huntington Beach (33u39.69N, 118uW), 4 Jun 1945, A. L. Melander (1-; USNM). Additional specimens examined from the western Nearctic Region (Fig. 10).— MEXICO. BAJA CALIFORNIA: Ensenada (31u51.59N, 116u389W), 24 Jun 1950, A. L. Melander (2U; USNM). UNITED STATES. CALIFORNIA. San Luis Obispo: Morro Bay (35u20.19N, 120u51.19W), 9 Aug 1950, A. L. Melander (1U; USNM); Morro Dunes (35u20.89N, 120u50.29W), 6 Sep 1945, A. L. Melander (1-, 1U; USNM). Santa Barbara: Carpinteria (34u23.99N, 119u31.19W), 12 Jun–23 Sep 1946, 1952, A. L. Melander (4U; USNM). Distribution.—Nearctic: United States (California). Neotropical: Mexico (Baja California). Remarks.—Specimens of this species are small and delicate and are distinctive in being light brown to dark yellowish in color, which is atypical among congeners. The male genitalia, especially in lateral view, closely resemble those of T. spinulosa. These two species are clearly distinct, however, based on the female cerci bearing short, thick setae in T. angustipennis while the female cerci of T. spinulosa bear fine setulae. Also, specimens of T. spinulosa, even those that are small and relatively frail, are heavily setulose over the entire body, while those of T. angustipennis generally have much weaker setae. Tethina horripilans (Melander) (Figs. 11–13) Rhicnoessa horripilans Melander 1952: 204.—Cole 1969: 387 [distribution, diagnosis] Tethina horripilans: Vockeroth 1965: 727 [generic combination; Nearctic cata- VOLUME 110, NUMBER 2 311 Figs. 11–12. Tethina horripilans. 11, External male terminalia (epandrium, cerci, and surstyli), posterior view (surstylus more from a posterolateral view). 12, Same, lateral view. log].—Vockeroth 1987: 1076–1077 [figs. of head, hindtibia, and wing].— Mathis and Munari 1996: 16 [world catalog]. Diagnosis.—This species is distinguished from congeners by the following combination of characters: Body length 2.64–4.61 mm; body gray with light gray microtomentum, males quite bristly in appearance. Head: Antenna dark orange gray; frons yellow orange, ocellar triangle and surrounding area gray microtomentose; 3–4 pairs of fronto-orbitals, anterior seta weak; gena white microtomentose, very high, gena-to-eye ratio nearly 0.66; palpus yellow, labellum and labrum dark brown to black. Thorax: Uniformly gray microtomentose, mesonotum may have a brownish cast; 4 irregular rows of strong acrostichal setae; scutellum gray. Coxae, femora and tibiae gray microtomentose, male with strongly swollen fore- and hindfemora, tarsi yellow except 5th tarsomere black. Abdomen: Uniformly gray micro- tomentose, extreme posterior edges of tergites white. Male terminalia (Figs. 11– 12): epandrium concolorous with rest of abdomen, surstylus robust, widely spatulate with a slight indentation ventrally in lateral view, armed with short, peglike spicules on medial surface of extreme apex, remainder of medial surface with longer setulae, aedeagus thin ribbonlike with micropubescence dorsally. Female terminalia: cerci grayish brown, with only weak setulae. Remarks.—Specimens of this species can be large, robust and quite bristly, especially males. The gena is the highest of all the west coast species of Tethina, being nearly 0.66 the eye height. Type material.—Melander (1952) noted that he had examined 170 specimens from Washington, Oregon, and California. The USNM collection has only 124 specimens from the Melander collection and the CAS has one, all of which have a yellow PARATYPE label, but only 87 specimens exactly match the dates he cited in his study (an additional three 312 Fig. 13. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Distribution map of Tethina horripilans in the western Nearctic Region. specimens were misidentified and are the new species described in this paper as T. sasakawai new species). We suspect that he simply did not mention an additional 37 specimens from the type locality, Ilwaco, Washington, that we include here in the paralectotype series. We here designate one of the male specimens of the syntypic series as lectotype. The lectotype is labeled ‘‘Ilwaco [46u18.69N, 124u02.69W], WASH[INGTON. Pacific:] July 1917 ALMelander/ALMelander Collection 1961[right third of label with green stippling]/HOLOTYPE Tethina horripilans Melander [red label]/LECTOTYPE - Tethina horripilans Melander by Foster and Mathis [handwritten except for ‘‘LECTOTYPE’’ and ‘‘By’’; black submarginal border]. The specimen is double mounted (minuten in a cardboard rectangle), is in excellent condition (abdomen removed, structures of male terminalia dissected and placed in an attached glycerin vial), and is deposited in the USNM. Paralectotypes are as follows: UNITED STATES. CALIFORNIA. San Francisco: San Francisco (37u48.69N, 122u21.19W), 22 Jun 1947, A. L. Melander (1-; USNM). OREGON. Lincoln: Waldport (44u25.69N, 124u04.19W), 13 Sep 1934, A. L. Melander (3-, 1U; USNM). WASHINGTON. Grays Harbor: Copalis (47u06.89N, 124u10.49W), 14 Aug–5 Sep 1921, 1934, A. L. Melander (35-, 16U; USNM). Jefferson: Kaloloch (47u36.39N, 124u22.49W), 5 Sep 1934, A. L. Melander (1-; USNM). Pacific: Ilwaco (46u18.69N, 124u02.69W), 25 May–6 Sep 1917, 1922, 1934, A. L. Melander (12-, 17U; CAS, USNM) 25 May–6 Sep 1917, 1922, 1934, A. L. VOLUME 110, NUMBER 2 Melander (12-, 25U; USNM); Long Beach (46u21.19N, 124u03.39W), 30 Aug 1921, A. L. Melander (1-; USNM). Additional specimens examined from the western Nearctic Region (Fig. 13).— UNITED STATES. WASHINGTON. Pacific: Grayland Beach State Park (46u47.39N, 124u05.69W), 28–29 Jun 1988, D. and W. N. Mathis (6-; USNM); Fort Canby State Park (46u17.59N, 124u04.39W), 29 Jun 1988, D. and W. N. Mathis (2-; USNM). OREGON. Lane: Heceta Head (44u08.19N, 124u07.49W; beach), 1 Aug 2005, D. and W. N. Mathis (12-, 4U; USNM); Florence (44u019N, 124u08.29W; beach), 1 Aug 2005, D. and W. N. Mathis (5-, 5U; USNM). Distribution.—Nearctic: United States (California, Oregon, Washington). Remarks.—Tethina horripilans is one of the most distinctive and readily identified species of Tethina in the Nearctic Region. The species’ extremely high gena to head ratio combined with its large size and robust, bristly habitus make it easy to distinguish from congeners. Tethina milichioides (Melander) (Figs. 14–23) Rhicnoessa milichioides Melander 1913: 299.—Hendel 1934: 43 [key], 48 [citation].—Melander 1952: 208 [citation]. Tethina milichioides: Sturtevant 1923: 6 [generic combination].—Foster 1976a: 345 [revision].—Mathis and Munari 1996: 17 [world catalog]. Phycomyza milichioides: Melander 1952: 198, 212 [generic combination, fig. of - terminalia].—Vockeroth 1965: 727 [Nearctic catalog].—Cole 1969: 386 [distribution, diagnosis]. Tethina woodi Foster 1976a: 342.— Mathis and Munari 1996: 19 [world catalog]. New synonym. Tethina steyskali Foster 1976a: 344.— Mathis and Munari 1996: 18 [world catalog]. New synonym. 313 Diagnosis.—This species is distinguished from congeners by the following combination of characters: Body length 1.40–3.00 mm; body gray microtomentose; setae all black. Head: (Figs. 20–22): Antenna and frons gray except extreme base of basal flagellomere yellowish; gena whitish to light brownish, narrow, gena-to-eye ratio varies from 0.27–0.47 generally in the 0.40 range. Thorax: Thorax uniformly gray microtomentose; 4 irregular rows of acrostichal setulae; scutellum uniformly gray. Wings slightly smoky hyaline with veins brownish. Femora uniformly gray microtomentose; fore- and hindfemora of both sexes clearly swollen; tibiae uniformly gray microtomentose; basitarsi yellow, remaining tarsal segments brown to black. Abdomen: Gray microtomentose with extreme apices of sternites whitish. Male terminalia (Figs. 14–19): epandrium concolorous with rest of abdomen, surstylus articulated with and broadly attached to epandrium, broadly spatulate in lateral view (Figs. 14–16); median margin bearing irregular row of sparse setulae along entire length, setulae moderately well developed; aedeagus thin, ribbon like, bearing dense velvety hairlike pubescence on dorsal surface; hypandrium as in Figs. 17–19. Female terminalia: cerci brown, with only weak setulae. Type material.—Melander (1913) described Rhicnoessa milichioides from three syntypes (two males and one of unknown sex since the abdomen is missing). Foster (1976a: 346) designated one of the males as the lectotype. It is labeled Seattle [47u36.49N, 122u19.99W] Wash[ington. King:], Aug[ust] 2, [19]08 [white label, the date is handwritten]/ ALMelander Collection 1961[right third of label with green stippling]/Lectotype Phycomyza milichioides (Mel.) G.A. Foster 1975 [top third of label in red]. It is in excellent condition (abdomen removed, structures of male terminalia dissected and placed in an attached 314 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 14–22. Tethina milichioides. 14, External male terminalia, lateral view (Washington: Ilwaco). 15, Same (California: Pismo Beach). 16, Same (Washington: Seattle). 17, Hypandrium, ventral view (Washington: Ilwaco). 18, Same (California: Pismo Beach). 19, Same (Washington: Seattle). 20, Head, lateral view (Washington: Ilwaco). 21, Same (California: Pismo Beach). 22, Same (Washington: Seattle). glycerin vial), and is deposited in the USNM. Paralectotypes (1-, 1? (abdomen removed); USNM) are labeled with an identical locality and date label and A. L. Melander collection label. The holotype male of Tethina woodi Foster is labeled ‘‘Ilwaco[,] WASH[ington. Pacific:] 12 Jul [19]22 ALMelander/ ALMelander Collection 1961 [green stippling on right side]/HOLOTYPE TETHINA WOODI Foster [red border; ‘‘TETHINA WOODI Foster’’ handwritten]. The holotype is double mounted (minuten in a vertical, rectangular card), is in good condition (abdomen removed and dissected; structures in an attached microvial), and is deposited in the USNM (73640). VOLUME 110, NUMBER 2 Fig. 23. 315 Distribution map of Tethina milichioides in the western Nearctic Region. The holotype male of Tethina steyskali Foster is labeled ‘‘PismoBeach 29/8/45 [29 Aug 1945] CAL[ifornia. San Luis Obispo:] ALMelander/ALMelander Collection 1961 [green stippling on right side]/HOLOTYPE TETHINA STEYSKALI Foster [red border; ‘‘TETHINA STEYSKALI Foster’’ handwritten]. The holotype is double mounted (minuten in a vertical, rectangular card), is in good condition (abdomen removed and dissected; structures in an attached microvial), and is deposited in the USNM (73639). Specimens examined from the western Nearctic Region (Fig. 23).—UNITED STATES. CALIFORNIA. Orange: Corona del Mar (33u35.69N, 117u52.59W), 2 Sep 1952, A. L. Melander (1-; USNM); Crystal Cove (33 34.89N, 117 50.79W, beach), 6 Apr 2005, D. and W. N. Mathis (9-, 4U; USNM). Humboldt: Clam Beach (40u59.79N, 124u06.79W9; Hwy 101), 6 Jul 1968, B. J. Peterson (5-, 2U; CNC). San Diego: Camp Pendleton (33u13.3N, 117u24.5W, beach), 4 Apr 2005, D. and W. N. Mathis (9-; USNM); La Jolla (32u51.89N, 117u15.39W, beach), 5 Apr 2005, D. and W. N. Mathis (4-, 1U; USNM). Coronado (32u37.69N, 117u08.39W; beach), 5 Apr 2005, D. and W. N. Mathis (14-, 6U; USNM); Leo Carillo (34u02.89N, 118u56.19W; beach), 7 Apr 2005, D. and W. N. Mathis (4-, 2U; USNM); Oceanside (33u11.89N, 117u23.29W; beach), 4 Apr 2005, D. and W. N. Mathis (1U; USNM). San Luis Obispo: Pismo Beach (35u08.69N, 120u38.59W), 26 Sep 1934, A. L. Melander (1-, 1U; USNM). Santa Barbara: Carpinteria (34u23.99N, 119u31.19W), 12 Jun 1953, A. L. Melander (2-, 3U; USNM). 316 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OREGON. Coos: Coos Bay (43u23.59N, 124u16.89W; beach), 30 Jul 2005, D. and W. N. Mathis (1-; USNM); Bastendorff Beach (43u20.79N, 124u20.99W; beach), 30 Jul 2005, D. and W. N. Mathis (6-; USNM). Lane: Heceta Head (44u08.19N, 124u07.49W; beach), 1 Aug 2005, D. and W. N. USNM); Florence Mathis (6-; (44u019N, 124u08.29W; beach), 1 Apr 2005, D. and W. N. Mathis (20-, 8U; USNM). Lincoln: Otter Crest Wayside (44u44.99N, 124u03.89W; beach), 2 Aug 2005, D. and W. N. Mathis (4-, 2U; USNM); Seal Rock (44u29.89N, 124u059W; beach), 2 Aug 2005, D. and W. N. Mathis (2U; USNM). Tillamook: Cape Kiwanda (45u12.99N, 123u58.39W; beach), 3 Aug 2005, D. and W. N. Mathis (3-, 1U; USNM). WASHINGTON. Pacific: Ilwaco (46u18.69N, 124u02.69W), 12 Jul 1922, A. L. Melander (1-; USNM). Distribution.—Nearctic: United States (California, Oregon, Washington). Remarks.—The variation noted in T. milichioides, especially the male surstylus, is as noted previously in T. albula and in several Palearctic species (Munari 2006). When Foster (1976a) described T. woodi and T. steyskali, he did not then have the extensive data that has been gleaned from more recent studies on the extent of variation in the male genitalia that occurs among some species of Tethina. In light of this new information, we now feel that the ‘‘milichioides group’’ represents a single species. Tethina prognatha (Melander) (Figs. 26) Rhicnoessa prognatha Melander 1952: 202, 206.—Cole 1969: 387 [distribution, diagnosis] Tethina prognatha: Vockeroth 1965: 727 [generic combination; Nearctic catalog].—Mathis and Munari 1996: 18 [world catalog]. Diagnosis.—This species is distinguished from congeners by the following combination of characters: Body generally gray, 3.0 mm; setae black. Head: Gena short, gena-to-eye ratio 0.35; labellum as long as head. Thorax: 4 irregular rows of acrostichal setulae; scutellum uniformly gray, lacking a spot. Femora gray; foretibia yellowish, basal tarsomere yellow, apical 4 tarsomeres brown. Abdomen: Male terminalia unknown. Female terminalia: cerci light gray becoming yellowish distally with thick, short setae and appearing to be fused into one larger cercus similar to T. angustifrons. Type material.—Melander (1952) described Rhicnoessa prognatha from a female holotype that remains the only specimen currently available to us. The holotype is labeled ‘‘Morro Dunes [35u20.89N, 120u50.29W] CAL[IFORNIA. San Luis Obispo:] 6/9/45 A. L. Melander/HOLOTYPE Rhicnoessa prognatha Melander [red label]/ALMelander Collection 1961[right third of label with green stippling]. The holotype is double mounted (minuten in a cardboard rectangle), in excellent condition, and is deposited in the USNM. Distribution (Fig. 26).—Nearctic: United States (California). Remarks.—Melander noted that the cerci of the holotype female are a ‘‘unique spinose hemitubular eighth segment of the abdomen which shows no trace of division into the usual two slender styles.’’ We did not dissect the holotype, being the only known specimen of this species. We have found, however, that dried females of T. angustifrons, a closely related species, often appear to have fused cerci. Upon dissection, however, it quickly became evident that the cerci are separate, long, slender, and approximate. In dried specimens the cerci can appear to be fused. The long labellum and quite prognathous head ally this species with T. milichioides, T. VOLUME 110, NUMBER 2 317 Figs. 24–25. Tethina sasakawai. 24, External male terminalia (epandrium, cerci, and surstyli), posterior view (surstylus more from a posterolateral view). 25, Same, lateral view. angustifrons, and other related congeners. Based on the evidence now available, especially our study of the female holotype, we are of the opinion that T. prognatha is a valid species. Its status, however, should be re-examined when additional specimens, especially males, become available. Tethina sasakawai Foster and Mathis, new species (Figs. 24–26) Diagnosis.—This species is distinguished from congeners by the following combination of characters: Body length 1.83–3.30 mm; body with gray to light brown microtomentum; thorax and abdomen evenly endowed with short but strong setae giving the fly a very bristly appearance, all setae and setulae black. Head: Antenna mostly orange except basal flagellomere brown around base of arista; frons orange; gena high, genato-eye ratio approx. 0.44. Thorax: 4 irregular rows of acrostichal setulae; abdomen uniformly gray microtomentose with a light golden cast; scutellum uniformly gray also with a light golden cast to the microtomentum. Wings hyaline with brownish veins. Femora entirely gray; fore- and hindfemora of male clearly swollen; tibiae uniformly gray, only yellowish at apex to varying extent; basal 3 tarsomeres yellow to orange, 4th tarsomere brown, apical tarsomere black. Abdomen: Gray microtomentose; sternites with apices yellow to white. Male terminalia (Figs. 24–25): epandrium concolorous with rest of abdomen, surstylus articulated with and broadly attached to epandrium, elongate and narrow in posterior view (Fig. 24), apex pointed, armed with 3–4 short peglike spicules, medial surface with longer setulae; surstylus in lateral view (Fig. 25) boot-shaped, anterior and posterior margins gently curved; aedeagus thin, ribbonlike, bearing some hairlike pubescence on dorsal surface. Female 318 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 26. Distribution map of Tethina prognatha (square) and T. sasakawai (dots) in the western Nearctic Region. terminalia: cerci entirely gray, with only weak setulae. Description.—Body length 1.83–3.30 mm; body uniformly dark gray microtomentose. Head: Ocellar triangle dull gray microtomentose; frons orange, parafrons silvery gray microtomentose; frontal lunule silvery; 4 strong, lateroclinate parafrontal setae; 3 pairs of proclinate frontal setae; row of 6–8 short but strong setae medial to the parafrontals; ocellar setae 1 pair, strong, additional 3–6 shorter ocellar setae; paravertical strong, widely separated and convergent. Antenna mostly orange, basal flagellomere with only area around base of arista brown. Gena moderately high, gena-to-eye ratio approx. 0.44, yellowish to golden microtomentose becoming whitish posteriorly, bearing a row of strong dorsoclinate setae in addition to a pair of strong convergent oral vibrissae; face with a pair of shiny tubercles above the oral; palpus yellow; clypeus brown microtomentose. Thorax: Mesonotum uniformly gray microtomentose, with a yellowish to golden cast dorsally and on scutellum and posterior half of anepisternum. Wing hyaline but slightly smoky. Coxae, femora and tibiae uniformly gray microtomentose and evenly setulose with long, strong setae; only extreme apex of femora and extreme base of tibiae slightly orange; fore- and hindfemora of male moderately to strongly swollen; apical tarsomere and half of 4th tarsomere black, basal three tarsomeres yellow to orange. Dorsocentrals 1 + 3, VOLUME 110, NUMBER 2 acrostichals in 4 irregular rows with 1 pair of strong prescutellars. Mesonotum evenly covered with numerous setae only slightly shorter than the dorsocentrals. Proepisternum with 2 dorsoclinate setae. Anepisternum and katepisternum strongly setose. Wing hyaline with brown veins. Abdomen: Uniformly gray microtomentose except posterior margins white. Evenly setose with long, strong setae. Male terminalia (Figs. 24–25): surstylus broadly articulated with ventral margin of epandrium and internally with subepandrial sclerite, anterior margin in lateral view strongly posteriorly curved, posterior margin in lateral view also strongly curved and with a row of small setulae along posterior margin and several very thick short setulae along the medioventral margin; cercus normal; aedeagus thin, ribbonlike, micropubescent; ejaculatory apodeme widely flared. Female cercus brown to black and weakly setulose. Type material.—The holotype male is labeled ‘‘USA. CA[LIFORNIA]. San Diego: Camp Pendleton (33u13.39N, 117u24.59W, beach), 4 Apr 2005 Wayne N. & D. Mathis/HOLOTYPE Tethina sasakawai - G. A. Foster & W. N. Mathis USNM [red label; species name and ‘‘-’’ and ‘‘Foster &’’ handwritten].’’ The holotype is double mounted (minuten in a vertically mounted white plastic cube), is in excellent condition, and is deposited in the USNM. Sixteen paratypes (12-, 4U; USNM) bear the same label data as the holotype. Other paratypes are as follows: CANADA. BRITISH COLUMBIA. Vancouver, Point Grey (49u179N, 123u149W), 31 Jul 1973, J. R. Vockeroth (1-; CNC); Vancouver, Stanley Park (48u409N, 124u519W), 8 Jun 1973, J. R. Vockeroth (1-; CNC); Galiano Island, Montague Harbour (48u539N, 123u249W), 20 Jul 1973, J. R. Vockeroth (4-, 3U; CNC). JAPAN. HOKKAIDO: Shari Beach, Shari-gun, 3 Aug 1967, T. Saigusa (2-, 319 2U; USNM; paratypes of T. thula Sasakawa). UNITED STATES. CALIFORNIA. Los Angeles: Leo Carillo (34u02.89N, 118u56.19W; beach), 7 Apr 2005, D. and W. N. Mathis (6-, 2U; USNM). Orange: Crystal Cove (33u34.89N, 117u50.79W; beach), 6 Apr 2005, D. and W. N. Mathis (8-, 1U; USNM). San Diego: Oceanside (33u11.89N, 117u23.29W; beach), 4 Apr 2005, D. and W. N. Mathis (20-, 8U; USNM); La Jolla (32u51.89N, 117u15.39W, beach), 5 Apr 2005, D. and W. N. Mathis (7-; USNM). San Francisco: San Francisco (37u48.69N, 122u21.19W), 22 Jun 1947, A. L. Melander (2-, 1U; syntypes of T. horripilans; USNM). Distribution (Fig. 26).—Nearctic: Canada (British Columbia), United States (California). Palearctic: Japan (Hokkaido). Etymology.—The species epithet, sasakawai, is a genitive patronym to recognize Professor Mitsuhiro Sasakawa for his numerous contributions to our knowledge of Acalyptrate Diptera, including Tethininae, and for his cordial friendship. Remarks.—This species is quite similar in external appearance to T. thula, although in general T. sasakawai is usually somewhat lighter in color. The gena of T. sasakawai tends to be more golden to yellowish and the mesonotum tends to have a golden yellow cast in lateral view while T. thula tends to have a metallic brown gena and a darker brown cast to the mesonotum. Examination of structures of the male terminalia, however, is often required to distinguish between these two species. Tethina spinulosa Cole (Figs. 27–30) Tethina spinulosa Cole 1923: 478.—Hendel 1934: 41 [citation].—Vockeroth 1965: 728 [Nearctic catalog].—Foster 320 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 27–29. Tethina spinulosa. 27, Head, lateral view. 28, External male terminalia (epandrium, cerci, and surstyli), posterior view (surstylus more from a posterolateral view). 29, Same, lateral view. 1976b: 2 [Neotropical catalog].— Mathis and Munari 1996: 18 [world catalog].—Foster and Mathis 2000: 544–546 [synonymy; review, neotropics]. Rhicnoessa spinulosa: Melander 1952: 202, 208 [generic combination; key, citation].—Cole 1969: 387 [distribution, diagnosis]. Tethina setulosa Malloch 1934: 454.— Foster 1976b: 2 [Neotropical catalog].—Mathis and Munari 1996: 18 [world catalog].—Foster and Mathis 1998: 624–625 [revision, Caribbean and Gulf of Mexico]; 2000: 544–546 [synonymy; review; neotropics]. Rhicnoessa setulosa: Hennig 1937: 139 [generic combination; citation]. Diagnosis.—This species is distinguished from congeners by the following combination of characters: Body length 1.85–2.70 mm; body generally with gray microtomentum; setae black. Head (Fig. 27): Gena short, gena-to-eye ratio about 0.40 or less. Thorax: 4 somewhat irregular rows of acrostichal setulae; scutellum uniformly gray, lacking apical spot. Femora and tibiae grayish to orange; hindfemur of male distinctly swollen, distinctly larger than fore- and midfemora; tibiae and basal 4 tarsomeres VOLUME 110, NUMBER 2 Fig. 30. 321 Distribution map of Tethina spinulosa in the western Nearctic Region. yellow, apical tarsomere brown. Abdomen: Usually gray microtomentose but may have several sternites orange or yellow giving the fly an overall yellowish appearance. Male terminalia (Figs. 28– 29): epandrium concolorous with rest of abdomen, surstylus articulated with and broadly attached to epandrium, spatulate in posterior view (Fig. 28), length about equal to width, median margin bearing dense patch of robust setulae along entire length (Fig. 28); surstylus in lateroblique view (Fig. 29) broadly rounded, constricted anteriorly, external surface bearing numerous setulae; aedeagus thin, ribbonlike. Female terminalia: cerci gray, with only weak setulae. Type material.—The holotype male of Tethina spinulosa Cole is labeled ‘‘[MEXICO. Baja California:] Las Animas Bay [28u499N, 113u219W] Gulf Cal. May 8 1921/EPVan Duzee Collector/HOLOTYPE spinulosa/ALLOTYPE spinulosa/Tethina spinulosa Type and allotype [two specimens on separate points; type data taken from Arnaud 1979: 345].’’ The holotype and allotype are double mounted (glued to separate paper points on same pin) and are deposited in the CAS (1,356). A number of Cole’s paratypes are in the USNM as follows: MEXICO. BAJA CALIFORNIA: Las Ánimas Bay, Gulf of California (28u499N, 113u219W), 8 May 1921, A. L. Melander (14-, 36U; USNM). BAJA CALIFORNIA SUR: Loreto (26u019N, 111u219W), 19 May 1921, E. P. VanDuzee (22-, 10U; USNM). Other paratypes that were noted by Cole, (Sal Si Puedes Island, 9 May 1921, and San Francisquito Bay, 10 May 1921), are not in the USNM. 322 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The holotype male of Tethina setulosa Malloch is labeled ‘‘Angol [crossed out] Chile DSBullock/Tocopilla [Antofagasta] Ap. 10, [19]31 Sea Beach [handwritten]/Type No. 50448 U.S.N.M. [red; ‘‘50448’’ handwritten]/Tethina setulosa Type Det. JRMALLOCH [species name and ‘‘Type’’ handwritten; black submargin].’’ The holotype is directly pinned, is in good condition (abdomen removed and dissected, the parts are in an attached microvial), and is deposited in the USNM (50448). Additional specimens examined from the western Nearctic Region (Fig. 30).— MEXICO. BAJA CALIFORNIA: Las Ánimas Bay, Gulf of California (28u499N, 113u219W), 8 May 1921, E. P. VanDuzee (1-, 4U; CAS). BAJA CALIFORNIA SUR: Loreto (26u019N, 111u219W; Sefton Orca Expedition to Gulf of California), 29 Mar 1953, P. H. Arnaud, Jr. (27-, 21U; CAS); Los Frailes (La Paz; 23u22.69N, 109u26.29W; Sefton Orca Expedition to Gulf of California), 18 Aug 1953, P. H. Arnaud, Jr. (4-, 1U; CAS). SONORA: Guaymas (64.5 km N; 28u31.79N, 110u569W), 24 Nov 1951, H. J. Teas (3-, 4U; USNM); Puerto de Lobos (30u179N, 112u529W), 18–19 Mar 1974, V. Roth, W. Brown (5-, 2U; USNM); Rocky Point (Puerto Peñasco; 31u199N, 113u329W; marsh), 21 Apr 1947, A. L. Melander (1-; USNM); Bahia San Carlos (27u57.59N, 111u049W), 10 Aug 1960, P. H. Arnaud, Jr., E. S. Ross, D. C. Rentz (6-, 6U; CAS). UNITED STATES. CALIFORNIA. Imperial: Palo Verde (33u25.99N, 114u43.99W), 15 Oct 1944, A. L. Melander (4-, 6U; USNM); Salton City (33u17.99N, 115u57.49W; shore of Salton Sea), 18 Mar 1986, J. E. Lowry, W. E. Steiner (1U; USNM). Orange: Balboa (33u34.69N, 117u54.19W), 13 Jul 1940, A. L. Melander (2-; USNM); Corona del Mar (33u35.69N, 117u52.59W), 25 Jul–19 Nov 1942, 1943, 1946, 1949, A. L. Melander (7-, 5U; USNM); Doheny Park (33u27.59N, 117u40.29W), 12 Oct. 1951, A. L. Melander (3-, 7U; USNM); Laguna Beach (33u32.39N, 117u46.69W 29), Apr–24 Oct 1932, 1943, 1951, 1957, 1958, J. M. Aldrich, A. L. Melander (21-, 16U; USNM); San Clemente (33u25.49N, 117u37.49W), 5 Oct 1950, A. L. Melander (4-, 6U; USNM); Seal Beach (33u44.39N, 118u06.29W), 26 Jul 1942, A. L. Melander (2-, 1U; USNM). San Diego: La Jolla (32u51.89N, 117u15.39W), 29 Aug–9 Oct 1952, 1984, D. King, A. L. Melander (13-, 9U; USNM); Mission Beach on Mission Bay (32u46.59N, 117u15.19W), 15 Jun 2003, S. D. and A. V. Gaimari (5-, 1U; CDFA); San Diego (32u42.69N, 117u09.39W), 3 Aug–10 Dec 1916, 1932, J. M. Aldrich, H. G. Dyer (1-, 8U; USNM); San Onofre State Beach (33u24.79N, 117u34.29W; 5 m), 19 Jun 1996, M. Gates (1-; USNM); Torrey Pines State Park (32u55.39N, 117u15.19W), 19 Jun 1996, M. Gates (2-; USNM). [FLORIDA. MONROE: Marathon Key (24u41.69N, 81u059W), 11 Feb 2000, W. N. Mathis (1-; USNM); Lower Matecumbe Key (24u51.49N, 80u43.79W), 10 Feb 2000, W. N. Mathis (3-; USNM). Pinellas: Tampa Bay (27u31.19N, 82u36.89W), 12 Feb 2000, W. N. Mathis (1-; USNM).] Distribution.—Nearctic: United States (California, Florida (new record)). Neotropical: Chile (Tarapaea to Antofagasta), Ecuador (Galápagos Islands), Mexico (Baja California, Baja California Sur, Sonora, Tabasco). Remarks.—This species is typically quite robust and very setose. Males have both fore- and hindfemora swollen, the hindfemur being much more swollen than that of the foreleg, however. The femora range in color from entirely gray to entirely orangish. All setae are generally very stout and well developed. Considerable variation is evident in the coloration of the abdominal sternites, VOLUME 110, NUMBER 2 323 Figs. 31–32. Tethina thula. 31, External male terminalia (epandrium, cerci, and surstyli), posterior view (surstylus more from a posterolateral view). 32, Same, lateral view. which vary from entirely gray to mostly yellowish orange, at least on the posterior half of the sternites. Tethina thula Sasakawa (Figs. 31–33) Tethina thula Sasakawa 1986: 436.— Mathis and Munari 1996: 19 [world catalog].—Munari 2002: 24 [citation, distribution note]. Rhicnoessa thula.—Beschovski and Nartshuk 1997: 138 [generic combination; Russia, figs. of -, U terminalia, head, and wing, description]. Diagnosis.—This species is distinguished from congeners by the following combination of characters: Body length 1.7–3.3 mm; body with gray microtomentum; all setae and setulae black. Head: Antenna mostly yellow except third flagellomere brown around base of arista; frons orange yellow; gena high, tending to be a light metallic brown, gena-to-eye ratio 0.44. Thorax: 4 irregular rows of acrostichal setulae; mesono- tum and scutellum with a brownish cast in lateral view. Wings lightly smoky hyaline with dark brown veins. Femora uniformly gray; fore- and hindfemora of male swollen; tibiae uniformly grey, extreme base or tip sometimes slightly yellow; basal tarsomere yellow, tarsomeres 2, 3, and 4 variably brown to black, apical tarsomere always black. Abdomen: Gray microtomentose, apices yellow. Male terminalia (Figs. 31–32): epandrium concolorous with rest of abdomen, surstylus articulated with and broadly attached to epandrium, roundly spatulate in posterior view (Fig. 31), evenly covered with setulae laterally and medially, with a row of 5–6 short peglike spicules along ventral mesal surface, surstylus in lateral view (Fig. 32) broad and rounded on posterior margin, anterior margin somewhat curved; aedeagus thin, ribbonlike. Female terminalia: cerci entirely gray with only weak setulae. Type material.—The holotype male of Tethina thula Sasakawa is labeled ‘‘[Japan.] HOKKAIDO [Notsuke-gun,] Bek- 324 Fig. 33. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Distribution map of Tethina thula in the western Nearctic Region. kai [Beach] 2. viii. 1967 [2 Aug 1967] T. SAIGUSA/- [glued to a rectangular, green label]/HOLOTYPE Tethina thula n. sp. Det. M.SASAKAWA [pink; ‘‘Tethina thula n. sp.’’ handwritten]/Holotype KPU Type No. 235 [pink; ‘‘KPU Type No. 235’’ handwritten].’’ The holotype is double mounted (glued to a paper triangle), is in excellent condition, and is deposited in the OMNH (formerly in Kyoto Prefectural University—KPU). Specimens examined from the western Nearctic Region (Fig. 33).—UNITED STATES. ALASKA. Kenai: Lowell Point (6.5 km S of Seward; 60u03.99N, 149u26.69W), 31 Jul 2003, D. and W. N. Mathis (42-, 8U; USNM); Deep Creek (60u029N, 151u429W; gravel to cobble beach), 2 Aug 2002, D. and W. N. Mathis (16-, 1U; USNM); Homer Spit (59u389N, 151u29.89W; beach), 2 Aug 2002, D. and W. N. Mathis (2-, USNM); Homer (59u38.59N, 1U; 151u33.29W; cobble beach), 3 Jul 2006, D. and W. N. Mathis (6-, 3U; USNM); Ninilchik (60u039N, 151u40.29W; beach), 2 Jul 2006, D. and W. N. Mathis (3-, USNM). Kodiak: Katmai 2U; (58u00.19N, 154u54.79W), July 1917, J. S. Hine (1-, 1U; USNM); Kukak Bay (58u17.79N, 154u38.89W), 4 Jul 1899, T. Kincaid (1-; USNM). Distribution.—Nearctic: United States (Alaska (New Record)). Palearctic: Japan (Hokkaido), Russia (Far East, South Sakhalin). Remarks.—Examination of the male type specimen of T. thula shows it to match Sasakawa’s (1986: fig. 2, p. 437) illustration. Fortunately the surstylus of the type is readily visible without need for dissection. We have also examined VOLUME 110, NUMBER 2 325 Figs. 34–38. Tethina willistoni. 34, Head, lateral view. 35, External male terminalia (epandrium, cercus, and surstylus), lateral view. 36, Same, lateral view. 37, Same, posterior view. 38, Same, posterior view. four paratypes generously provided by Dr. M. Sasakawa and now deposited in the USNM. One of these males has been dissected and examined. We have discovered that the structures of the male terminalia of this paratype do not match Sasakawa (1986) or Beschovski’s (1997) illustrations. These structures do, however, match those of the new species, T. sasakawai, described above and the specimens are listed under that species. The external differences between these two species are slight and overlapping, and it is not difficult to see why the paratype series was partially mixed. Tethina willistoni Melander (Figs. 34–39) Anthomyza cinerea Williston 1896: 444 [preoccupied, Loew 1862]. 326 Fig. 39. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Distribution map of Tethina willistoni in the western Nearctic Region. Rhicnoessa willistoni Melander 1913: 298 [new name for T. cinerea of Williston, not Loew].—Melander 1952: 201, 209 [citation]. Tethina willistoni.—Foster 1976b: 3 [generic combination; Neotropical catalog].—Mathis and Munari 1996: 19 [world catalog].—Foster and Mathis 1998: 615–618 [revision; Caribbean and Gulf of Mexico] .—Mathis and Foster 2007: 420–424 [review; fauna of Delmarva states]. Rhicnoessa bermudaensis Melander 1952: 203.—Mathis and Foster 2007: 421 [synonymy]. Tethina bermudaensis.—Vockeroth 1965: 727 [generic combination; Nearctic catalog].—Mathis and Munari 1996: 15 [world catalog].—Foster and Mathis 1998: 611–613 [revision, lectotype designation; Caribbean and Gulf of Mexico, fig. of - terminalia]. Rhicnoessa variseta Melander 1952: 201, 209.—Cole 1969: 387 [distribution, diagnosis].—Foster and Mathis 1998: 615 [synonymy; revision, Caribbean]. Tethina variseta.—Vockeroth 1965: 728 [generic combination; Nearctic catalog].—Mathis and Munari 1996: 19 [world catalog]. Tethina carioca Prado and Tavares 1966: 433 [figs. of - terminalia and wing].— Foster and Mathis 1998: 615 [synonymy]. Diagnosis.—This species is distinguished from congeners by the following combination of characters: Body length 1.65–3.00 mm; body generally whitish gray to gray, microtomentose; setae generally white to slightly off white but sometimes with all setae black. Head (Fig. 34): Gena high, greater than 0.5 eye height. Thorax: 4 irregular rows of VOLUME 110, NUMBER 2 acrostichal setulae; scutellum uniformly gray. Femora mostly yellow to mostly gray; hindfemur of male similar to or only slightly more swollen than fore- and midfemora; tibiae yellow; basal 4 tarsomeres yellow, apical tarsomere brown. Abdomen: Male terminalia (Figs. 35–38): Surstylus articulated with and broadly attached to epandrium, broadly spatulate/triangular in posterior view (Fig. 37), length 2–33 width, apex broadly rounded; medial margin bearing numerous short, stout setulae along entire length; surstylus in lateral view (Figs. 36–37) narrow, tapered to apical point, posterior margin almost straight; basal portion produced anteriorly as a broadly rounded lateral lobe bearing several short setulae mesally; aedeagus thick, straplike. Type Material.—The neotype male of A. cinerea Williston (designated by Foster & Mathis 1998: 616) is labeled ‘‘W.I. St.Vincent: Wallilabou-beach [13u159N, 61u169W], 27 March 1989 Wayne N. Mathis/NEOTYPE Anthyomyza cinerea - Williston by Foster and Mathis [red, handwritten].’’ The neotype is double mounted (minuten in a plastic block), is in excellent condition, and is deposited in the USNM. The lectotype male of R. variseta Melander (designated by Foster & Mathis 1998: 616) is labeled ‘‘[USA.] CoronaDelMar 29/6/42 [29 Jun 1942] CAL[ifornia] A L Melander/ALMelander Collection 1961 [green stippling on right side]/HOLOTYPE Rhicnoessa variseta Melander [red]/LECTOTYPE Rhicnoessa variseta - Melander By Foster and Mathis [handwritten except for ‘‘LECTOTYPE’’ and ‘‘By’’; black submarginal border].’’ The lectotype is double mounted (minuten in a rectangular card on end), is in excellent condition, and is deposited in the USNM. The holotype male of Tethina carioca Prado & Tavares is labeled ‘‘[Brazil. Ilha do] Governador: Galeão[,] Rio[de Janeiro]-Brasil. 11.X.66 [11 Oct 1966] 327 Lopes and Prado/Tethina carioca n.sp Prado and Tavares det/Holotypus [red]/ N. 13.356 [number handwritten] DIPTERA Inst.Oswaldo Cruz [black margin].’’ The holotype is double mounted (minuten partially wound around base pin), is in excellent condition (abdomen removed, dissected, parts are in an attached microvial), and is now deposited in MZSP. The lectotype male of Rhicnoessa bermudaensis Melander (designated by Foster & Mathis 1998: 612) is labeled ‘‘BERMUDA Cooper ISL [32u21.29N, 64u39.69W] 25 Jan [19]34 ALMelander/ ALMelander Collection 1961 [right third of label with green stippling]/PARATYPE Rhicnoessa bermudaensis Melander [yellow]/LECTOTYPE - Rhicnoessa bermudaensis Melander By Foster and Mathis [handwritten except for ‘‘LECTOTYPE’’ and ‘‘By’’; black submarginal border].’’ The lectotype is double mounted (minuten in a cardboard rectangle), is in excellent condition (the abdomen has been removed and dissected; the parts are in an attached microvial), and is deposited in the USNM. Specimens examined from the western Nearctic Region (Fig. 39).—UNITED STATES. CALIFORNIA. Los Angeles: Long Beach (33u469N, 118u11.49W), 21 May 1944, A. L. Melander (1-; USNM). Orange: Balboa (33u35.29N, 117u549W), 15 Jul 1940, A. L. Melander (1-; USNM); Corona del Mar (33u35.69N, 117u52.59W), 29 Jun–2 Sep 1940, 1942, 1952, A. L. Melander (4-, 1U; USNM); Huntington Beach (33u39.69N, 118uW), 4 Jun 1945, A. L. Melander (2-; USNM); Seal Beach (33u44.39N, 118u06.29W), 26 Jul 1942, A. L. Melander (3-, 5U; USNM). Distribution.—Nearctic: Bermuda, United States (California, Connecticut, Delaware, Florida, Maryland, Massachusetts, North Carolina, South Carolina, Virginia). Neotropical: Bahamas, 328 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Belize, Brazil (Rio de Janeiro), Cuba, Curaçao, Ecuador, Mexico (Chihuahua, Tabasco), Panama, Peru, Tobago, Turks and Caicos, West Indies (Anguilla, Antigua, Barbados, Barbuda, Dominica, Dominican Republic, Grand Cayman, Grenada, Jamaica, Montserrat, Puerto Rico, St. Croix, St. Lucia, St. Vincent). Remarks.—The variation in setal coloration and size of T. willistoni is remarkable. While we have seen virtually no variation in structures of the male terminalia, the variation in external characters is as follows: specimens that are more robust have mostly stout, black setae and in general present a very ‘‘bristly’’ habitus (similar to T. spinulosa and T. horripilans). Smaller, more delicate specimens have only the apical scutellar setae black with all other setae being white. Many specimens fall between these two extremes, making it virtually impossible to distinguish between T. willistoni and other species on the basis of external structures alone. Specimens we have seen from the study area exhibit the mostly whitish habitus with whitish wings and brown veins. Most setae are white with a few others, especially the apical scutellars, being black. ACKNOWLEDGMENTS We gratefully acknowledge the assistance and cooperation of many organizations and individuals who contributed to the field work and production of this paper. To our colleagues and their institutions listed below, who loaned specimens, we express our sincere thanks. Without their cooperation this study could not have been completed: David A. Grimaldi and Julian Stark (AMNH); Jon K. Gelhaus, Donald F. Azuma, and Jason D. Weintraub (ANSP); Paul H. Arnaud, Jr. and Anthea Carmichael (CAS); Stephen D. Gaimari (CDFA); James E. O’Hara and Jeff Cumming (CNC); Philip D. Perkins (MCZ); Roberto Lamas (MZSP); and Rikio Matsumoto and Mitsuhiro Sasakawa (OMNH). Hollis B. Williams provided technical support and produced the maps. James F. Edmiston advised on the production of the maps. The illustrations were carefully inked by Young T. Sohn from pencil drawings. For reviewing a draft of this paper we thank Lorenzo Munari, Oliver S. Flint, Jr., and J. Richard Vockeroth. LITERATURE CITED Ardö, P. 1957. Studies in the marine shore dune ecosystem with special reference to the Dipterous fauna. Opuscula Entomologica, Supplementum 14: 1–255. Arnaud, P. H., Jr. 1979. A catalog of the types of Diptera in the collection of the California Academy of Sciences. Myia 1: 505 pp. Beschovski, V. L. 1993. Taxonomic and systematic notes on the genera Tethina Haliday, 1838, and Rhicnoessa Loew, 1862 (Insecta: Diptera: Tethinidae). Reichenbachia 30(16): 103–107. ———. 1994. Contribution to the study of the west Palaearctic Tethinidae (Diptera). Acta Zoologica Bulgarica 47: 16–29. Beschovski, V. L. and E. P. Nartshuk. 1997. The Tethinidae species in the collection of the Zoological Institute in St. Petersburg (Insecta: Diptera: Tethinidae). Reichenbachia 32(22): 129–141. Clausen, P. J. and E. F. Cook. 1971. A revision of the Nearctic species of the tribe Parydrini (Diptera: Ephydridae). Memoirs of the American Entomological Society Number 27, 150 pp. Cole, F. R. 1923. Expedition of the California Academy of Sciences to the Gulf of California in 1921. Diptera from the Islands and Adjacent Shores of the Gulf of California. II. General Report. Proceedings of the California Academy of Sciences 12(25): 457–481. Cole, F. R. (with the collaboration of E. T. Schlinger). 1969. The flies of western North America. University of California Press, Berkeley and Los Angeles, xi+693 pp. Collin, J. E. 1911. Additions and corrections to the British list of Muscidae Acalyptratae [part]. Entomologist’s Monthly Magazine 46: 229– 234. Cumming, J. M., B. J. Sinclair, and D. M. Wood. 1995. Homology and phylogenetic implications of male genitalia in Diptera-Eremoneura. Entomologica Scandinavica 26: 121–149. VOLUME 110, NUMBER 2 Curran, C. H. 1934. The families and genera of North American Diptera. The Ballou Press, New York, 512 pp. Curtis, J. 1837. A guide to an arrangement of British insects; being a catalogue of all the named species hitherto discovered in Great Britain and Ireland, London, vi+294 pp. Czerny, L. 1928. 55. Tethinidae. In Lindner, E., ed. Die Fliegen der palaearktischen Region 5(2): 1–8. Stuttgart. de Meijere, J. C. H. 1928. Vierde Supplement op de Nieuwe Naamlijst van Nederlandsche Diptera. Tijdschrift voor Entomologie 71: 11–83. Foster, G. A. 1976a. Notes on the phylogeny of the Nearctic Tethinidae and a review of the genus Neopelomyia Hendel and the Tethina milichioides Group (Diptera). Proceedings of the Entomological Society of Washington 78(3): 336–352. ———. 1976b. 74. Family Tethinidae, pp. 1–4. In Papavero, N., ed. A catalogue of the Diptera of the Americas south of the United States. Museu de Zoologia, Universidade de São Paulo, São Paulo. Foster, G. A. and W. N. Mathis. 1998. A revision of the family Tethinidae (Diptera) from the Caribbean, Gulf of Mexico, and Bermuda. Proceedings of the Entomological Society of Washington 100(4): 601–632. ———. 2000. Notes on Neotropical species of Tethina Haliday (Diptera: Tethinidae). Proceedings of the Entomological Society of Washington 102(3): 542–548. ———. 2003. A revision of the genera Pelomyia Williston and Masoniella Vockeroth (Diptera: Tethinidae). Smithsonian Contributions to Zoology 619, 63 pp. Frey, R. 1919. Mitteilungen über südamerikanische Dipteren. Vetenskaps-Societetens Förhandlingar 60A 14(1918): 1–35. Grimaldi, D. A. 1987. Phylogenetics and taxonomy of Zygothrica. Bulletin of the American Museum of Natural History 186: 103–268. Haliday, A. H. 1838. New British insects indicated in Mr. Curtis’s Guide [part]. Annals and Magazine of Natural History 2: 183–190. Hardy, D. E. and M. Delfinado. 1980. Tethinidae, pp. 369–379. In Hardy, D. E. and M. D. Delfinado, eds. Insects of Hawaii, 13. Diptera: Cyclorrhapha III. University Press of Hawaii, Honolulu, 451 pp. Hendel, F. 1902. Ueber die systematische Stellung der Dipteren-gattungen Pseudopomyza Strobl u. Rhicnoëssa Lw. Wiener Entomologische Zeitung 21(10): 261–264. ———. 1911. Über von Professor J. M. Aldrich erhaltene und einige andere amerikanische Dipteren. Wiener Entomologische Zeitung 30(2–3): 19–46. 329 ———. 1917. Beiträge zur Kenntnis der acalyptraten Musciden. Deutsche Entomologische Zeitschrift 1917(1): 33–47. ———. 1934. Revision der Tethiniden (Dipt. Muscid. acal.). Tijdschrift voor Entomologie 1934: 37–54. Hennig, W. 1937. Systematisch-tiergeographische Beiträge zur Kenntnis der Tethiniden (Dipt., Acalypt.). Entomologischen Rundschau 54(9)(1936): 136–140. Johnson, C. W. 1910. Order Diptera, pp. 703–814. In Smith, J. B., ed. The insects of New Jersey. New Jersey State Museum Annual Report 1909, pp. 15–888. ———. 1913. Insects of Florida. I. Diptera. Bulletin of the American Museum of Natural History 32(3): 37–90. ———. 1925. Fauna of New England. 15. List of the Diptera or two-winged flies. Occasional Papers of the Boston Society of Natural History 7: 326. ———. 1930. A list of the insect fauna of Nantucket, Massachusetts. The Nantucket Maria Mitchell Association 3(2): 1–175. Loew, H. 1862. Ueber einige bei Varna gefangene Dipteren. Wiener Entomologische Zeitung 6(6): 161–175. ———. 1865. Ueber die europäischen Arten der Gattung Rhicnoëssa. Berliner Entomologische Zeitschrift 9: 34–39. ———. 1869. Diptera Americae septentrionalis indigena. Berliner Entomologische Zeitschrift 13: 1–52. Malloch, J. R. 1913. A synopsis of the genera of Agromyzidae, with descriptions of new genera and species. Proceedings of the U.S. National Museum 46: 127–154. ———. 1934. Tethinidae, pp. 452–460. In Edwards, F., ed. Diptera of Patagonia and south Chile. British Museum (Natural History), London. Mathis, W. N. and G. A. Foster. 2007. Canacidae (Diptera) from the Delmarva States. Proceedings of the Biological Society of Washington 120(4): 387–428. Mathis, W. N. and L. Munari. 1996. World catalog of the family Tethinidae (Diptera). Smithsonian Contributions to Zoology 584: 1–27. Mathis, W. N. and M. Sasakawa. 1989. 102. Family Tethinidae, pp. 667–668, pp. 803–804. In Evenhuis, N. L., ed. Catalog of the Diptera of the Australasian and Oceanian Regions. E. J. Brill and B. P. Bishop Museum special publication 86, Honolulu, 1155 pp. McAlpine, J. F. 1981. Morphology and Terminology—Adults, pp. 9–63. In McAlpine, J. F., et al., eds. Manual of Nearctic Diptera. Agriculture Canada, Research Branch, Monograph 27, Vol. 1, Ottawa, 674 pp. 330 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Melander, A. L. 1913. A synopsis of the dipterous groups Agromyzinae, Milichinae, Ochthiphilinae and Geomyzinae. Journal of the New York Entomological Society 21(4): 283–300. ———. 1952. The North American species of Tethinidae (Diptera). Journal of the New York Entomological Society 59: 187–212. Munari, L. 1981. Tre Nuove Tethina Haliday Raccolte dal Prof. A. Giordani Soika in Asia Minore e Senegal (Diptera, Tethinidae). Bolletino del Museo Civico di Storia Naturale di Venezia (1980) 31: 139–144. ———. 1990. Contributo alla conoscenza dei Tethinidae afrotropicali. IV. Tethinidae raccolti ad Aldabra dalla ‘‘Aldabra Atoll Royal Society Expedition (1967–68)’’ e nel Sud Africa da R.E. Turner e B. and P. Stuckenberg, con descrizione di due nuove specie (Diptera, Acalyptratae). Società Veneziana di Scienze Naturali—Lavori 15: 51–68. ———. 2002. Beach flies (Diptera: Tethinidae) of the Palaearctic Region: An annotated checklist, including world distribution. Società Veneziana di Scienze Naturali—Lavori 27: 17–25. ———. 2006. New synonymies and lectotype designations in Western Palaearctic Tethinidae, with some remarks on the intraspecific variability of the surstylus of Tethina strobliana (Mercier, 1923). (Diptera: Brachycera, Acalyptrata). Bolletino del Museo Civico di Storia Naturale di Venezia 57(2006): 101–115. Prado, A. P. do. and O. Tavares. 1966. Sôbre duas espécies novas do gênero ‘‘Tethina’’ Haliday, 1838 (Diptera, Tethinidae). Revista Brasileira de Biologia 26(4): 429–439. Sabrosky, C. W. 1999. Family-group names in Diptera. An annotated catalog. Myia, Vol. 10, Backhuys Publishers, Leiden, 576 pp. [including a bibliography by F. C. Thompson, N. L. Evenhuis, and C. W. Sabrosky]. Sasakawa, M. 1986. A revision of the Japanese Tethinidae (Diptera). Kontyû 54(3): 433–441, 4 figures. Steyskal, G. C. and M. Sasakawa. 1977. Family Tethinidae, pp. 395–395. In Delfinado, M. D. and D. E. Hardy, eds. A catalog of the Diptera of the Oriental Region. Vol. III. Suborder Cyclorrapha (excluding Division Aschiza). University Press of Hawaii, Honolulu, 854 pp. Sturtevant, A. H. 1923. New species and notes on synonymy and distribution of Muscidae Acalypteratae (Diptera). American Museum Novitates 76: 1–12. Thompson, F. C. and W. N. Mathis. 1981. Haliday’s generic names of Diptera first published in Curtis’ A Guide to…British Insects (1837). Journal of the Washington Academy of Sciences 70(2): 80–89. Vockeroth, J. R. 1965. Family Tethinidae, pp. 726–728. In Stone, A., ed. A Catalog of the Diptera of America North of Mexico. United States Department of Agriculture, Agriculture Handbook 276, 1696 pp. ———. 1987. 101. Tethinidae, pp. 1073–1078. In McAlpine, J. F., et al., eds. Manual of Nearctic Diptera. Research Branch, Agriculture Canada, Monograph 28, Ottawa. Vol. 2, pp. iv+675–1332. Williston, S. W. 1893. List of Diptera of the Death Valley Expedition. North American Fauna 7: 253–259. ———. 1896. XI. On the Diptera of St. Vincent (West Indies). Transactions of the Entomological Society of London 3: 253–446. ———. 1908. Manual of North American Diptera. Third ed., 405 pp. New Haven, Connecticut.

RELATED PAPERS

RELATED TOPICS

Log In

Review of the Genus Tethina Haliday (Diptera: Canacidae: Tethininae) from Western North America

Review of the Genus Tethina Haliday (Diptera: Canacidae: Tethininae) from Western North America

Review of the Genus Tethina Haliday (Diptera: Canacidae: Tethininae) from Western North America

Review of the Genus Tethina Haliday (Diptera: Canacidae: Tethininae) from Western North America

Review of the Genus Tethina Haliday (Diptera: Canacidae: Tethininae) from Western North America

Related Papers

RELATED PAPERS

RELATED TOPICS