Shore (Insecta: Diptera: ) of Plummers Island, Maryland Author(s): Wayne N. Mathis and Dianne Mathis Source: Bulletin of the Biological Society of Washington, 15(1):173-191. Published By: Biological Society of Washington DOI: http://dx.doi.org/10.2988/0097-0298(2008)15[173:SFIDEO]2.0.CO;2 URL: http://www.bioone.org/doi/full/10.2988/0097-0298%282008%2915%5B173%3ASFIDEO %5D2.0.CO%3B2

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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. Shore Flies (Insecta: Diptera: Ephydridae) of Plummers Island, Maryland

Wayne N. Mathis and Dianne Mathis Department of Entomology, NHB 169, P.O. Box 37012, Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A., e-mail: [email protected]

Abstract.—Twenty-nine species of shore flies in 17 genera have been col- lected and identified from Plummers Island. Although relatively diverse, the shore-fly fauna known from the Island represents only 21% of the species known from the surrounding states, indicating that we probably are dealing with sampling error and that many more species are likely to be found there. Key words.—Inventory, diagnosis, distribution, systematics.

Ephydrids or shore flies are well known among gender. Although many specimens examined for this true flies (order Diptera) because of their rather study are in the National Museum of Natural History, unique ability to proliferate in environments that are Smithsonian Institution, Washington, D.C. (USNM), normally inimical to life. Breeding habitats for some we also borrowed and studied numerous specimens, species include mineral and thermal springs, petro- especially primary types, that are deposited in the leum pools, highly saline lakes, salt marshes, and following museums: AMNH—American Museum of small carrion (Foote 1995). Their unusual ability to Natural History, New York, New York; ANSP— breed in such a diverse array of habitats prompted Academy of Natural Sciences of Philadelphia, Penn- Oldroyd (1964:189), a renowned English dipterist, to sylvania; BMNH—The Natural History Museum, write: ‘‘Evidently we are seeing in the Ephydridae a former British Museum (Natural History), London, family of flies in the full flower of its evolution, and England; KU—Snow Entomological Museum, Uni- as such they offer attractive material for study, not versity of Kansas, Lawrence, Kansas; MCZ—Muse- only to the dipterist, but also to students of um of Comparative Zoology, Harvard University, physiology and behavior.’’ Obviously we concur and Cambridge, Massachusetts; NMW—Naturhistorisch- submit this faunal survey to document the diversity es Museum, Vienna, Austria; NRS—Naturhistoriska of shore flies on Plummers Island. Riksmuseet, Stockholm, Sweden; OHSU—Ohio Shore flies are diverse and often abundant in the State University, Columbus, Ohio; SMN—Staatlich- Potomac River gorge, but being small flies, usually es Museum fu¨r Naturkunde in Stuttgart, Stuttgart, less than 3 mm, they are not often collected or iden- Germany; ZIL—Zoological Institute, Lund Univer- tified. The North American fauna includes 464 spe- sity, Lund, Sweden; ZMO—Zoological Museum, cies (not including nomina dubia) and is now rea- University of Oslo, Oslo, Norway. sonably well known, although some genera, such as For each species, including junior synonyms that Allotrichoma Becker, have not been revised in the occur in the Nearctic Region, we list the status and last 50 years, and we are aware of several undescri- depository of the primary type(s) and have used the bed or unreported species. According to the Mary- abbreviations HT (ϭ holotype), LT (ϭ lectotype), land Natural Heritage Program, there are no shore and ST (ϭ syntype). In the synonymy provided for flies on the rare and endangered or threatened species each species, we include papers where nomenclatural in the state. changes were made and selected others that provide This faunal survey is based on study of 174 spec- information on the identification (especially if figures imens, representing 29 species in 17 genera. Twenty- were included) and natural history of the species. nine species represents 21% of the fauna from the Delmarva states. Specimens have been collected Systematics from April to October. We highlight the collecting atrimana (Loew) efforts of Raymond C. Shannon (1894–1945), who collected most of the 174 specimens. Ten species are atrimana Loew 1878:197 [United States. represented by specimens that he alone collected. District of Columbia and Texas; ST (, MCZ We provide a diagnosis for each species and note (11141)]. specific data for all specimens examined, such as the Leptopsilopa atrimana: Cresson 1942:126 [generic date of collection, collector, depository of the speci- combination].—1946a:155, 162 [review; key].— mens and the number of specimens we examined by Wirth 1965:741 [Nearctic catalog].—Deonier

173 174 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1965:502 [natural history].—Scheiring & Foote see, Texas, Virginia, Wisconsin, West Virginia). Neo- 1973:159 [natural history].—Steinly & Runyan tropical: Guatemala, West Indies (Cuba). 1979:139–147 [life history]; Mathis & Zatwarnicki Natural history.—Deonier (1965) and Scheiring & 1995:35 [world catalog]; 2006:90–95 [revision]. Foote (1973) reported respectively from studies con- ducted in Iowa and Ohio what little is known on the Diagnosis.—This species is distinguished from natural history of this species. They collected speci- congeners by the following combination of charac- mens from marsh-reed, sedge-meadow, and Eragros- ters: Head: Face above cruciate facial setae with tis-mat habitats in Iowa, and from limnic-wrack hab- transverse striae. Thorax: Wing infuscate over cross- itats in Ohio. In addition, Scheiring & Foote (1973) vein dm-cu; foreleg, including forecoxa and tarso- reared larvae in the laboratory on decaying lettuce meres, entirely black; mid- and hindfemora yellow- but noted that the larvae were probably feeding pri- ish, concolorous with tibiae. Abdomen: Male Termin- marily on saprophytic microorganisms. They also re- alia: Epandrium robustly developed, in posterior marked that adults were found abundantly in a flower view as an inverted U, slightly wider ventrally; in bed of chrysanthemums in Tucson, Arizona. lateral view subrectangular, essentially parallel-sided, Remarks.—Malloch (1934:314) wrote that this with height about twice width, in posterior view as ‘‘. . . species had been placed in a separate genus, an inverted U, ventral arms thickly developed; cercus Leptopsilopa Cresson, but the only characters cited in posterior view semihemispherical, in lateral view for the separation from Psilopa consist of the minute more or less lunate, slightly narrower ventrally than cross striation of the face, and the thickened foretarsi dorsally; postsurstylus robust, prominent, in posterior of both sexes, which hardly entitle the group to even view with dorsomedial projection, medial margin subgeneric segregation in our opinion.’’ Although we concave, bearing several setulae, in lateral view sub- have some evidence to substantiate the monophyly of rectangular with rounded angles; pregonite in lateral New World Leptopsilopa, we acknowledge that it view very elongate, rodlike to digitiform, almost par- may be an included lineage within the larger complex allel sided process, bearing 2 apical, moderately long now recognized as Psilopa (Mathis & Zatwarnicki setulae; postgonite in lateral view elongate, L-shaped, 2006). An obvious solution would be to combine spe- with base extended posterodorsally, apical extension cies of both genera into a single genus, perhaps rec- irregularly sided, posterior margin with subapical, an- ognizing subgenera or species groups. We have not gulate bulge, apical one-third bearing setulae, espe- done so for two reasons. First, the Old World species cially on subapical bulge, apex bluntly rounded, are very inadequately known, and second, no one has thumblike, bearing shorter setulae; subepandrial plate conducted comprehensive, comparative studies of all in lateral view narrow, apical one-third slightly included species, especially structures of the male curved and tapered to acute point, in posterior view and female terminalia. For the present, therefore, we very broadly and shallowly U-shaped with each arm defer acting upon a ‘‘merger’’ until the Old World spatulate; aedeagus in posterior view with basal two- fauna is better known and we have had an opportu- thirds as a trapezoid, narrower at base, slightly wider nity to analyze the evidence more comprehensively. apically, apical one-third as a bluntly pointed trian- In distribution, this is mostly an eastern species gle, in lateral view L-shaped, with extension narrow, and L. varipes is the western equivalent. There are, about twice length of basal portion, tapered gradu- however, a few records of L. atrimana from the West. ally, apex bluntly rounded; phallapodeme in lateral view irregularly triangular, keel pointed, in posterior (Paralimna) punctipennis (Wiedemann) view like an hourglass with base wider than apex; hypandrium in lateral view concave, pouchlike, punctipennis Wiedemann 1830:590 [not & pouch broadly rounded, posterior margin in ventral given (? United States); LT (designated by Cres- view conspicuously emarginate. son 1929:184), NMW]. Paralimna punctipennis Specimen examined.—MARYLAND. Montgom- : Cresson 1929:184 [generic combination].—1946b:229 [review].—Wirth 1965: ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 748 [Nearctic catalog].—Mathis & Zatwarnicki 23 Apr 1914, R. C. Shannon (1(; ANSP). 1995:124 [world catalog]. Distribution.—Nearctic: Canada (Manitoba, On- Paralimna appendiculata Loew 1862:138 [United tario, Quebec), United States (Alabama, Arizona, States. ‘‘Middle States’’ and ‘‘Georgia’’; ST (&, California, Connecticut, District of Columbia, Flori- MCZ (11134)]; Cresson 1929:184 [synonymy]. da, Georgia, Iowa, Illinois, Indiana, Kansas, Ken- tucky, Louisiana, Massachusetts, Maryland, Maine, Diagnosis.—This species is distinguished from Michigan, Minnesota, Mississippi, Missouri, Nebras- congeners by the following combination of charac- ka, Nevada, New Jersey, New Hampshire, New Mex- ters: Body generally bicolored, dorsum, especially ico, New York, North Carolina, Ohio, Oklahoma, frons and mesonotum, mostly grayish brown to Pennsylvania, Rhode Island, South Carolina, Tennes- brown, lateral surfaces mostly gray, sometimes sil- NUMBER 15 175

very white. Head: Arista bearing 11–13 dorsal Remarks.—This is apparently a rare species, and branches; eye round, about as wide as high; gena until we collected the male from Great Falls (Clay high, height subequal or slightly greater than length Pond) and Pocahontas State Park, Virginia, it was of basal flagellomere; frons mottled, dark to whitish only known from females. brown. Thorax: Scutellum guttate, mostly brown, es- pecially base of setae; anepisternum and katepister- fascitibia (von Roser) num mostly concolorous (anepisternum with dorsal margin sometimes brown), faintly yellowish gray to Notiphila fascitibia von Roser 1840:62 [Germany. silvery gray; wing mostly hyaline; crossvein dm-cu Wu¨rttemberg; HT &, SMN]. lacking infuscate halo; femora mostly gray; forefe- Hydrellia fascitibia: Becker 1905:198 [generic com- mur of male bearing anteroventral, comblike row of bination]. distinctly flattened, long setae; and posteroventral Hydrellia cruralis Coquillett 1910:131 [United surface of male forefemur bearing patch of long setae States. New Jersey. Burlington: Riverton; HT &, at basal 1/3. USNM (13102)].—Berg 1950:378–382 [immature Specimens examined.—MARYLAND. Montgom- stages, biology].—Deonier 1971:53–55 [revision, ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), immature stages; host plants].—Zatwarnicki 1988: 20 May–17 Aug 1914, 1988, 1989, A. Freidberg, F. 595 [synonymy]. K. Knab, R. C. Shannon, P. J. Spangler (3(,4&; USNM). Diagnosis.—This species is distinguished from Distribution.—Nearctic: United States (California, congeners by the following combination of charac- Delaware, District of Columbia, Florida, Georgia, Il- ters: Head: Frons tannish to grayish black with par- linois, Iowa, Maryland, Minnesota, New Jersey, New afrons lighter, usually grayer; ocellar setae present; 2 York, North Carolina, Ohio, South Dakota, Virginia, fronto-orbitals, anterior seta proclinate, posterior seta West Virginia). reclinate. Scape and pedicel dark brown, basal fla- Remarks.—This is one of the more common spe- gellomere partially to wholly yellow to orange; 5–9 cies on Plummers Island. It is often found abundantly (usually 6–7) aristal rays. Face in anterior view light on exposed muddy and/or sandy banks and shore- yellow brown or silvery, contrasted with silvery to lines. gray parafacial, face in lateral view very slightly con- vex with indistinct antennal grooves; maxillary pal- Hydrellia decens Cresson pus yellowish, broadly triangular, spathulate. Thorax: 3–4 presutural and 4 postsutural dorsocentral setae; Hydrellia decens Cresson 1931:107 [United States. mesonotum conspicuously darker than pleural areas, Maryland. Montgomery Co.: near Plummers Is- pleural areas substantially gray, not densely micro- & land; HT , USNM (43455)]; 1944:170 [re- tomentose and appearing velvety; forecoxa dark, gray view].—Wirth 1965:744 [Nearctic catalog]; Deon- to blackish brown; tibiae moderately grayish yellow ier 1971:55–56 [revision].—Mathis & Zatwarnicki to yellow; at least apical 2–3 tarsomeres dark. Ab- 1995:69 [world catalog]. domen: Male terminalia: Fused surstylus pentagonal, Diagnosis.—This species is distinguished from basal 3rd rectangular with lateral margins slightly an- congeners by the following combination of charac- gled laterally, apical two-thirds triangular, anterior ters: Head: Parafrons and mesofrons except for ocel- margin narrowly rounded and lacking a medial notch. lar triangle, densely microtomentose, appearing vel- Specimens examined.—MARYLAND. Montgom- vety black or dark reddish brown; ocellar setae pre- ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), sent; 2 fronto-orbitals, both setae proclinate. Pedicel 28 Apr–1 Sep 1914, 1962, K. Krombein, R. C. Shan- concolorous with frons; 9–11 aristal rays. Face in non (11(,7&; ANSP, USNM). anterior view light yellowish brown to yellowish Distribution.—Nearctic: Canada (Ontario, Que- gray; maxillary palpus blackish brown. Thorax: 3–4 bec), United States (Alabama, Alaska, Connecticut, presutural and 2 postsutural dorsocentral setae; me- Florida, Idaho, Illinois, Iowa, Kansas, Maryland, sonotum and abdomen (latter to a lesser extent) dark, Massachusetts, Michigan, New Jersey, Ohio, Penn- shiny, metallic bluish green to greenish blue; pleural sylvania, Texas, Virginia, West Virginia). Palearctic: areas faintly bluish gray; legs dark, femora with some Austria, Belgium, Bulgaria, Czech Republic, Finland, grayish microtomentum, otherwise blackish; tarsi France, Germany, Hungary, Italy, Poland, Romania, with some brownish orange at joints of tarsomeres. Sweden, Switzerland. Specimens examined.—MARYLAND. Montgom- Remarks.—This species has a Holarctic distribu- ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 2 tion and is especially widespread in the Nearctic Re- Sep 1914, R. C. Shannon (3&; ANSP, USNM). gion, although it apparently does not occur in the Distribution.—Nearctic: United States (Maryland, Southwest. On Plummers Island, this is the most fre- Virginia). quently collected species of Hydrellia. 176 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Hydrellia formosa Loew mature stages; host plant]; 1998:139–149 [revi- sion]. Hydrellia formosa Loew 1861:355 [United States. Hydrellia hypoleuca Loew 1862:151 [United States. & ‘‘Pennsylvania’’; HT , MCZ (11153)].—Cresson ‘‘Middle States’’; ST &, MCZ (11158)].—Grigar- 1944:163 [review].—Deonier 1971:61–63 [revi- ick 1959:4 [synonymy]. sion], 1998:130–135 [revision]; Wirth 1965:744 Hydrellia obscuriceps Loew 1862:152 [United [Nearctic catalog].—Mathis & Zatwarnicki 1995: States. ‘‘Middle States’’; ST (, MCZ (11157)].— 72 [world catalog]. Grigarick 1959:4 [synonymy]. Diagnosis.—This species is distinguished from Hydrellia scapularis Loew 1862:153 [United States. & congeners by the following combination of charac- ‘‘United States’’; ST , MCZ 11155].—Grigarick ters: Head: Frons mostly densely microtomentose, 1959:4 [synonymy]. appearing velvety black except for semi-glossy ocel- Diagnosis.—This species is distinguished from lar triangle; ocellar setae lacking; 2 fronto-orbitals, congeners by the following combination of charac- anterior seta proclinate, posterior seta reclinate. An- ters: Head: Frons mostly light gray to bluish gray tenna generally black, somewhat concolorous with microtomentose, parafrons darker brown; ocellar se- frons except for some yellowish coloration along tae present; 2 fronto-orbitals, anterior seta proclinate, venter of basal flagellomere; 7–11 (usually 9) aristal posterior seta reclinate. Antenna usually dark grayish rays. Face in anterior view unicolorous, silvery to brown; 4–8 (usually 5–6) aristal rays. Face in ante- slightly grayish white microtomentose, in lateral view rior view variable, light to dark metallic yellowish slightly receding, with indistinct antennal grooves; brown, in lateral view nearly vertical to distinctly maxillary palpus yellowish, short, length about or convex, very rounded, medial elevation to a distinct less than twice width. Thorax: 3–4 presutural and 2– carina on dorsal portion, antennal grooves usually 3 postsutural dorsocentral setae; scutellar disc, noto- distinct; maxillary palpus yellowish, broadly trian- pleuron (and adjacent supra-alar area), and ventral gular, spathulate. Thorax: 3–5 presutural and 2–3 area of anepisternum and katepisternum densely mi- postsutural dorsocentral setae; mesonotum conspicu- crotomentose, black, appearing velvety; femora and ously darker than pleural areas, not densely micro- tibiae concolorous, black; tarsi yellowish. Abdomen: tomentose and appearing velvety; wing length usu- Male terminalia: Fused surstylus shallowly bilobed, ally less than 0.8 mm; forecoxa dark, gray to blackish widely and moderately shallowly concave medially, brown; tibiae dark gray to brown, generally conco- width only slightly longer than length. lorous with femora and tarsi. Abdomen: Male ter- Specimens examined.—MARYLAND. Montgom- minalia: Fused surstylus only slightly wider than ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 5 long, broadly bilobed, somewhat similar to H. valida Jun–5 Aug 1913, 1963, W. L. McAtee, R. C. Shan- but with deep, medial emargination slightly narrower non, G. C. Steyskal (2(,2&; USNM); Plummers Is- and deeper. land (near; at light; 38Њ58.2ЈN, 77Њ10.6ЈW), 23 May– Specimens examined.—MARYLAND. Montgom- 26 Jul 1914, 1915, R. C. Shannon (4(,3&; USNM). Њ58.2ЈN, 77Њ10.6ЈW), Distribution.—Nearctic: Canada (Nova Scotia, ery Co.: Plummers Island (38 Ontario, Quebec), United States (Alabama, Arkansas, 17 May–8 Sep 1907, 1918, 1963, K. V. Krombein, ( & Connecticut, District of Columbia, Florida, Georgia, W. L. McAtee, R. C. Shannon (3 ,4 ; USNM). Indiana, Iowa, Kansas, Louisiana, Maine, Maryland, Distribution.—Australasian/Oceanian: Australia Massachusetts, Michigan, Mississippi, Missouri, (Queensland). Nearctic: Bermuda, Canada (Alberta, New Jersey, New York, North Carolina, Ohio, Penn- British Columbia, Manitoba, Nova Scotia, Ontario, sylvania, South Carolina, Tennessee, Texas, Virginia, Quebec, Saskatchewan), United States (Alaska, Ala- Vermont, West Virginia). bama, Arkansas, California, Connecticut, District of Remarks.—This species is easily recognized and is Columbia, Florida, Georgia, Idaho, Indiana, Iowa, one of the more beautiful species of shore flies. The Kansas, Louisiana, Maine, Maryland, Massachusetts, dense patches of microtomentum that appear velvety Michigan, Minnesota, Missouri, Mississippi, Mon- are especially attractive. The species is fairly com- tana, Nevada, New Hampshire, New Jersey, New mon and widespread in temperate eastern North Mexico, New York, North Carolina, North Dakota, America, including Plummers Island. Ohio, Oregon, Pennsylvania, South Carolina, South Dakota, Tennessee, Texas, Utah, Virginia, Vermont, Washington, West Virginia, Wisconsin, Wyoming). Hydrellia griseola (Falle´n) Neotropical: Colombia. Oriental: Nepal, Philippines, Notiphila griseola Falle´n 1813:250 [Sweden; ST Taiwan. Palearctic: Afghanistan, Algeria, Austria, (sex?), ZIL]. Azores, Belgium, Canary Islands, China (Hefei, Xin- Hydrellia griseola: Macquart 1835:523 [generic ing, NE Tibet), Cyprus, Czech Republic, Denmark, combination].—Deonier 1971:63–69 [revision, im- Egypt, Estonia, Faroe Islands, Finland, France, Ger- NUMBER 15 177 many, Great Britain, Greece, Hungary, Iceland, Italy, Hydrellia ischiaca Loew Japan (Hokkaido, Honshu), Macedonia, Madeira Is- lands, Malta, Morocco, Netherlands, Norway, Oman, Hydrellia ischiaca Loew 1862:150 [United States. & Poland, Romania, Russia (European Territory), Slo- ‘‘Middle States’’; LT (designated by Deonier vakia, Spain, Sweden, Switzerland, Tunisia, Ukraine, 1971:72), MCZ (11154)].—Cresson 1944:165 [re- Yugoslavia. view].—Deonier 1971:71–76 [revision, immature Remarks.—In temperate zones around the world, stages; host plants]; 1998:156–164 [revision].— this is one of the most common shore flies and at Wirth 1965:744 [Nearctic catalog].—Mathis & Za- times it has been the cause of considerable damage twarnicki 1995:78–79 [world catalog]. in rice (Grigarick 1959). The species is also one of Diagnosis.—This species is distinguished from the more common species in the Delmarva States congeners by the following combination of charac- where numerous host plants of the species also occur. ters: Head: Parafrons dark brown, frontal vitta and Deonier (1971, 1998) noted that there is considerable fronto-orbits light gray to greenish gray; ocellar setae variation in this species, especially in the color of the present; 2 fronto-orbitals, anterior seta proclinate, face. Structures of the male terminalia are virtually posterior seta reclinate. Antenna blackish brown to identical in all color morphs. black; 7–10 (usually 7–8) aristal rays. Face in ante- rior view metallic yellowish brown to silvery white, Hydrellia insulata Deonier in lateral view somewhat convex at midfacial level, with shallow to distinct antennal grooves; maxillary Hydrellia insulata Deonier 1971:71 [United States. palpus yellowish, roundedly spatulate. Thorax: 4–5 Maryland. Montgomery Co.: Plummers Island, Po- presutural and 2 postsutural dorsocentral setae; scu- ( tomac River; HT , USNM (70535)]; 1998:153– tellar disc, notopleuron (and adjacent supra-alar 155 [revision].—Mathis & Zatwarnicki 1995:78 area), and ventral area of anepisternum and katepis- [world catalog]. ternum not densely microtomentose and appearing velvety; forecoxa yellowish; femora grayish black; Diagnosis.—This species is distinguished from tibiae yellowish to blackish brown; tarsi, except for congeners by the following combination of charac- apical 2 blackish brown tarsomeres, yellowish. Ab- ters: Head: Parafrons slightly to distinctly densely domen: Male terminalia: Fused surstylus longer than microtomentose, appearing velvety black; frontal vit- wide, parallel sided over most of length, apical one- ta and fronto-orbits golden brown to moderately fourth obtusely tapered to notched apex. brown; ocellar setae present; 2 fronto-orbitals, both Specimens examined.—MARYLAND. Montgom- setae proclinate. Antenna dark brown; 7–10 (usually ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 5 8) aristal rays. Face in anterior dark, yellowish gold Aug–5 Sep 1913, 1915, W. L. McAtee, R. C. Shan- to silvery light gray, in lateral view slightly convex non (4&; USNM); Plummers Island (near; 38Њ58.2ЈN, with slightly, rounded, medial elevation on dorsal 77Њ10.6ЈW), 22 May 1915, R. C. Shannon (1&; half and with distinct antennal grooves; maxillary USNM). Tho- palpus yellowish, broadly triangular, spathulate. Distribution.—Nearctic: Canada (Nova Scotia, rax: 3–5 presutural and 2 postsutural dorsocentral se- Ontario, Quebec), United States (Alaska, California, tae; mesonotum dark brown, pleural areas grayish Connecticut, Georgia, Indiana, Illinois, Iowa, Maine, brown, not densely microtomentose and appearing Maryland, Massachusetts, Michigan, Minnesota, velvety; forecoxa dark, gray to blackish brown. Ab- New Hampshire, New York, North Carolina, Ohio, domen: Male terminalia: Fused surstylus about as Pennsylvania, South Carolina, Tennessee, Texas, Vir- long as wide, lateral margins essentially parallel sid- ginia, West Virginia). Palearctic: Czech Republic, ed, anterior margin broad, like steps, with lateral, Germany, Great Britain, Hungary, Italy, Japan (Hok- short lobes then medial area flat except for narrowly kaido), Poland, Romania, Russia (Far East), Sweden. rectangular, medial notch. Remarks.—Among species of Hydrellia, this is Specimens examined.—MARYLAND. Montgom- one of the more common and widespread in the Del- Њ Ј Њ Ј ery Co.: Plummers Island (38 58.2 N, 77 10.6 W); marva States. There is considerable variation, how- wasp nest, Potomac River, 7 Aug 1962, K. V. Krom- ever, in structures of the male terminalia, and the sta- ( & bein (6 ,5 ; ANSP, USNM). tus of these varying populations has not been clari- Distribution.—Nearctic: United States (Mary- fied. land). Remarks. —Thus far, this species has been re- Hydrellia tibialis Cresson trieved only from nests of wasps, and the few spec- imens we examined were collected in conjunction Hydrellia tibialis Cresson 1917b:341 [United States. with studies on wasps of Plummers Island and the Idaho. Latah: Moscow; HT (, ANSP (6141)].— provisioning of wasp nests. Deonier 1971:99–102 [revision, immature stages; 178 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON

host plant]; 1998:230–237 [revision].—Wirth Diagnosis.—This species is distinguished from 1965:744 [Nearctic catalog].—Mathis & Zatwar- congeners of the subgenus by the following nicki 1995:91 [world catalog]. combination of characters: Thorax: Crossveins, es- pecially dm-cu, somewhat to conspicuously clouded, Diagnosis.—This species is distinguished from infuscate. Abdomen: Tergites somewhat microtomen- congeners by the following combination of charac- tose, especially along posterior margins of tergites. ters: Head: Only parafrons densely microtomentose, appearing velvety black, anterior portion of mesof- Male terminalia as follows: Epandrium and fused rons like ocellar triangle; ocellar setae present; 2 presurstylus in lateral view oriented along same fronto-orbitals, anterior seta proclinate, posterior seta plane, nearly straight, not projected or angulate; reclinate. Antenna mostly dark brown except dorso- epandrium in lateral view nearly parallel sided except lateral portion of pedicel densely microtomentose, for wide, anterior projection, oriented anteroventrally, appearing velvety black, similar to parafrons; 5–6 ar- forming a moderately deep U-shaped concavity with istal rays (rarely 7). Face in anterior view unicolo- presurstylus, epandrium in posterior view about a rous, silvery white microtomentose, in lateral view wide as high, roundedly quadrate, usually with nearly vertical but with low, rounded, medial carina paired, short projections immediately ventrad of cer- and shallow but distinct antennal grooves; maxillary cal cavity; presurstylus in lateral view as a broadly palpus dark brown to black, long blade or cleaver rounded, ventral projection, bearing a short, truncate, shaped. Thorax: 4–5 presutural and 2 postsutural dor- subapical, anterior projection, in posterior view socentral setae; mesonotum, pleural areas, and ab- height about equal to that of cercal cavity, lateral domen dark brown to dark greenish gray; legs dark margins conspicuously sinuous, ventral margin emar- greenish black to black. Abdomen: Male terminalia: ginate, like an inverted W, setulose along mediob- Fused surstylus trapezoidal, broadly truncate apically, lique margins, medial surface variable, usually mod- as long as wide. erately produced as a single, broadly produced pro- Specimens examined.—MARYLAND. Montgom- jection but often nearly flat or with 2 symmetrical, ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), shallow projections; postsurstylus in lateral view nar- 28 Apr 1914, R. C. Shannon (1(; USNM). rowly rounded basally, becoming wider apically, api- Distribution.—Nearctic: Canada (Alberta, British cal margin truncate, bearing serulae along ventral Columbia, Manitoba, Ontario, Quebec), United margin and with a robust, short setula at apicoventral States (Alabama, Alaska, Arizona, Arkansas, Cali- corner, in ventral view broad basally, irregularly ta- fornia, Colorado, Delaware, District of Columbia, pered to apex, bearing setulae along apical margin, Florida, Idaho, Illinois, Indiana, Iowa, Kansas, Lou- robust, short setula inserted subapically; subepandrial isiana, Maine, Maryland, Michigan, Minnesota, Mis- plate in lateral view rodlike, elongate, very shallowly sissippi, Missouri, Nebraska, New Hampshire, New arched, in ventral view shorter than lateral view, Jersey, New York, North Carolina, Ohio, Oklahoma, more rodlike, more evenly, shallowly arched; aedea- Oregon, Pennsylvania, Tennessee, Texas, Virginia, gus in lateral view tubular, although with ventral mar- Washington, West Virginia, Wyoming). Neotropical: gin stepwise angulate, in ventral view symmetrically Bolivia, Chile, Costa Rica, Mexico, Trinidad and To- tubular, with basal one-third tapered to slightly more bago, West Indies (Dominica). Palearctic: Austria, narrow base, apical two-thirds very slightly expand- Finland. ed, almost parallel sided, apex broadly attenuate to Remarks.—This is one of the most common spe- broadly rounded; phallapodeme in lateral view with cies in the Nearctic Region; it also has been reported keel elongate, moderately expanded, and greatly from Europe and extends into the Neotropical Re- slanted ventrally, extension to base of aedeagus elon- gion. gate, almost parallel sided, with a short, medial pro- jection at attachment end with hypandrium, in ventral Notiphila (Dichaeta) caudata Falle´n view irregularly hour-glass shaped, hypandrial attach- Notiphila caudata Falle´n 1813:249 [Sweden. Ska˚ne; ment projection flatter and wider than aedeagal at- HT (, ZIL].—Mathis & Zatwarnicki 1995:103 tachment, aedeagal end U-shaped, angularly concave, [world catalog]. hypandrial and keel end broadly produced, expanded, Dichaeta caudate: Meigen 1830:62 [generic combi- apex nearly straight, truncate; gonite (probably the nation]; Cresson 1917a:65–66 [review]; 1946b: pregonite) in ventral view T-shaped, very broad ba- 230–231 [review].—Wirth 1965:748 [Nearctic cat- sally and with a digitiform projection that bears 2 alog].—Eastin & Foote 1971:271–279 [life history, apically setulae, in lateral view L-shaped with a immature stages].—Krivosheina 1986:810–811 broad base and digitiform projection; hypandrial arm [revision], 1993:229–230 [immature stages]. in lateral view narrowed and curved on apical half Dichaeta caudata caudate: Sturtevant & Wheeler with apex sharply recurved, darkened, well sclero- 1954:236–237 [subspecific status]. tized, in ventral view transversely bandlike with lat- NUMBER 15 179

eral extensions that are recurved and bear numerous evenly, pocket formed by arms broadly U-shaped; setulae. gonite short, narrow, bearing 2–3 apical setulae; bas- Specimens examined.—MARYLAND. Montgom- iphallus about as wide as long, short, apex expanded. ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 5 Specimens examined.—MARYLAND. Montgom- May 1915, R. C. Shannon (1(,1&; USNM). ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW; Distribution.—Nearctic: Canada (Manitoba, On- cell in wasp nest), 7 Aug 1962, K. V. Krombein (7(, tario, Quebec, Saskatchewan). United States (Alaska, 9&; USNM). Colorado, Connecticut, District of Columbia, Florida, Distribution.—Nearctic: United States (Delaware, Idaho, Illinois, Indiana, Iowa, Louisiana, Maryland, District of Columbia, Florida, Indiana, Iowa, Ken- Massachusetts, Michigan, Minnesota, Mississippi, tucky, Maryland, New Jersey, New York, North Car- Montana, Nebraska, New Hampshire, New Jersey, olina, Ohio, Virginia). New York, North Dakota, Ohio, Oregon, Pennsyl- vania, South Dakota, Utah, Vermont, Virginia, Wis- Diclasiopa lacteipennis (Loew) consin, Washington). Palearctic: Austria, Belgium, lacteipennis Loew 1862:145 [United Czech Republic, Denmark, Estonia, Finland, France, States. District of Columbia: ‘‘Washington’’ [D. Germany, Greece, Great Britain, Hungary, Italy, Ja- C.]; LT ( (designated by Zatwarnicki 1991:303), pan (Hokkaido, Honshu), Kazakhstan, Kyrgyzstan, MCZ (11146)]. Netherlands, Poland, Romania, Russia (European Diclasiopa lacteipennis: Cresson 1929:167 [generic Territory, Far East), Slovakia, Spain, Sweden, Swit- combination].—Wirth 1965:739 [Nearctic cata- zerland, Tadzhikistan, Turkmenistan, Ukraine, Yu- log].—Mathis & Zatwarnicki 1995:164 [world cat- goslavia. alog]. Remarks.—Some intraspecific and age-related var- iation has been observed in this species. We have also Diagnosis.—This species is distinguished from noted some variation in the shape of the posterior or congeners by the following combination of charac- apical margin of the fused surstylar process, espe- ters: Head: Face and parafacial of male yellow; lat- cially the medial portion. The medial portion in some eral vertical series of facial setae moderately well de- specimens can be more slightly extended, very shal- veloped, longest seta about half length of setae in lowly bi- or trilobed, slightly narrower or even trun- medial row, inserted between level of medial setae. cate to very shallowly concave. The variation is Thorax: Postsutural supra-alar setae, anterior seta slight and was not expressed in any detectable pat- small, posterior seta well developed, nearly twice tern. The few specimens demonstrating variation usu- length of presutural supra-alar seta. Abdomen: Third ally occurred microsympatrically with greater num- abdominal tergite long, comparable to fourth. Male bers of more typical conspecific specimens. terminalia: Epandrium in posterior view as an in- This is the most common and widespread species verted U, somewhat angulate at corners, very thin in the subgenus Dichaeta and its distribution is Hol- dorsally, becoming wider ventrally, ventral margin arctic. For example, this is the only known congener curved medially; cerci in posterior view irregularly of the subgenus in the western Palearctic Region as oval, dorsomedial angle somewhat pointed, ventral well as North America. margin rounded; aedeagus in lateral view L-shaped, robust, truncate apically, in ventral view asymmetri- Notiphila (Notiphila) carinata Loew cal, right basal lobe much larger than left lobe; phal- lapodeme in lateral view with wide, enlarged keel, Notiphila carinata Loew 1862:137 [United States. process toward base of aedeagus elongate, narrow, ‘‘Middle States’’; HT &, MCZ (11133)].—Wirth slender, process toward hypandrium short, digitiform, 1965:746 [Nearctic catalog].—Deonier et al. 1979: in ventral view narrow, almost parallel sided; gonite 798–814 [biology].—Mathis 1979:37–39 [revi- in lateral view with recurved apex, emargination of sion].—Mathis & Zatwarnicki 1995:05 [world cat- recurve bearing a setula, otherwise shoelike; hypan- alog]. drium in lateral view like a parallelogram, moderate- Diagnosis.—This species is distinguished from ly deeply incised anteriorly and posteriorly, in ventral congeners by the following combination of charac- view more or less broadly U-shaped, lateral arms ters: Head: Facial ratio 0.84 or less; facial setae slightly flared, rounded and bearing short, stoutly weakly developed, hairlike, smaller than genal seta; processes medially and anterolaterally. gena short, gena-to-eye ratio less than 0.21 (averag- Specimens examined.—MARYLAND. Montgom- ing 0.15). Thorax: Mesonotum immaculate; lateral ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 9 margin of scutellum essentially concolorous with May–27 Sep 1914, 1949, C. W. Sabrosky, R. C. scutellar disc; anepisternum usually with a rather Shannon (1(,2&; ANSP, USNM). well-defined, darkened area. Abdomen: Male termin- Distribution.—Afrotropical: Kenya, Sudan. Nearc- alia: Surstylar arms projected straight, not tapered tic: Canada (Ontario), United States (Alabama, Ari- 180 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON zona, Colorado, District of Columbia, Georgia, Illi- 14 May–24 Aug 1912, 1913, 1914, J. R. Malloch, R. nois, Indiana, Kansas, Iowa, Massachusetts, Mary- C. Shannon (8(,9&; ANSP, USNM). land, Michigan, Minnesota, New Jersey, New Mexi- Distribution.—Afrotropical: Cape Verde Islands, co, New York, Ohio, Oregon, Pennsylvania, South Kenya, Rodriguez Islands, Zaire. Nearctic: Canada Carolina, South Dakota, Tennessee, Texas, Utah, Vir- (Newfoundland, Ontario, Quebec), United States ginia, Washington, West Virginia). Palearctic: Al- (Alaska, Alabama, Arkansas, Arizona, California, geria, Austria, Belgium, Bulgaria, Croatia, Czech Re- Connecticut, Delaware, District of Columbia, Florida, public, Finland, France, Germany, Great Britain, Georgia, Illinois, Indiana, Iowa, Kansas, Louisiana, Greece, Hungary, Italy, Netherlands, Morocco, Po- Massachusetts, Maryland, Maine, Michigan, Missou- land, Romania, Slovenia, Spain, Sweden, Switzer- ri, Nebraska, New Hampshire, New Jersey, New land, Yugoslavia. Mexico, Nevada, New York, North Carolina, North Dakota, Ohio, Oregon, Pennsylvania, South Carolina, (Falle´n) South Dakota, Tennessee, Texas, Utah, Virginia, Washington, West Virginia). Neotropical: Argentina, Notiphila obscurella Falle´n 1813:251 [Sweden; ST Bahamas, Bolivia, Brazil, Chile, Colombia, Costa &, ZIL]. Rica, Ecuador, Guatemala, Mexico (Veracruz), Pan- Discocerina obscurella: Haliday 1839:224 [generic ama, Paraguay, Peru, Venezuela, West Indies (Cuba, combination].—Foote & Eastin 1974:401–408 Dominica, Jamaica, Puerto Rico, St. Thomas). Pale- [preimaginal stages].—Zatwarnicki & Mathis arctic: Austria, Algeria, Azores, Belgium, Bulgaria, 2001:22 [obscurella group]. Canary Islands, Croatia, Czech Republic, Estonia, Psilopa (Clasiopa) obscurella: Stenhammar 1844: Finland, France, Germany, Great Britain, Hungary, 160 [generic combination]. Iran, Italy, Macedonia, Morocco, Netherlands, Po- Clasiopa obscurella: Schiner 1864:57 [generic com- land, Romania, Russia (European Territory, Far bination]. East), Slovakia, Slovenia, Spain, Sweden, Switzer- Discocerina (Discocerina) obscurella: Wirth 1965: land, Turkey, Yugoslavia. 738 [Nearctic catalog].—Mathis & Zatwarnicki Remarks.—This may be the most common and 1995:166–168 [world catalog]. widespread species in the Holarctic Region and its Discocerina parva Loew 1862:146 [United States. occurrence on Plummers Island was expected. District of Columbia: ‘‘Washington’’; HT &, MCZ (11149)].—Sturtevant 1923:11 [synonymy]. Hydrochasma leucoproctum (Loew) Discocerina parva variety nigriventris Cresson 1916: Discocerina leucoprocta Loew 1861:355 [United 148 [United States. California. Alameda: Berkeley States. ‘‘Maryland’’; HT &, MCZ (11148)]. Hills; HT (, ANSP (6100)].—Cresson 1949:260 Hydrochasma leucoproctum: Cresson 1942:113 [ge- [synonymy]. neric combination, as H. leucostoma].—Wirth Discocerina (Discocerina) obscurella nitidiventris: 1965:739 [Nearctic catalog].—Mathis & Zatwar- Cresson 1946a:143 [subspecies status]. nicki 1995:183 [world catalog]. Diagnosis.—This species is distinguished from Diagnosis.—This species is distinguished from congeners by the following combination of charac- congeners by the following combination of charac- ters: Head: Antenna mostly yellowish although dor- ters: Head: Gena moderately high, height usually sal surface of pedicel and basal flagellomere blackish; slightly higher than length of basal flagellomere. Tho- facial series of setae 3 or more; face lacking vertical rax: Wing hyaline. Abdomen: Tergites lacking stripes, entirely microtomentose, microtomentum wedgelike marking laterally but tergites 2–4 with gray to silvery gray; parafacial bearing setulae, par- wide medial area extensively dark slate gray to black; afacial microtomentum creamy whitish, slightly con- fifth tergite mostly light gray to silvery gray with trasted in coloration with facial microtomentum; posterior margin blackish brown to slate black, sim- frons mostly microtomentose, lacking metallic blue ilar to coloration of medial area on tergites 1–4; me- or green stripes. Thorax: Mesonotum moderately dial coloration on tergites 1–4 wide, occupying most densely microtomentose, grayish brown; pleural ar- of dorsum, dark grayish to slate black. eas gray; supra-alar seta present; and anterior noto- Specimens examined.—MARYLAND. Montgom- pleural seta inserted in anterior notopleural angle; fo- ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), recoxa mostly dark colored, brownish to blackish 30 May–4 Jul 1907, 1914, W. L. McAtee (2&; brown; male forefemur bearing about 4 long, peglike USNM). setae apically along posteroventral surface; abdomen Distribution.—Nearctic: United States (Delaware, subshiny gray to grayish black. District of Columbia, Florida, Georgia, Indiana, Kan- Specimens examined.—MARYLAND. Montgom- sas, Maryland, Michigan, New Jersey, Pennsylvania, ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), Texas, Virginia, West Virginia). Neotropical: Argen- NUMBER 15 181

tina, Bolivia, Colombia, Costa Rica, Ecuador, Gua- plant].—Mathis & Zatwarnicki 1995:132 [world temala, Mexico, Panama, Venezuela, West Indies (Ja- catalog]. maica). (Ochteroidea) granulose: Wirth 1965: Remarks.—This is the more common congener in 735 [Nearctic catalog]. the region and in eastern North America generally from Michigan and Pennsylvania south into northern Diagnosis.—This species is distinguished from South America. There are two undescribed congeners congeners by the following combination of charac- in the region that also may occur on Plummers Is- ters: Head: Arista with 5–6 dorsal branches; face at land. narrowest point between eyes narrow, width subequal to combined length of pedicel and basal flagellomere; face medially with vertical, granulose stripe; face in Athyroglossa (Athyroglossa) glaphyropus Loew lateral view evenly arched, lacking midheight prom- Athyroglossa glaphyropus Loew 1878:197 [United inence; genal height at ventral margin of eye short, States. ‘‘Texas’’; LT ( (designated by Cresson about equal to length of basal flagellomere; basal fla- 1922:340), MCZ (11145)].—Cresson 1946a:134 gellomere concolorous with black pedicel. Thorax: [review].—Wirth 1965:735 [Nearctic catalog].— Mesonotum conspicuously granulose, shiny. Wing Mathis & Zatwarnicki 1995:132 [world catalog]. hyaline; halter black. Foretibia entirely black; mid- Ochtheroidea glaphyropus: Thaxter 1917:680–681 and hindtibiae yellowish. Abdomen: Abdominal ter- [generic combination].—Cresson 1918:61 [re- gites not creased dorsolaterally. view]; 1922:340–341 [revision]. Specimens examined.—MARYLAND. Montgom- ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 8 Diagnosis.—This species is distinguished from its May–8 Aug 1914, 1922, J. R. Malloch, R. C. Shan- congeners by the following combination of charac- non (3(,6&; ANSP, USNM). ters: Head: Arista with 7–8 dorsal branches; face at Distribution.—Nearctic: Canada (Quebec), United narrowest point between eyes narrow, width subequal States (District of Columbia, Idaho, Indiana, Iowa, to combined length of pedicel and basal flagellomere; Massachusetts, Maryland, Michigan, New York, face almost entirely shiny, smooth, lateral margin North Carolina, Ohio, Pennsylvania, Virginia, Ver- next to parafacial with pits into which setulae are mont, West Virginia). Neotropical: West Indies (Gre- inserted; face in lateral view evenly and shallowly nada). arched; genal height at ventral margin of eye short, about equal to length of basal flagellomere; basal fla- Allotrichoma (Allotrichoma) simplex (Loew) gellomere concolorous with black pedicel. Thorax: Mesonotum very finely granulose, shiny. Wing hya- Discocerina simplex Loew 1861:354 [United States. line; halter whitish. Foretibia entirely black; mid- and ‘‘Maryland’’; ST &, MCZ (11150)]. hindtibiae yellowish. Abdomen: Abdominal tergites Allotrichoma simplex: Johnson 1925:271 [generic not creased dorsolaterally. combination].—Cresson 1942:108 [list, general Specimens examined.—MARYLAND. Montgom- distribution in North America]; 1949:260 [synon- ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), ymy].—Deonier 1965:501 [natural history].— 24 Aug 1913, R. C. Shannon (1&; USNM). Wirth 1965:736 [Nearctic catalog].—Runyan & Distribution.—Nearctic: United States (Alabama, Deonier 1979:123–137 [discussion, figs. of head District of Columbia, Florida, Georgia, Illinois, Iowa, and immature stages].—Mathis & Zatwarnicki Louisiana, Maryland, New York, Pennsylvania, 1995:153 [world catalog]. South Carolina, Texas, Virginia). Neotropical: Ar- gentina, Belize, Bolivia, Brazil, Costa Rica, Ecuador, Diagnosis.—This species is distinguished from Guatemala, Guyana, Mexico (Colima, Nayarit, San congeners by the following characters: Abdomen: Luis Potosı´, Tabasco), Panama, Peru, Trinidad and Male terminalia: Epandrium in posterior view round- Tobago, West Indies (Cuba, Dominica, Dominican ed dorsally, constricted laterally at base of surstyli; Republic, Grenada, Guadeloupe, Jamaica, Puerto epandrium in lateral view somewhat rectangular dor- Rico, St. Croix). sally with anterior and posterior margins parallel-sid- ed, ventral portion with an anterior bulge and a nar- Athyroglossa (Athyroglossa) granulosa (Cresson) row, sharp incision at about level or base of surstylus; surstyli (ventral extensions of epandrium) in posterior Ochteroidea granulosa Cresson 1922:341 [United view as relatively wide at base third, thereafter ven- States. Pennsylvania. Delaware: Swarthmore trally abruptly narrowed, especially along medial (creek); HT (, ANSP (6345)]. margin, to form an elongate process that is slightly Athyroglossa granulose: Cresson 1944:160 [generic spatulate apically, apical portion bearing several short combination].—Grimaldi & Jaenike 1983:86 [host setulae; surstylus in lateral view moderately narrow, plant].—Bunyard & Foote 1990:118 [host slightly expanding with apex obliquely pointed and 182 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON bearing several short setulae; cerci in posterior view ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 5 approximate dorsally then extended laterally to form Oct 1913, W. L. McAtee (1&; USNM). a lyrelike structure that curves laterally than ventrally Distribution.—Nearctic: Canada (Alberta, Mani- and medially, apex of cercus bearing 4 short setulae; toba, Northwest Territories, Ontario, Quebec, Sas- cercus in lateral view shallowly sinuous, nearly par- katchewan), United States (Colorado, Connecticut, allel sided, apex slightly expanded, bluntly rounded; Delaware, District of Columbia, Idaho, Illinois, In- aedeagus in ventral view elongate, narrowly ovate, diana, Iowa, Kansas, Maryland, Maine, Massachu- very slightly tapered apically, in lateral view elon- setts, Michigan, Minnesota, Missouri, Montana, Ne- gate, nearly parallel sided, apex bluntly rounded; braska, New Hampshire, New Jersey, New Mexico, phallapodeme in lateral view triangular, keel abruptly New York, North Dakota, Ohio, Oklahoma, Oregon, tapered to narrow, digitiform apex; gonite in ventral Pennsylvania, Rhode Island, South Dakota, Tennes- view barlike, slightly wider basally, apex with short, see, Texas, Utah, Vermont, Virginia, Washington, lateral process, apex bearing setula, in lateral view West Virginia, Wisconsin, Wyoming). wide basally, gently but distinctly arched, tapered to apical point. Zeros flavipes (Williston) Specimens examined.—MARYLAND. Montgom- ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), ? flavipes Williston 1896:403 [West Indies. St. ( 12 Apr 1914, W. L. McAtee (1(,1&; ANSP). Vincent; LT (designated by Mathis & Edmiston Distribution.—Nearctic: United States (California, 1991:834), BMNH].—Aldrich 1905:625 [Nearctic Delaware, Iowa, Maryland, Massachusetts, Nevada, catalog].—Jones 1906:184 [catalog]. New Jersey, Ohio, Vermont, Virginia, Washington). Zeros flavipes: Cresson 1943:14 [generic combina- Palearctic: Austria, Bulgaria, China (Tibet), Czech tion], 1947:45 [list].—Wirth 1965:746 [Nearctic Republic, France, Hungary, Israel, Italy, Morocco, catalog].—Scheiring & Connell 1979:609–612 Poland, Spain, Russia (European Territory), Switzer- [egg].—Mathis & Edmiston 1991:834 [review of land, Yugoslavia. Williston’s St. Vincent species].—Woodley & Hil- Natural history.—Deonier (1965:501) found this burn 1994:32 [list, Bermuda].—Mathis & Zatwar- species to be ‘‘occasional’’ in mud shore and sand nicki 1995:189 [world catalog].—Mathis 1997:60 shore habitats but rare in sedge meadow and Era- [review, Belize].—Hollmann-Schirrmacher 1998: grostis mat habitats. 26–29 [revision]. Remarks.—This species apparently has a Holarctic Diagnosis.—This species is distinguished from distribution, and Palearctic specimens have been congeners by the following combination of charac- named at least three times, with A. simplex being the ters: Thorax: Vein R2ϩ3 nearly straight, not distinctly senior synonym. In males of A. simplex the cercus undulating; cells R2ϩ3 and R4ϩ5 with 3 infuscate spots has four apical setulae, and the mesonotal stripes are or transverse bars, these generally not connected to more evident in A. strandi than in A. simplex. form longer, transverse bands; crossvein dm-cu pre- In the Nearctic Region, this species is common and sent. Abdomen: Tergites shiny black, mostly bare of widespread. microtomentum. Specimens examined.—MARYLAND. Montgom- anatolikos Clausen ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 6 Ochthera anatolikos Clausen 1977:468 [United Jul 1963, G. C. Steyskal (1&; USNM). States. Ohio. Franklin County; HT (, OHSU].— Distribution.—Afrotropical: Zaire. Nearctic: Ber- Mathis & Zatwarnicki 1995:144 [world catalog]. muda, Canada (Ontario), United States (Florida, Iowa, Maryland, Ohio, Texas, Virginia, West Virgin- Diagnosis.—This species is distinguished from ia). Neotropical: Argentina, Belize, Bolivia, Brazil, congeners, especially O. tuberculosa, by the follow- Colombia, Costa Rica, Ecuador, Mexico, Panama, ing combination of characters: Body length 5.00– Trinidad and Tobago, West Indies (Cuba, Dominica, 6.10 mm. Head: Face lacking distinct, shiny black Dominican Republic, Grenada, Haiti, Jamaica, Puerto spots, bars, stripes, or combination thereof; clypeus Rico, St. Lucia, St. Vincent). large, apex rounded or somewhat pointed. Thorax: Foretarsus black; forefemur bearing 4–5 flexor setae picta (Falle´n) along anteroventral surface. Abdomen: Gonites small, slender-shanked; base of aedeagus acute dorsally, Notiphila picta Falle´n 1813:254 [Sweden; ST (&, curving to posterior points; female sternite 4 small, NRS].—Sturtevant & Wheeler 1954:241 [revision, narrow, subequal in width to sternite 2; sternite 5 key]. rounded anteriorly, tapered to wider, truncate poste- (Hydrina) picta: Haliday 1839:405 [generic rior margin. combination]. Specimens examined.—MARYLAND. Montgom- picta: Loew 1860:25 [generic combina- NUMBER 15 183

tion].—Hollmann-Schirrmacher 1998:105–107 [re- sia (European Territory, Far East), Slovakia, Slove- vision]. nia, Spain, Sweden, Switzerland, Ukraine. Philygriola picta Hendel 1917:42 [generic combina- Remarks.—We have studied 241 specimens from tion]. North America and 101 specimens from Europe. Fa- Nostima (Philygriola) picta: Cresson 1930:101 [ge- cial and leg coloration is highly variable, even among neric combination]; 1944:176 [review].—Sturte- specimen series collected at the same date and locale. vant & Wheeler 1954:241 [review]. Genitalia from all the Nearctic and European speci- Nostima picta: Wirth 1965:745 [Nearctic catalog]. mens are virtually identical and separation into Ne- Nostima picta picta: Mathis & Zatwarnicki 1995:192 arctic and European species or subspecies with no [world catalog]. evidence of reproductive isolation presently seems Nostima scutellaris Cresson 1933:68 [United States. unwarranted. Nostima picta is apparently a morpho- Indiana. Laporte: Michigan City; HT (, USNM logically diverse Holarctic species. (44806)].—Edmiston & Mathis 2005:55 [synony- my]. Loew Nostima (Philygriola) scutellaris Cresson 1944:176 Philygria debilis Loew 1861:357 [United States. [review]. ‘‘Pennsylvania’’; ST (, MCZ (11160)].—Deonier Nostima scutellaris scutellaris: Sturtevant & Wheeler 1965:501 [biology].—Wirth 1965:745 [Nearctic 1954:241 [revision].—Wirth 1965:745 [Nearctic catalog].—Scheiring & Foote 1973:160 [habitat in catalog].—Mathis & Zatwarnicki 1995:193 [world Ohio].—Steinly 1984:753–754 [biology]; 1990: catalog]. 188–193 [biology].—Mathis & Zatwarnicki 1995: Nostima scutellaris occidentalis Sturtevant & Wheel- 195 [world catalog].—Hollmann-Schirrmacher er 1954:242 [United States. California. Riverside: 1998:66–71 [revision].—Edmiston & Mathis 2005: Pasadena; HT &, ANSP (6702)].—Mathis & Za- 12–15 [review]. twarnicki 1995:193 [world catalog].—Edmiston & Hydrina debilis: Slosson 1902:8 [generic combina- Mathis 2005:55 [synonymy]. tion].—Cresson 1944:177 [review].—Sturtevant & Wheeler 1954:237–238 [review]. Diagnosis.—Nostima picta specimens are distin- Philygria fuscicornis Loew 1862:155 [United States. guished from congeners by the following combina- ‘‘Middle States’’; ST &, MCZ (11159)].—Johnson tion of external characters: Head: Gena with dense 1925:273 [synonymy]. microtomentum ventrally. Thorax: Velvety black mi- Hydrina fuscicornis: Slosson 1902:8 [generic com- crotomentum on posterior margin of scutum and over bination]. entire scutellar disc; mesonotum striped laterally; Cressoniella montana Saether 1970:107 [United wing crossveins paler than veins, white, at least an- States. Colorado. Boulder: North Boulder Creek, terior portion. Femora and tibiae mostly black, apices northern branch; ST &, ZMO].—Mathis & Wirth sometimes paler. Abdomen: Tergite 5 of male lacking 1977:520 [synonymy]. silvery white microtomentum along posterior margin. Diagnosis.—This species is distinguished from Specimens examined.—MARYLAND. Montgom- Nearctic congeners by the following combination of ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW; at external characters: Small shore flies, body length light), 19 May–10 Oct 1914, R. C. Shannon (1(,1&; 1.30–1.86 mm; yellowish brown to brown with yel- ANSP, USNM). lowish-silver and gray microtomentum. Head: Para- Distribution.—Nearctic: Canada (Alberta, British facial area yellowish brown. Thorax: Wing mostly Columbia, Manitoba, New Brunswick, Ontario, Que- hyaline, infuscate spots over crossveins only; veins bec, Saskatchewan), United States (Alabama, Arizo- lacking spurious veins. Abdomen: Basal 1–4 tergites na, Arkansas, California, Colorado, District of Co- densely microtomentose, grayish brown, fifth tergite lumbia, Georgia, Idaho, Illinois, Indiana, Iowa, Kan- of male sparsely microtomentose to bare, subshiny to sas, Louisiana, Maryland, Michigan, Minnesota, Mis- shiny. sissippi, Missouri, New Hampshire, New Jersey, New Specimen examined.—MARYLAND. Montgom- York, North Carolina, Ohio, Oregon, Pennsylvania, ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 9 South Dakota, Tennessee, Texas, Virginia, Washing- May 1913, W. L. McAtee (1&; USNM). ton, Wisconsin, West Virginia). Neotropical: Guate- Distribution.—Nearctic: Canada (Alberta, British mala, Me´xico (Jalisco, Veracruz). Palearctic: Af- Columbia, Manitoba, Ontario, Quebec, Saskatche- ghanistan, Austria, Azores, Belgium, Bulgaria, Ca- wan), Mexico (Baja California), United States (Ala- nary Islands, Czech Republic, Egypt, Estonia, Fin- bama, Arizona, Arkansas, California, Connecticut, land, France, Germany, Great Britain, Hungary, Italy, Colorado, Delaware, District of Columbia, Idaho, Il- Japan (Hokkaido, Honshu), Macedonia, Madeira Is- linois, Indiana, Iowa, Kansas, Maine, Maryland, lands, Morocco, Netherlands, Poland, Romania, Rus- Massachusetts, Michigan, Minnesota, Mississippi, 184 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Missouri, Montana, Nebraska, Nevada, New Hamp- ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 5 shire, New Jersey, New Mexico, New York, North Aug 1913, R. C. Shannon (1(; USNM). Carolina, North Dakota, Ohio, Oklahoma, Oregon, Distribution.—Nearctic: Canada (Ontario, Que- Pennsylvania, South Carolina, South Dakota, Ten- bec), United States (California, Connecticut, District nessee, Texas, Utah, Vermont, Virginia, Washington, of Columbia, Illinois, Maryland, Massachusetts, West Virginia, Wisconsin, Wyoming). Michigan, New Hampshire, New Jersey, New York, Remarks.—Philygria debilis is one of the most Oregon, Pennsylvania, Rhode Island, Utah, Vermont, common and widespread Nearctic species, probably Virginia, Washington). Palearctic: Austria, Belgium, because of its occurrence on lawns associated with France, Germany, Great Britain, Netherlands, Poland, human habitations and recreation, such as golf cours- Russia (European Territory), Sweden. es (Cresson 1944, Zack 1998). Also, the overall body Remarks.—Although widespread over much of and wing length often exceeds 2 mm, allowing this temperate North America, this species is not collect- species to be easily collected with larger mesh nets. ed commonly and is presumably uncommon in na- Although common and widespread, the immature ture. We know nothing about its biology. stages of this species are unknown, and information on its natural history is limited to habitat descrip- argentata (Walker) tions. Presumably the larvae feed on blue-green al- gae. Notiphila argentata Walker 1853:407 [United States; This species is sexually dimorphic, and not rec- LT (sex?; designated by Mathis 1983:178), ognizing this dimorphism is probably the reason why BMNH]. Loew described this species twice. : Osten Sacken 1878:203 [generic combination].—Crawford 1914:37 [para- sitoid: Cyrtogaster glasgowi Crawford (ϭ C. try- Philygria punctatonervosa (Falle´n) pherus (Walker), Pteromalidae)].—Sturtevant & Notiphila punctatonervosa Falle´n 1813:254 [Sweden; Wheeler 1954:204–205 [review, distribution].— LT ( (designated by Mathis & Zatwarnicki 1990: Wirth 1964:5–6 [revision]; 1965:753 [Nearctic cat- 902), NRS]. alog].—Scheiring & Foote 1973:163 [natural his- Ephydra (Hydrina) punctatonervosa: Haliday 1839: tory].—Mathis 1983:179 [figs. of male termina- 405 [generic combination]. lia].—Mathis & Zatwarnicki 1995:230 [world cat- Philygria punctatonervosa: Loew 1860:25 [generic alog].—Mathis & Winkler 2003:410–411 [review]. combination].—Mathis & Zatwarnicki 1995:198– Brachydeutera dimidiata Loew 1862:163 [United & 199 [world catalog].—Hollmann-Schirrmacher States. District of Columbia: ‘‘Washington’’; LT 1998:110–112 [revision]. (designated by Mathis 1983:178), MCZ].—Osten Hydrina punctatonervosa: Becker 1926:57 [generic Sacken 1878:203 [synonymy]. combination]. Diagnosis.—This species is distinguished from Philygria opposita Loew 1861:356 [United States. congeners by the following combination of charac- ‘‘Pennsylvania’’; LT ( (designated by Mathis & ters: Head: Facial carina high, sharply defined, es- Zatwarnicki 1990: 902), MCZ (11161)].—Wirth pecially ventrally. Thorax: Brown color of anepister- 1965:745 [Nearctic catalog]; Mathis & Zatwarnicki num continued ventrally to about dorsal one-sixth to 1990:901 [synonymy]. one-third, thereafter sharply delimited from pale gray Hydrina opposite: Cresson 1944:177 [generic com- coloration on ventral pleural region. Abdomen: Ap- bination]. parent merger of fused surstyli with epandrium in lateral view broadly rounded; gonite digitiform api- Diagnosis. —This species is distinguished from cally, narrowly rounded. Nearctic congeners by the following combination of Specimens examined.—MARYLAND. Montgom- external characters: Small shore flies, body length ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 1.30–1.86 mm; yellowish brown to brown with yel- 28 Jun–17 Aug 1907, 1961, A. K. Fisher, P. J. Span- lowish-silver and gray microtomentum. Head: Para- gler (6&; USNM). facial area yellowish to silvery. Thorax: Wing con- Distribution.—Nearctic: Bermuda, Canada (Que- spicuously spotted (over crossveins r-m, dm-cu, and bec), United States (District of Columbia, Florida, spurious veins); veins R2ϩ3,R4ϩ5, and M bearing nu- Iowa, Michigan, Maine, Maryland, Massachusetts, merous spurious veins. Abdomen: Basal 1–3 tergites New Jersey, North Carolina, Ohio, Texas, Virginia). sparsely microtomentose, faintly grayish brown, mostly subshiny, becoming progressively shinier pos- (Parydra) aquila (Falle´n) teriorly; 4th and 5th tergites mostly to entirely bare, shiny. Ephydra aquila Falle´n 1813:247 [Sweden; LT & Specimens examined.—MARYLAND. Montgom- (designated by Clausen & Cook 1971:38), NRS]. NUMBER 15 185

Parydra aquila: Haliday in Walker 1856:345 [generic deagus boat-shaped, not fused to phallapodeme; gon- combination].—Clausen & Cook 1971:35–41 [re- ites triangular with rounded apices; subepandrial vision].—Krivosheina 1987:540–543 [biology, im- plate usually complete dorsally, narrow, with trian- mature stages].—Meyers & Deonier 1993:7–30 gular, medial fragment. [parasitoids: Kleidotoma parydrae Beardsley (Eu- Specimens examined.—MARYLAND. Montgom- coilidae), Anaphes species (Mymaridae)].—Mathis ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), & Zatwarnicki 1995:222–223 [world catalog]. 3–18 Aug 1912, 1913, J. R. Malloch, R. C. Shannon Napaea aquila: Becker 1926:99 [generic combina- (4(,2&; ANSP, USNM). tion]. Distribution.—Nearctic: Canada (Alberta, British Parydra bituberculata Loew 1862:165 [United Columbia, Manitoba, New Brunswick, Newfound- States. ‘‘Middle States’’; LT ( (desginated by land, Nova Scotia, Ontario, Quebec, Saskatchewan), Cresson 1949:246), MCZ (11168)].—Clausen & United States (Arizona, California, Colorado, Con- Cook 1971:35 [synonymy]. necticut, District of Columbia, Idaho, Illinois, Indi- Napaea (Parydra) bituberculata bituberculata: Stur- ana, Iowa, Kansas, Kentucky, Maryland, Massachu- tevant & Wheeler 1954:227 [generic combination]. setts, Maine, Michigan, Minnesota, Missouri, Mon- Parydra nitida Cresson 1915:70 [United States. Ida- tana, Nebraska, Nevada, New Hampshire, New Jer- ho. Latah: Potlatch; HT (, ANSP (6071)].—Clau- sey, New Mexico, New York, North Carolina, North sen & Cook 1971:36 [synonymy]. Dakota, Ohio, Oregon, Pennsylvania, South Dakota, Napaea (Parydra) bituberculata nitida: Sturtevant & Tennessee, Texas, Vermont, Virginia, Washington, Wheeler 1954:227 [generic combination, revised West Virginia, Wisconsin, Wyoming). Palearctic: status]. Austria, Belgium, Czech Republic, Denmark, Fin- Parydra tibialis Cresson 1916:150 [United States. land, France, Germany, Great Britain, Hungary, Italy, Arizona. Coconino: Oak Creek Canyon; HT (,KU Japan (Hokkaido, Honshu), Kazakhstan, Morocco, (694)].—Clausen & Cook 1971:40 [synonymy]. Netherlands, Norway, Poland, Romania, Russia (Eu- Napaea (Parydra) tibialis: Sturtevant & Wheeler ropean Territory, Far East, Siberia), Slovakia, Swe- 1954:229 [generic combination]. den, Switzerland, Ukraine, Yugoslavia. Parydra papulata Cresson 1949:247 [United States. Remarks.—This is a common and very widespread Washington. San Juan: Friday Harbor; HT (, species with a Holarctic distribution. ANSP (6631)].—Clausen & Cook 1971:39 [syn- onymy]. Parydra (Parydra) breviceps Loew Napaea (Parydra) bituberculata papulata: Sturtevant Parydra breviceps Loew 1862:167 [United States. & Wheeler 1954:228 [generic combination, revised ‘‘Middle States’’; LT & (designated by Cresson status]. 1949:244), MCZ (11174)].—Wirth 1965:750 [Ne- Diagnosis.—This species is distinguished from arctic catalog].—Clausen & Cook 1971:43–46 [re- congeners by the following combination of charac- vision].—Mathis & Zatwarnicki 1995:223 [world ters: Moderately large to very large species, body catalog]. length 3.70–5.50 mm; generally brown to blackish Napaea breviceps: Johnson 1925:274 [generic com- brown species. Head: Fronto-orbital setae short, bination]. length one-third or less the shortest diameter of eye; Parydra limpidipennis Loew 1878:201 [United ( face in lateral view conspicuously convex; facial mi- States. ‘‘District of Columbia’’; LT (designated crotomentum varying (best seen in anterior view) by Cresson 1949:244), MCZ (11170)].—Cresson from nearly uniformly densely microtomentose to 1949:243 [synonymy]. dorsal half more thinly microtomentose, becoming Parydra vicina Cresson 1940:9 [United States. Cal- ( densely microtomentose ventrally, microtomentum ifornia. Alameda: Berkeley Hills; HT , ANSP white to silvery white. Thorax: Apical scutellar pro- (6546)].—Sturtevant & Wheeler 1954:223 [syn- cess lacking; lateral setae present, short, inserted onymy]. Napaea (Napaea) breviceps vicina: Sturtevant & close to apical setae; apical scutellar setae and tub- Wheeler 1954:223 [generic combination, revised ercules present, setae short, distance between apical status]. scutellar tubercules greater than their length; katepis- ternal spinelike seta present. Wing with moderately Diagnosis.—This species is distinguished from pronounced pattern, infuscation especially evident congeners by the following combination of charac- over crossveins. Femora and tibiae generally black ters: Moderately small to large species, body length with some moderately sparse grayish to tannish mi- 2.5–3.6 mm; brown to blackish-brown species. Head: crotomentum; tarsi generally black but basal tarso- Fronto-orbital setae long, length subequal to one-half meres with some reddish or orange coloration, es- shortest diameter of eye; face in lateral view shallow- pecially ventrally. Abdomen: Male terminalia: Ae- ly convex; facial microtomentum golden or bright 186 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON coppery. Thorax: Scutellum lacking an apical pro- aedeagus boat shaped, not fused to phallapodeme; cess; lateral scutellar tubercules lacking; apical setae gonites short, but somewhat angular apices; subepan- present, extended from moderately well-developed drial plate incomplete dorsally, comprising 2, lateral, tubercules. Wing lacking pronounced pattern, gener- triangular fragments. ally hyaline, infuscation generally limited to cross- Specimens examined.—MARYLAND. Montgom- veins. Femora black; tibiae and tarsi brownish red to ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), orangish red. Abdomen: Male terminalia with aedea- 21 Jul–18 Aug 1907, 1912, J. R. Malloch, W. L. gus long, boat shaped, not fused to phallapodeme; McAtee (20(,15&; ANSP, USNM). gonites long, rounded apically; subepandrial plate Distribution.—Nearctic: Canada (Alberta, Mani- connected dorsally, as a large, triangular plate. toba, Ontario, Quebec, Saskatchewan), United States Specimens examined.—MARYLAND. Montgom- (Alabama, Colorado, Delaware, District of Columbia, ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), Florida, Illinois, Indiana, Iowa, Kansas, Louisiana, 14 May–31 Aug 1914, 1963, K. V. Krombein, R. C. Maine, Maryland, Massachusetts, Michigan, Minne- Shannon (2&; USNM). sota, Mississippi, Missouri, Montana, Nebraska, New Distribution.—Nearctic: Canada (Ontario, Que- Hampshire, New Jersey, New York, North Carolina, bec). United States (Arizona, Arkansas, California, North Dakota, Ohio, Oregon, Pennsylvania, Rhode Colorado, Connecticut, Delaware, District of Colum- Island, South Dakota, Tennessee, Texas, Virginia, bia, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Wisconsin). Neotropical: Mexico (Chiapas, Hidalgo, Kentucky, Louisiana, Maryland, Massachusetts, Mexico, Morelos, Tabasco, Veracruz). Michigan, Minnesota, Mississippi, Missouri, Nebras- Remarks.—This is the most common species of ka, Nevada, New Hampshire, New Jersey, New York, Parydra on Plummers Island and often can be abun- North Carolina, Ohio, Oklahoma, Oregon, Pennsyl- dant. Elsewhere in the Nearctic Region, this species vania, South Carolina, South Dakota, Tennessee, Tex- is widespread, although it does not occur in the West as, Utah, Virginia, West Virginia, Wisconsin). Neo- beyond the Rocky Mountains. The well-developed tropical: Mexico (Baja California). scutellar tubercles are especially characteristic of this species and distinguish this large species from con- Parydra (Parydra) quadrituberculata Loew geners. Parydra quadrituberculata Loew 1862:165 [United stenhammari (Zetterstedt) States. ‘‘Middle States’’; LT & (designated by Cresson 1949:250), MCZ (11172)].—Wirth 1965: Ephydra stenhammari Zetterstedt 1846:1842 [Swe- 749 [Nearctic catalog].—Clausen & Cook 1971: den. Ska˚ne, Mellby, Espero¨d; LT & (designated by 57–59 [revision].—Deonier & Regensburg 1978: Andersson 1971:55), ZIL]. 341–353 [biology].—Mathis & Zatwarnicki 1995: stenhammari: Loew 1860:40 [generic com- 227 [world catalog]. bination]. Napaea (Parydra) quadrituberculata: Sturtevant & Stictoscatella stenhammari: Collin 1930:139 [generic Wheeler 1954:228 [generic combination]. combination]. Eustigoptera stenhammari: Cresson 1930:128 [ge- Diagnosis.—This species is distinguished from neric combination]. congeners by the following combination of charac- Lamproscatella stenhammari: Hendel 1930:153 [ge- ters: Moderately large to large species, body length neric combination]. 3.6–4.6 mm; generally brown to blackish brown spe- Limnellia stenhammari: Cresson 1935:364 [generic cies. Head: Fronto-orbital setae short, length one- combination].—Wirth 1965:758 [Nearctic cata- third or less the shortest diameter of eye; face in lat- log].—Andersson 1971:55–56 [review].—Mathis eral view convex; facial microtomentum varying 1978:280–285 [revision].—Mathis & Zatwarnicki (best seen in anterior view) from nearly uniformly 1995:260–261 [world catalog]. densely microtomentose to dorsal half more thinly Ephydra oscitans Walker 1849:1106 [Canada. Ontar- microtomentose, becoming densely microtomentose io: Hudson Bay, Albany River, St. Martin’s Falls; ventrally, microtomentum white to silvery white. HT (sex?; abdomen missing), BMNH].—Mathis Thorax: Apical scutellar process lacking; lateral and 1978:280 [synonymy]. apical scutellar setae and tubercules present, setae Scatella oscitans: Osten Sacken 1878:204 [generic short; distance between apical scutellar tubercules combination]. less than their length; katepisternal spinelike seta pre- Ilythea oscitans: Coquillett 1900:260 [generic com- sent. Wing lacking pronounced pattern, generally hy- bination]. aline, infuscation generally limited to crossveins. Limnellia oscitans: Wirth 1965:758 [generic combi- Legs generally black with some grayish microtomen- nation]. tum and tibiae with silvery gray microtomentum ba- sally and apically. Abdomen: Male terminalia with Diagnosis.—This species is distinguished from NUMBER 15 187

congeners by the following combination of charac- Scatella (Scatella) picea (Walker) ters: Head: Face with setulose portion mostly uni- Ephydra picea colorous, brown to grayish brown, occasionally with Walker 1849:1105 [Canada. Ontario: Hudson Bay, Albany River, St. Martin’s Falls; ST faintly colored cinereous band along ventral margin; &, BMNH]. antennal groves shallowly impressed, but more so : Cole & Lovett 1921:335 [generic than in L. anna. Thorax: Wing maculation pattern as combination].—Wirth 1965:757 [Nearctic cata- follows: cell r with checker board pattern of 5–6 1 log].—Mathis & Zatwarnicki 1995:276 [world cat- dark and 5–6 whites; cell r ϩ with 3 dark spots along 4 5 alog]. anterior margin; basitarsus of legs concolorous with Scatella lugens Loew 1862:171 [United States. tibiae, black; halter brownish yellow to black. Ab- ‘‘Middle States’’; ST (&, MCZ (11178)].—Cres- domen: Structures of male terminalia as follows: son 1935:365 [synonymy]. epandrium in posterior view longer than wide, round- ed dorsally, truncate ventrally; surstylus large, gen- Diagnosis.—This species is distinguished from erally subrectangular, ventrolateral margin rounded, congeners by the following combination of charac- setulose, not fused indistinguishably with ventral ters: Head: Face mostly densely tan to brownish mi- margin of epandrium; aedeagus long than wide, with crotomentose; gena short, less than one-third eye a V-shaped notch dorsally in lateral view. height; genal seta well developed, conspicuous. Tho- Specimens examined.—MARYLAND. Montgom- rax: Bearing a well-developed pair of presutural ac- ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), rostichal setae; well-developed dorsocentral setae all 28 Oct 1913, R. C. Shannon (1(; USNM). postsutural; basal scutellar setae much shorter than Distribution.—Nearctic: Canada (British Colum- apical setae. Wing with conspicuous spots; coloration bia, Manitoba, Newfoundland, Northwest Territories, mostly faintly brownish; wing cell r2ϩ3 with a hyaline Ontario, Quebec, Saskatchewan), Greenland, United area toward apex; costal section I of male normally States (Alaska, Idaho, Indiana, Iowa, Massachusetts, developed, not extensively and conspicuously thick- Maryland, Maine, Minnesota, Montana, New Hamp- ened. Abdomen: Tergites moderately to thinly micro- shire, New Jersey, North Carolina, Ohio, Oregon, tomentose, partially subshiny. Pennsylvania, South Carolina, South Dakota, Virgin- Specimens examined.—MARYLAND. Montgom- ia, Washington). Palearctic: Algeria, Austria, Czech ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), 8 Republic, Estonia, Finland, France, Germany, ? Great Apr 1914, R. C. Shannon (1&; ANSP). Britain, Hungary, Japan (Hokkaido), Netherlands, Distribution.—Nearctic: Canada (Ontario, Que- Norway, Poland, Russia (European Territory, Far bec), United States (Alabama, Connecticut, Dela- East), Spain, Sweden. ware, District of Columbia, Florida, Iowa, Massachu- setts, Maine, New Hampshire, New Jersey, New York, Ohio, Oregon, Rhode Island, Virginia). Scatella (Scatella) obsoleta Loew

Scatella obsoleta Loew 1861:358 [United States. Discussion District of Columbia: ‘‘Washington’’; ST &, MCZ While the shore-fly fauna of Plummers Island is (11175)].—Sturtevant & Wheeler 1954:180 [re- relatively diverse, with 29 species in 17 genera, we view].—Wirth 1965:757 [Nearctic catalog].— expect several additional species and probably genera Olafsson 1991:28–29 [revision].—Mathis & Za- to be found on the Island. This faunal survey, for twarnicki 1995:274–275 [world catalog]. example, was based on only 174 specimens, and these were collected primarily in the first half of the Diagnosis.—This species is distinguished from 20th century. With the exception of two specimens congeners by the following combination of charac- that Amnon Freidberg collected in the 1980s, there ters: Head: Mesofrons bare to sparsely microtomen- are no specimens in the collections that we examined tose, subshiny to shiny, broadly reaching ptilinal su- that were collected in the 1970s, 1980s, 1990s, or ture; gena moderately high, about one-fourth eye thus far in this current decade. Moreover, no spe- height. Thorax: Dorsocentral setae 2 (0ϩ2); usually cialist on the family has collected on the Island. Thus, with a well-developed, presutural pair of acrostichal the number of species is reflective of a lack of sys- setae; wing mostly hyaline, spots very obscure to tematic collecting by specialists. Certainly the poten- nonexistent; costal section I of male greatly swollen; tial for discovery of more shore-fly species exists. In coloration brown dorsally but gray to whitish gray on the Delmarva states, for example, we recorded over pleuron. 140 species, and we anticipate that a significant num- Specimens examined.—MARYLAND. Montgom- ber of these eventually will be found on the Island. ery Co.: Plummers Island (38Њ58.2ЈN, 77Њ10.6ЈW), Across the river on the Virginia side of the Potomac 29 Jul 1963, G. C. Steyskal (1(; USNM). River (Turkey Run and Great Falls), we have recently 188 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON identified 62 shore-fly species in 30 genera from a Bunyard, B., & B. A. Foote. 1990. Acalyptrate Diptera reared from single season’s collecting (2006), and many if not higher fungi in northeastern Ohio.—Entomological News 101:117–121. most of these species will undoubtedly occur on Clausen, P. J. 1977. A revision of the Nearctic, Neotropical, and Plummers Island. The need for specialized collecting Palearctic species of the genus Ochthera including one Ethi- on the Island is evident. opian species, and one new species from India.—Transac- As noted in the introduction, there are no shore tions of the American Entomological Society 103:451–530. flies listed by the state of Maryland as rare or en- , & E. F. Cook. 1971. A revision of the Nearctic species of the Tribe Parydrini (Diptera: Ephydridae).—Memoirs of the dangered. While we are not advocating that any of American Entomological Society 27:1–150. the shore flies from Plummers Island be listed by Cole, F. R., & A. L. Lovett. 1921. An annotated list of the Diptera Maryland as rare or endangered, there are three shore (flies) of Oregon.—Proceedings of the California Academy flies from the Island that are represented by few spec- of Sciences series 4, 11(15):197–344. imens, and for two of these, Hydrellia decens and H. Collin, J. E. 1930. Some new species of the dipterous genus Sca- tella Dsv. and the differentiation of Stictoscatella gen. nov. insulata, the type locality is Plummers Island. The (Ephydridae).—Entomologist’s Monthly Magazine 66:133– first species, H. decens, is known from a few females 139. only with the possible exception of a single male Coquillett, D. W. 1900. Report on a collection of dipterous from central Virginia. Wasps, as part of provisioning from Puerto Rico.—Proceedings of the United States Na- their nests, collected the second species, H. insulata, tional Museum 22:249–270. . 1910. New genera and species of North American Dip- and all specimens we examined were recovered from tera.—Proceedings of the Entomological Society of Wash- wasp nests. The third species highlighted here is Phil- ington 12:124–131. ygria punctatonervosa. This is a Holarctic species Crawford, J. C. 1914. Three new Hymenoptera (Chalcidoidea).— that is found only rarely in North America, although Insecutor Inscitiae Menstruus 2(3):36–38. it is apparently widespread in the Nearctic Region Cresson, E. T., Jr. 1915. Descriptions of new genera and species of the dipterous family Ephydridae.–II.—Entomological (Mathis & Zatwarnicki 1995). One of the few sites News 26:68–72. in eastern North America where this species has been . 1916. Studies in the American Ephydridae (Diptera). I. collected is Plummers Island. Revision of the species of the genus Paralimna.—Trans- In summary, although diverse, we anticipate that actions of the American Entomological Society 42:101– more shore-fly species will be discovered on the Is- 124. land. Even among the 29 species that have been iden- . 1917a. Studies in the American Ephydridae (Diptera). II. A revision of the species of the genera Notiphila and Di- tified, there are species that are seldom collected and chaeta.—Transactions of the American Entomological So- are known primarily or only from this Island. ciety 43:27–66. . 1917b. Descriptions of new genera and species of the Acknowledgments dipterous family Ephydridae. IV.—Entomological News 28: 340–341. . 1918. Costa Rican Diptera collected by Philip P. Calvert, We gratefully acknowledge the assistance and co- Ph.D., 1909–1910. Paper 3. A report on the Ephydridae.— operation of many organizations and individuals who Transactions of the American Entomological Society 44: contributed to production of this paper. To Jon K. 39–68. Gelhaus and Jason D. Weintraub (Academy of Nat- . 1922. Studies in American Ephydridae (Diptera). III. A ural Sciences of Philadelphia, Pennsylvania) and their revision of the species of Gymnopa and allied genera con- institution, who gave access to their collection, we stituting the subfamily Gymnopinae.—Transactions of the American Entomological Society 47:325–343. express our sincere thanks. Hollis B. Williams pro- . 1929. Studies in the dipterous family Ephydridae. Paper vided technical support. We also thank John W. II.—Transactions of the American Entomological Society Brown, Allen L. Norrbom, David R. Smith, F. Chris- 55:165–195. tian Thompson, and Hollis B. Williams for reviewing . 1930. Descriptions of new genera and species of the dip- an earlier draft of this paper. terous family Ephydridae. Paper VIII.—Entomological News 41:76–81. . 1931. Descriptions of new genera and species of the dip- Literature Cited terous family Ephydridae. Paper IX.—Entomological News 42:104–108. Andersson, H. 1971. The European species of Limnellia (Dipt., . 1933. 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Meyers, D. M., & D. L. Deonier. 1993. A behavioral-ecological . 1853. Diptera. Pp. 253–414 in W. W. Saunders, ed., In- study of Kleidotoma parydrae Beardsley (Hymenoptera: secta Saundersiana: or Characters of Undescribed Insects in Eucoilidae), with notes on Anaphes sp. (Hymenoptera: My- the Collection of William Wilson Saunders, Esq., F. R. S., maridae) parasites of Parydra spp. (Diptera: Ephydridae).— F. L. S., &c. 1(4), Van Voorst, London. Contributions of the American Entomological Institute 27: . 1856. Catalogue of the dipterous insects collected at Sa- 7–30. rawak, Borneo, by Mr. A. R. Wallace, with descriptions of Olafsson, E. 1991. Taxonomic revision of western Palaearctic spe- new species.—Proceedings of the Linnean Society of Lon- cies of the genera Scatella R.-D. and Lamproscatella Hen- don Zoology 1:105–136. del, and studies on their phylogenetic positions within the Wiedemann, C. R. W. 1830. 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Erster Nachtrag zu dem in Jahre 1843 book, 276, U.S. Department of Agriculture, Washington, bekannt gemachten verzeichnisse in Wu¨rttemberg vorkom- D.C., 1696 pp. mender zweiflu¨gliger Insekten.—Correspondenzblatt der . 1970. A new Athyroglossa with a key to North American ko¨niglich Wu¨rttembergischen Landwirthschaftlichen Ver- species (Diptera: Ephydridae).—Journal of the Kansas En- eins (new series) 17(1):49–64. tomological Society 43:315–319. Runyan, J. T., & D. L. Deonier. 1979. A comparative study of Zack, R. S. 1998. Shore flies (Diptera: Ephydridae) of the Hanford Pseudohecamede and Allotrichoma. Pp. 123–137 in D. L. Site, Washington.—Northwest Science 72:127–141. Deonier, ed., First Symposium on the Systematics and Ecol- Zatwarnicki, T. 1988. Materials to the knowledge of the genus ogy of Ephydridae (Diptera), North American Benthological Hydrellia Robineau-Desvoidy (Ephydridae, Diptera).—Pol- Society, Oxford, Ohio, 147 pp ϩ iii. skie Pismo Entomologiczne 58:587–634. 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Additional list of insects taken in alpine re- gion of Mt. Washington.—Entomological News 13:319– Checklist of Shore Flies from Plummers Island 321. (5 subfamilies, 13 tribes, 17 genera, 29 species) Steinly, B. A. 1984. Shore fly (Diptera: Ephydridae) community structure in a xeric grass habitat.—Proceedings of the En- Subfamily Discomyzinae Acloque tomological Society of Washington 86:749–759. Tribe Discomyzini Acloque . 1990. Shore-fly (Diptera: Ephydridae) community struc- Leptopsilopa atrimana (Loew) ture in selected terrestrial habitats of Ohio and Illinois.— Subfamily Robineau-Desvoidy Proceedings of the Entomological Society of Washington Tribe Dryxini Zatwarnicki 92:188–193. Paralimna punctipennis (Wiedemann) , & J. T. Runyan. 1979. The life history of Leptopsilopa Tribe Hydrelliini Robineau-Desvoidy atrimana (Diptera: Ephydridae). Pp. 139–147 in D. L. 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Tribe Hecamedini Mathis Subfamily Ephydrinae Zetterstedt Allotrichoma simplex (Loew) Tribe Dagini Mathis Tribe Ochtherini Dahl Brachydeutera argentata (Walker) Ochthera anatolikos Clausen Tribe Parydrini Wirth and Stone Subfamily Cresson Parydra aquila (Falle´n) Tribe Ilytheini Cresson P. breviceps Loew Zeros flavipes (Williston) P. quadrituberculata Loew Tribe Philygriini Lizarralde de Grosso Tribe Scatellini Wirth and Stone Nostima picta (Falle´n) Limnellia stenhammari (Zetterstedt) Philygria debilis Loew Scatella obsoleta Loew P. punctatonervosa (Falle´n) S. picea (Walker)