2 January 2007 PROC. ENTOMOL. SOC. WASH. 10% I ). 2007. pp. 208-222

THE IDENTITY AND DISTRIBUTION OF PLENA (HINE) AND E. NEMORALIS (HINE) (DIPTERA: )

The Museum, Department of Entomology, University of Arkansas, Fayetteville. Arkansas 72701-1201, U.S.A. (e-mail: [email protected])

Ahsrrvr~~~.-Eflercrplencr (Hine) and E r?o~~oroli.s(Hine) from the eastern and south-central United States are redescribed and illustrated. These members OF the Effi.rin .stan?inecr group are similar in appearance, and specimens of E. plena often have been misidentified as E. rzmzoro1i.r. Several characters for reliable identification of males are offered. Thc distributions of the species are mappcd. A lectotype is designated from the syntype series of Eras r?~~r?~oruli.~Hine. Key Words: Diptera, robber , Asilidae, Efferirr. Erau, Pogonieffirirr, Pogorrio<:fi /?ria, Nearctic

Efleriu, with about 110 species recog- species groups, based largely on wing nized in the United Statcs (Poole 1996) characteristics. and more than 110 species in the more American species were assigned to southern reaches of the Americas (Mar- Era..: until Martin (1961) showed that tin and Papavero 1970), is one of the Macquart should have erected a new largest genera of Asilidae in the New for the American species. He World. Coquillett (1893) erected the recognized that Eflrriu is the first generic genus to receive a small group of North name available for the North and South American species previously referred to American species. Hull (1962), spparent- the old Palearctic genus Elvrr Scopoli, ly unaware of Martin's (1961) work, while leaving other North American stated that the New World species only species in that Old World genus. He superficially resemble the Old World subsequently (Coquillett 1910) designat- Er0.x species and proposed the new genus ed E carididn Coquillett as type species Nercr.~to receive the majority of Amer- of his new North American genus. ican species, while reserving Efjerio for Assignment of Western Hemisphere the species included in Hine's Err,.\- robber to Erax began with Mac- anoma1u.s group - the species in which quart (1838) and continued for a long the recurrent branch of vein R4 joins period. He assigned 29 species, most of Rz+, rather than ending blindly in cell them American, to the genus. Williston r~+~.Martin (1965) soon synomymized (1891) listed 74 species from South Neras under Effiria. In his revision of 4merica. In his Er0.r revision, Hine the North American species, Wilcox (1919) recognized 76 valid North Amer- (1966) followed Martin's (1961, 1965) ican species of Emx- sensu Macquart. interpretation of the genus Effhrir~and and he placed them in eight informal Hine's (1919) species group classifica- VOLUME 109, NUMBER I 209 tion. He described 30 new species and On closer examination, it was found that recognized a total of 100 species in North males of the two species differ in the America north of Mexico. color pattern of the third abdominal Artigas and Papavero (1997) divided tergite and in the form of the gonocoxite Efferia. They continued to recognize the and the apex of the epandrium. genera Diplosynapsis Enderlein and Por- Hine (1911) described Erax nernoralis nsil~~sCurran, removed Nernx from from several specimens, both male and synonymy, reserved Efferia for anom- female, collected at New Roads, Pointe ola-group species, and described the five CoupAe Parish, Lousiana, on July 15, new genera Albibarbefferia, Ariclefferia, 1905. Brolnley (1951) described Erax Cavinefferia, Pogoniefferia, and Tuhercu- monki from a male holotype collected lelferia to receive other species. (Artigas at Donna, Hidalgo County, Texas, on and Papavero (1997) used the spelling October 1, 1933. He distinguished it Pogoniefferia most frequently, but they from E. nemoralis by its smaller size, apparently misspelled it as Pogorzioef- more slender build, hyaline wings, white feria in one instance; the former spelling mystax and beard, darker legs, and more is here selected as correct.) These genera rounded male genitalia. Wilcox (1966) are nearly identical to Hine's (1919) did not study specimens of E. monki species groups, based on superficial before publishing his work on North differences in setation and wing venation American Efferia. He merely quoted that can vary among illdividuals of the Bromley's description, and did the best same species. Although the latest catalog he could in constructing the keys. Brom- of robber fly genera recognizes all of the ley (1951) indicated that the holotype Artigas and Papavero genera (Geller- was in the American Museum of Natural Grimm 2003), it is perhaps better to treat History, hut thorough searches of that them as synonyms of Efferia until in museum, the National Museum of Nat- depth phylogenetic analysis provides ural History, the California Academy of additional data upon which to base Sciences, the Texas A&M University a conclusion. Collection, and several other New species of Efferia are still being museums and collections have failed to recognized and described in western produce any type specimens of E. monki. North America (Bullington and Lavigne A syntype series of E. nemoralis and the 1984; Forbes 1987, 1988), but it is rare to holotype of E. plana were located in the find new species in the East. While Charles A. Triplehorn Insect Collection, surveying the robber flies of the south- Ohio State University. I now regard E. central United States, it became evident morzki as a species inquirerzda, of doubtful that trays of Efferia nemoralis (Hine) at identity and needing further investiga- the University of Arkansas Arthropod tion. The holotype and some other Museum and other collections actually specimens were collected in the Gulf contained two species. Differences in Coastal region late in the season, sug- coloration of the vestiture of the mystax gesting that it might be junior a synonym and legs initially suggested the presence of E. nemoralis. of two species. A specimen with a mystax Study of the genitalia of the E plena that is interpreted as being "yellow or holotype and E. nemoralis lectotype, yellowish" keys to E. nemoralis, and designated herein, has led me to the a specimen with a mystax that is inter- conclusion that specimens of E. plena preted as being white keys to E. nzonlci have often been misidentified as E. (Bromley) in Wilcox's (1966) seminal nemoralis and placed in trays with monograph on North American Efferia. properly identified E. nernoralis. For this 210 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON reason, E. plena was recorded from only veins R4 and Rs is located basad to the Kansas and Oklahoma by Martin and level of the basal section of vein Mz Wilcox (1965) in their catalog of North (apex of cell d), and the male abdomen American robber flies, and its wide- has several silvery pollinose segments spread occurrence in Arkansas was not with long, parted, white hairs. Within recognized (Scarbrongh 1972, Whitcomb the staminea group, they are segregated and Bell 1964). Hine (1916) described from many other species by the black Erax plenus from a male holotype femora and white or yellow mystax. Both collected in Douglas County, Kansas. Efferia plena and Efferia nemoralis occur No date was given. That specimen and widely in Arkansas, where the former several others studied by Hine have species is apparently most abundant in a light colored abdominal tergite 7. One June, and the latter species is most specimen from Onaga, Pottawatomie abundant in July and August. Efferia County, Kansas has a dark colored plena has a wide distribution from Mary- tergite 7, but Hine (1916) dismissed this land and North Carolina in the East to condition as intraspecific variation. Kansas, Oklahoma, and Texas in the However, Wilcox (1966) admitted to no Midwest. Efferh nemoralis is strictly variation, and he identified only speci- a south-central species, occurring also mens with a "silvery pollinose" segment in Mississippi, Louisiana, northeastern 7 as Efferia plena. Most specimens of E. Texas, and southeastern Oklahoma. The plena that I have seen have a dark- ranges of the two species overlap in colored tergite 7, and they are identified Arkansas and Mississippi. Several char- as E. nenzorulis when using Wilcox's acters for reliable identification of males (1966) key. are described. No reliable characters In his revision of the genus Erax, Hine have been found separating the females (1919) separated male E. plena from male of the two species. E. nentoralis by the color pattern of abdominal tergites 1-3. The holotype of E. plena has the first three abdominal segments dark, mostly with black hair Terminology largely follows that found above (Fig. 1) That species is thus in the Manual of Nearctic Diptera (McA1- distinguished from E nemoralis, which pine 1981, Wood 1981). Antennomere to Hine describes as having male abdominal scape ratios were formulatedby dividing segments 1-2 and a basal triangle of the length of each antennomere by the segment 3 dark and largely clothed with length of the first antennomere (the scape) black hair, but the apex of 3 is white and and then listing the numbers in order. covered with long white hair parted in Specimens from the following collec- the middle and directed outward tions were studied: Colorado State Uni- (Fig. 4A). Unfortunately, there is a great versity, Department of Bioagricultural deal of variation in the color pattern of Sciences and Pest Management, C. P. abdominal tergite 3 of E. plena speci- Gillette Museum of Arthropod Diversi- mens, with many showing a well de- ty, Ft. Collins (CSUC); Mississippi State veloped white posterior band and long, University, Mississippi Entomological white parted hairs (Fig. 2). Even the Museum, Mississippi State (MEMU); holotype has some posterolateral white National Museum of Natural History, areas and some long, white hairs (Fig. 1) Smithsonian Institution, Washington, D. Efferia plena and Efferia nemoralis are C. (USNM); Charles A. Triplehorn In- members of Hine's (1919) staminea sect Collection, Ohio State University, group of species. The junction of wing Columbus (OSUC); Oklahoma State VOLUME 109, NUMBER 1 211

Fig. I. E,!fi//i'l.Mipienit, holotypc ilhdomcn. cit~rvi~/

University, K. C. Emerson Museum, - Abdominal tergite j shining dark brown to Stillwater (OSEC); Kansas State Univer- black donally on anterior half, almost devoid of black setae, with bmad, postenor, sity, Department of Enton~ology,Man- silvery pollinose band V-notched at midline hattan (KSUC); Texas A & M Univer- (Fig. 4A); epandrium with deep apical sity Insect Collection, College Station notch, as in Fig. 4R; gonocoxite with poster- (TAMU); University of Arkansas Ar- oventral lobe narrow, more or less parallel- thropod Museum, Fayetteville (UAAM); sided; apex surpassing that of anterodorsal University of Kansas Natural History lobe: gonostylus long. surpassing gonocoxite by distance approximating length of apical Museum, Snow Entomological Collec- gonocoxite lobes (Fig. 4C); aedeagus sharply tions, Lawrence (SEMC); University of hooked and somewhat sigmoid basally Michigan Museum of Zoology, Insect (Fig. 4D) ...... E[fe/er.in nemorolb (Hine) Division, Ann Arbor (UMMZ); Univer- sity of Missouri. Department of Ento- Efferiu plerza (Hine) mology, Wilbur R. Enns Entomology (Figs. 1-3) Museum, Columbia (UMEE). Erax plenus Hine 1916: 21; Hine 1919: 148. Efferia plena: Wilcox 1966: 214. Pogonieffeeria plena: ~rtigasand Papa- vero 1997; 73. Both E. plena and E. nemoralis key to Male.-Body length: 19.4-28.6 mm couplet 9 in Wilcox's (1966) key to males of the stamineo group. They can be (mean ? S.D.: 23.0 2 1.8; N = 63). separated as follows: Head: Width 1.5 times eye height. Face golden tomentose, with large gih- 1. Abdominal tergite 3 mostly shining black bosity about as deep as length of scape or dark brown dorsally and bearing nlany extending nearly two-thirds of distance black setae (Fig. 2A); epandrium with shallow apical notch, as in Fig. 2R; gono- from oral margin to antenna1 bases. coxite with postel.oventra1 lobe broad Mystax consisting of long pale yellow basally, tapered to rounded apex; apex to deep golden yellow bristles covering not surpassing that of anterodorsal lobe; entire facial gibbosity, sometimes with 1- gonostylus short, not surpassing gonocox- 4 black bristles in ventolateral corners. ite by distance approximating length of apical gonocoxite lobes (Fig. 2C); aedeagus Gena brown, thinly golden pollinose. smoothly rounded basally (Fig. 2- Frons brown, subshining medially, gold- D) ...... Ererio plena (Hine) en pollinose laterally, with 3-5 long 212 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHTNGTON

black bristles and patch of many black, each divided by narrow line of pollen shorter, hairlike setae laterally. Vertex, along transverse suture, and pair of including ocellar tubercle, golden polli- thinly pollinose, central, brown vittae nose. Ocellar tubercle bearing 24long, extending from pronotum to region just strong, black bristles and several weaker posterior to transverse suture, pair sep- ones. Postcranium white tomentose. arated by narrow line of golden pollen Dorsal postocular setae black; lateral along mid-dorsal line of scutum. Postalar postocular setae pale yellow. Lower callus densely golden pollinose, covered occiput, postgena, and base of labium with short black setae and 3-6 long, with dense vestiture of long, slender, strong black bristles. Scutellum convex, golden plumose hairs. Proboscis black; brown. golden pollinose, with 6-12 palpus black, usually with black setae strong, long, erect, black marginal bristles dorsally and apically, golden setae ven- and many short black setae on disc. trally near base. Antenna, including Mediotergite brown, thinly golden polli- style, 2.0-2.5 mm long, brown; scape nose; anatergite gray pollinose; katater- setose dorsally, laterally, and ventrally, gite gray pollinose, with dense fan of long with setae usually black dorsally, often black and golden bristles. All mesopleural golden yellow laterally and ventrally; sclerites brown, golden pollinose. Anepis- pedicel setose apically; first flagellomere ternum with long, fine, golden to dark attenuate; antennomere to scape ratios brown, apically-crinkled hairs along an- 1.0-0.5-1.0-1.5. terior, dorsal and posterior margins; Thorax: Cervical sclerites brown, anterior and dorsal hairs erect; posterior sparsely golden pollinose, covered with hairs reclinate. Anterior basalare brown, long, pale, plumose hairs. Pronotum thinly pollinose, with long, fine, black, sparsely golden pollinose, covered with apically-crinkled hairs along anterior sparse pile of long, slender pale and margin; posterior brown, thickly golden black hairs; antepronotum with trans- pollinose. Katepisternum with long, kine, verse row of about 6-11 black bristles; black, apically-crinkled hairs posterodor- postpronotal lobe brown, densely golden sally. Anepimeron with few long, fine, pollinose, covered with short, black, black or pale, apically-crinkled hairs erect to reclinate setae. Propleuron posteriodaorsally. Katepimeron and brown. golden pollinose, with vestiture meron lacking setae or hairs. Basal of long, declinate, plumose white hairs swelling of pleural wing process golden on anterior portion of proepisternum; pollinose. Subalar sclerite brown. Mete- declinate golden hairs on posterior por- pisternum golden pollinose, covered with tion of proepisternum; and reclinate several long, fine, black or pale, apically- golden to brown hairs on proepimeron. crinkled hairs and dense crescentic patch Scutum brown in ground color, covered of short brown hairs posteroventrally. throughout with vestiture of short or Metepimeron gray pollinose, with very long, black, erect or slightly reclinate fine, apically-crinkled, erect golden hairs. setae; postsutural hairs and setae longer Legs: Coxae brown in ground color, than presutural hairs and setae; lateral golden pollinose; forecoxa densely cov- margin of scutum, in addition to several ered with long, golden, declinate bristles weaker black setae, with 3-4 strong, anteriorly; midcoxa with long, golden, black presutural setae and 1-3 strong, declinate bristles anteroventrally and black postsutural, supra-alar setae; scu- some stronger golden bristles laterally; tum mostly covered with dense golden hind coxa with fine, long, apically- pollen except for pair of broad, postero- crinkled, pale hairs anterolaterally and lateral, thinly pollinose or bare patches, posteriorly, and about 3 long black to VOLUME 109; NUMBER I 213

Fig. 2. e/foin plena, male. A, Tergites 3-5. left to right (Arkansas: Cr~igheadCo., near Joneshoro). B-D. Male penitalia (Arkansas: Benton Co.. Siloam Springs, Chesney Prairie Natural Area). B. Epandrinm, right latcral view. C, Gonocoxite and gonostylus, right lateral view. D, Aedeagus, right lateral view. Abbreviations: adl. anterodonal lobe of gonocoxite: gs, gonostylus; pvl, posteroventral lobe of gonocanite. pale bristles posterolaterally. Femora 4-5 stout, black bristles and 2-3 stout, dark brown to black, subshining. Fore- black bristles on basal half of anterior femur lacking stout bristles, with many face. Hind femur with many short golden long, black hairlike setae dorsally, ante- setae anterodorsally on basal three- riorly, and ventroapically, and with fourths; many long, dark, apically-crin- many long, golden, hairlike setae poster- kled, hairlike setae posteroventrally; ven- iorly and ventrally near base, these tral row of 7-9 stout black bristles; and ventral setae much longer than others. anterior row of 2-3 stout black bristles. Midfemur similar to forefemur, but with Tibiae reddish brown, darkened at api- anteroventral and posteroventral rows of ces. Foretibia with many long, golden, 214 PROCEEDINGS OF THE EN TOMOLOGICAL SOCIETY OF WASHINGTON apically-crinkled, hairlike setae poster- hairlike setae; posterior band narrow, oventrally; dense brush of short, fine, with setae parted at midline, falling over golden hairs anteroventrally; rows of sides of tergite. Tergite 4 (Fig. 2A) stout, black bristles anterodorsally and almost entirely silvery pollinose; devoid posterodorsally; and circle of several of black; covered with many long, white, stout, black bristles at apex. Midtibia apically-crinkled, hairlike setae parted at with many long, golden, apically-crin- midline and falling over sides of tergite. kled, hairlike setae ventrally; row of 4-6 Tergites 5 (Fig. 2A) and 6 almost entirely stout black bristles posterodorsally; and silvery pollinose; devoid of black setae circle of several stout, black bristles at and covered with many short, fine, white apex. Hind tibia with many long, golden, setae parted at midline and falling over apically-crinkled, hairlike setae ventrally; sides of tergite. Tergite 7 (Fig. 1) some- dense elongate patch of short golden times silvery pollinose, but usually large- setae posterodorsally; and stout black ly shining brown except for silvery to bristles as follows: 1 anterodorsally and 1 grayish brown pollinose anterior V- posterodorsally near base, 1-2 antero- shaped area and narrow lateral band; dorsally at about midlength, 1 anteriorly covered with many short, fine, recum- at about apical fourth, 2 anteroventrally bent, posteriorly directed, black setae. along apical third, and 5 circling apex Sternites 1-3 grayish brown pollinose, except posterodorsally. Tarsi uniformly except for narrow, silvery pollinose brown; length of first tarsomere about posterior marginal band on sternite 3. 1.GI .3 times length of second and third Sternites 47 silvery pollinose. Sternites combined. All claws black with brown 14 covered with many long, white, base. All pulvilli pale yellow. apically-crinkled, hairlike setae; sternites Wing: 11.&18.0 mm long (mean ? 5-7 with shorter, white, hairlike setae. S.D.: 14.5 t 1.4; N = 63). Very lightly Sternite 8 shining brown, with narrow infuscated, nearly hyaline; sometimes posterior band of long, slender, black more heavily infuscated in subcostal cell. bristles directed posteriorly. Halter with stem yellow, knob often Epandrium with shallow apical notch, darker. as in Fig. 2B. Gonocoxite with poster- Abdomen: Tergites dark brown to oventral lobe broad basally, tapered to black in ground color. Tergite 1 brown rounded apex; apex not surpassing that in ground color; covered with grayish of anterodorsal lobe. Gonostylus short, brown pollen and fine black setae not surpassing gonocoxite by distance throughout; with 5-8 stout black bristles approximating length of apical gonocox- and several long, pale, apically-crinkled ite lobes (Fig. 2C). Aedeagus smoothly hairlike setae posterolaterally. Tergite 2 rounded basally (Fig. 2D). mostly shining dark brown to black, with Type Material.-The holotype male is broad lateral and narrow posterior gray- in the Charles A. Triplehorn Insect ish brown pollinose band; covered dor- Collection at Ohio State University. Tt is sally with many long, black, hairlike labeled "Douglas Co., Kan. 900 ft., F. H. setae; with narrow lateral band of long, Snow.fI'YPE [red label]/Erax plenus Hine pale, hairlike setae. Tergite 3 (Fig. 2A) [handwritten, with black border]." The mostly shining dark brown to black, specimen is 27.4 mm long, and its wings usually with broad lateral grayish brown are 15.5 mm long. Tergite and sternite 7 pollinose band and narrow posterior are silvery pollinose. A paratype male in silvery pollinose band; covered mostly the same collection is labeled "Osborne with black setae; with posterior and Co, Kan. 1557 ft., F. X. Williams, Aug. 3, lateral bands of white, apically-crinkled, 19 12 [date handwritten1lPARATYPE VOLUME 109, NUMBER 1 215

Fig. 3. Distribution of efferio plenn by county

[red label]. It also has a silvery pollinose grass and forbs, 1.5 July 2004, N. Lavers tergite and sternite 7. (1 8,UAAM); Jonesboro, 5 June 1964, Other specimens examined.-AR- A. G. Scarbrough (1 6, UAAM), 28 KANSAS. Benton Co.: Chesney Prairie June 1964, A. G. Scarbrough (1 6, Natural Area, near Siloam Springs, 7-12 UAAM), 31 May 2004, N. Lavers (1 6, June 2004, J. K. Barnes (1 6, UAAM); UAAM); near Jonesboro, 16 June 1971, 12-18 June 2004, J. K. Barnes (3 6, A. G. Scarbrough (2 6, UAAM). Drew UAAM). Boone Co.: Baker Prairie Co.: Seven Devils Wildlife Management Natural Area, 2 July 2003, M. D. Area, understory trail, above swamp, 23 Warriner (1 6, UAAM). Craighead August 2004, H. Raney (1 6, UAAM). Co.: Bono Bog, 22 May 2003, N. Lavers Fanlkner Co.: Bell Slough Wildlife (1 8, UAAM); Hatchie Coon, on mud in Management Area, 26 May 2003, H. overflow area, 1 August 2004, N. Lavers Raney (1 8, UAAM); Camp Robinson (1 8, UAAM); Hatchie Coon, 1 August Wildlife Management Area, 1 June 2003, 2004, N. Lavers (2 8,UAAM); Hatchie H. Raney (1 6, UAAM). Franklin Co.: Coon Wildlife Management Area, tall Cherokee Prairie Natural Area, 19 May 216 PROCEEDINGS OF THE EN TOMOLOGlCAL SOCIETY OF WASHINGTON

2003, M. D. Warriner (1 6, UAAM). NORTH CAROLINA. Mecklenburg Lafayette Co.: Lafayette Wildlife Man- Co.: Charlotte, 5 June 1927 (5 8, agement Area, roadcut along fields, 15 USNM). August 2004, H. Raney (1 6, UAAM). OKLAHOMA. Comanche Co.: Fort Poinsett Co.: Lake Hogue, 27 June 2004, Sill, East Range, Parks Hill, 12 June N. Lavers (1 8, UAAM). Sharp Co.: 2003, J. Owens, J. Schmidt (I 8,CSUC); Strawberry River Preserve, in open field Fort Sill, East Range, nr Hoyle Bridge, near river, 22 May 2004, H. Raney (I 6, pinned with presumed prey: Colicrs phi- UAAM). lodice Godart, clouded sulphur butterfly, INDIANA. Clark Co.: State Forest, 12 June 2003, J. Owens, J. Schmidt (I 6, 13 June 1933, Montgomery (1 6, CSUC). Craig Co.: Bluejacket, 17 June UMMZ). 1931 (1 8,OSEC); Centralia. 26 June KANSAS. Clay Co. (1 6, OSUC). 1931 (1 8, OSEC). Jefferson Co.: Douglas Co.: 5 mi. NE of Lawrence, 18 Waurika Lake, 13 July 1983, H. C. Reed June 1971, V. P. Gapud (1 6, SEMC). & J. Nelson (2 8, OSEC). McCurtain Osage Co.: 12 June 1923, H. Darby (I 6, Co.: Idabel, 9 June 193 1, (1 8,OSEC). UAAM), 14 June 1923, H. Darby (1 6, Nowata Co.: Nowata, 28 June 1935, C. UAAM), 16 June 1923, R. H. Bealner A. Sooter (1 6, OSEC). Okmulgee Co.: (1 6,UAAM). Riley Co.: 5 June 1955, Bryant, 11 June 1934, C. A. Sooter (I 8, W. W. Gibson (1 6, KSUC). Pottawa- USNM), 13 June 1934, C. A. Sooter (2 6, tomie Co.: Onaga, 20 August 1901 (1 8, USNM); Henryetta, 7 June 1934, C. A. OSUC). Sooter (I 6, USNM), 15 June 1934, C. MARYLAND. Prince George's Co.: A. Sooter (1 6, USNM). Osage Co.: College Park, 9 June 1935, C. T. Greene Pawhuska, 3 June 1934, A. E. Pritchard (1 6.USNM). (1 6,USNM). Pushmataha Co.: Tuska- MISSISSIPPI. Oktibbeba Co.: 3 June homa, 23 May 1928, R. H. Beamer (I 6, 1989, T. L. Schiefer (3 6, MEMU); 3 mi. UAAM). W of Adaton, 20 June 1996, T. L. TENNESSEE. Coffee Co.: Manche- Schiefer (1 8,MEMU), 12 June 1998, ster, 20 June 1941, W. S. Craig (1 8, T. L. Schiefer (1 6, MEMU); 6 mi. SW UMEE). of Starkville, 6-7 July 1984, R. L. & B. TEXAS: Calhoun Co.: Port Lavaca, B. Brown (1 6, MEMU); Agricultural 12 August 1925 (1 8, OSUC), College, 16 May 1921, W. F. Turman (I Distribution (Fig. 3).-Nearctic: Unit- 6, MEMU); John Stars Memorial For- ed States (Arkansas, Indiana, Kansas, est, 14 June 1995, D. M. Pollock (1 6, Maryland, ~ississi'~pi,Missouri, North MEMU); Longview, 8 June 1919, F. Carolina, Oklahoma, Tennessee, Texas). Oswalt (1 6, MEMU); State College, 1 Existing collect~onrecords seem to in- May 1940, C. M. Wells (I 8,MEMU). dicate that E. plena is relatively rare east MISSOURI. Audrain Co.: Mexico, 1 of the Mississippi, but abundant in August 1993, T. Woods (1 6, UMEE). Kansas, Oklahoma. Arkansas, and Mis- Callaway Co.: Tucker Prairie, 19-20 souri. It is tempting to speculate that this June 1968 (2 8,UMEE), 2-6 July 1968 is a prairie species that spread eastward (5 8, UMEE), 8 July 1968 (2 6, with deforestation. UMEE)? 12 July 1968 (3 6, UMEE), Note.-The mystax and the long 14 July 1968 (1 8,UMEE), 18 July 1968 ventral hairs of the femora and tibiae (1 8, UMEE). Lawrence Co.: Mt. usually are golden yellow, although in Vernon, 16 June 1954, W. R. Enns (2 some specimens they appear to be paler. 8, UMEE). Pike Co.: New Hartford, 24 They are usually distinctly more richly June 1938, W. S. Craig (I 8, USNM). colored than those of E. nemoralis. VOLUME 109, NUMBER I 217

Fig. 4. Gff2~tirrn<.moililic, male. A, Tsrgites 3-5. left to right, lectotype (Louisiana: Point Coup& Parish, New Roads). B-D. Male genitalia (Arkansas: Hempstead Co. Rick Evans Graltdvicw Prairie). B. Epandrium, right lateral view. C, Gnnocoaite and gonostylus. right lateral view. D, Aedeagus. right lateral view. A specimen from Fort Sill, Comanche Male.-Body length: 20.5-27.9 mm Co., Oklahoma is pinned with its pre- (mean 2 S.D.: 24.2 -C 1.9; N = 49). sunled prey, an adult clouded sulphur Head: Width 1.5 times eye height. Face butterfly, Colius pl?ilodice Godart. white tornentose, with large gibbosity about as deep as length of scape extending Efferia nen?orrrlis (Hine) nearly two-thirds of distance from oral (Figs. 45) margin to antenna1 bases. Mystax con- El-CIXnemoralis Hine 191 1: 3 11; Hine sisting of long white or slightly yellowed 1919: 147. bristles covering entire facial gibbosity, Efferia nemoralis: Wilcox 1966: 208. sometimes with 1 or a few black bristles in Pogoniejyeria nenzoralis: Artigas and ventolateral corners. Gena brown, thinly Papavero 1997: 73. pale pollinose. Frons brown, subshining 218 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON medially, golden pollinose laterally, with dense golden pollen except for pair of 3-5 long black bristles and patch of many broad, posterolateral, thinly pollinose or black, shorter, hairlike setae laterally. bare patches, each divided by narrow line Vertex, including ocellar tubercle, golden of pollen along transverse suture, and pollinose. Ocellar tubercle bearing 24 pair of thinly pollinose, central, brown long, strong, black bristles and several vittae extending from pronotum to re- weaker ones. Postcranium white tomen- gion just posterior to transverse suture, tose along ocular margin, with thin pale the pair separated by narrow line of pollen elsewhere. Dorsal postocular setae golden pollen along mid-dorsal line of black; lateral postocular setae pale yellow scutum. Postalar callus densely golden or brown. Lower occiput, postgena, and pollinose, covered with short black setae base of labium with dense vestiture of and 24 long, strong black bristles. long, slender, white, plumose hairs. Pro- Scutellum convex, brown, golden polli- boscis black; palpus black, with black nose, with 6-10 strong, long, erect, black setae dorsally and apically, white setae marginal bristles and short black setae on ventrally near base. Antenna, including disc. Mediotergite brown, thinly polli- style, 2.2-2.6 mm long, brown; scape nose; anatergite gray pollinose; katater- setose dorsally, laterally, and ventrally, gite gray pollinose, with dense fan of long the setae usually black dorsally, often black and pale bristles. All mesopleural white laterally and ventrally; pedicel sclerites brown, at least partly golden or setose apically; first flagellomere attenu- gray pollinose. Anepisternum with long, ate; antennomere to scape ratios 1.0-0.5- fine, golden to dark brown, apically- 0.9-2.1. crinkled hairs along anterior, dorsal and Thorax: Cervical sclerites brown, gold- posterior margins; anterior and dorsal en pollinose, covered with long, white, hairs erect; posterior hairs reclinate. plumose hairs. Pronotum sparsely golden Anterior basalare brown, thinly polli- pollinose, covered with sparse pile of nose, with long, fine, black, apically- long, slender pale and black hairs; ante- crinkled hairs along anterior margin; pronotum with transverse row of about posterior basalare yellowish brown, sil- 11-20 black bristles; postpronotal lobe very pollinose. Katepisternum with long, brown, densely golden pollinose, covered fine, black, apically-crinkled hairs poster- with short, black, erect to reclinate setae. odorsally. Anepimeron with few long, Propleuron brown, golden pollinose; fine, black or pale, apically-crinkled with vestiture of long, declinate, plumose hairs posteriodorsally. '~ate~imeronand white hairs on anterior portion of proe- meron lacking setae or hairs. Basal pisternum; declinate golden hairs on swelling of pleural wing process silvery posterior portion of proepisternum and pollinose. Subalar sclerite brown. Mete- anterior portion of proepimeron; and pisternum golden pollinose, covered with reclinate black hairs on posterior portion several long, fine, black or pale, apically- of proepimeron. Scutum brown in crinkled hairs and dense crescentic patch ground color, covered throughout with of short brown hairs posteroventrally. vestiture of short or long, black, erect or Metepimeron gray pollinose, with very slightly reclinate setae; postsutural hairs fine, apically-crinkled, erect white to and setae longer than presutural hairs golden hairs. and setae; lateral margin of scutum, in Legs: Coxae brown in ground color, addition to several weaker black setae, gray to golden pollinose; forecoxa dense- with 24strong, black presutural setae ly covered with long, white, declinate and 1-3 strong, black postsutural, supra- bristles anteriorly; midcoxa with long, alar setae; scutum mostly covered with white, declinate bristles anteroventrally VOLUME 109, NUMBER 1 219 and longer, stronger, white to pale Wing: 13.7-18.8 mm long (mean C golden bristles laterally; hind coxa with S.D.: 15.8 t 1.0; N = 49). Lightly sparse, fine, long, apically-crinkled, infuscated; more heavily infuscated in white hairs anterolaterally and poster- subcostal cell. Halter with stem and knob iorly and about 3 long black to pale yellow. bristles posterolaterally. Femora dark Abdomen: Tergites dark brown to brown to black, subshining. Forefemur black in ground color. Tergite 1 brown lacking stout bristles, with many long, in ground color; covered with grayish black hairlike setae dorsally, anteriorly, brown pollen and fine black setae and ventroapically, and with many long, throughout; with 5-8 stout black bristles pale, hairlike setae posteriorly and ven- and several long, pale, apically-crinkled trally near base, these ventral setae much hairlike setae posterolaterally. Tergite 2 longer than others. Midfemur similar to mostly shining dark brown to black, with forefemur, but with anteroventral and broad lateral and narrow posterior gray- posteroventral rows of 65stout, black ish brown pollinose band; covered dor- bristles and 1-2 stout, black bristles on sally with many long, black, hairlike basal half of anterior face. Hind femur setae; with narrow lateral band of long, with many short yellow setae anterodor- pale, hairlike setae. Tergite 3 (Fig. 4A) sally on basal three-fourths; many long, shining dark brown to black dorsally on pale, apically-crinkled, hairlike setae anterior half; with broad silvery pollinose posteroventrally; ventral row of 5-9 band laterally and along posterior half, V- stout black bristles; and anterior row of notched at midline; almost devoid of 45stout black bristles. Tibiae yellowish black setae and bearing many long, white, brown, darkened at apices. Foretibia apically-crinkled, hairlike setae parted at with many long, pale, apically-crinkled, midline and falling over sides of tergite. hairlike setae posteroventrally; dense Tergite 4 (Fig. 4A) mostly silvery polli- brush of short, tine, golden hairs ante- nose except for anterior, mid-dorsal V- roventrally; rows of stout, black bristles notch; devoid of black setae and covered anterodorsalty and posterodorsally; and with many long, white, apically-crinkled, circle of several stout, black bristles at hairlike setae parted at midline and falling apex. Midtibia with many long, pale, over sides of tergite. Tergites 5 (Fig. 4A) apically-crinkled, hairlike setae ventrally; and 6 mostly silvery pollinose except for row of 4-6 stout black bristles poster- usually brown mid-dorsal line or V-notch; odorsally; and circle of several stout, devoid of black setae and covered with black bristles at apex. Hind tibia with many short, fine, white setae parted at marry long, pale, apically-crinkled, bair- midline and falling over sides of tergites. like setae ventrally; dense, elongate patch Tergite 7 largely shining brown except for of short golden setae posterodorsally; silvery pollinose anterodorsal V-shaped and stout black bristles as follows: 1 area and narrow lateral band; covered anterodorsally and 1 posterodorsally with many short, fine, recumbent, poster- near base, 1 anterodorsally at about iorly directed, black setae. Sternites 1 and midlength, 1 anteriorly at about apical 2 brownish gray pollinose, except for fourth, 2 anteroventrally along apical narrow, silvery pollinose posterior mar- third, and 5 circling apex except poster- ginal band on sternite 2. Sternites 3-7 odorsally. Tarsi uniformly brown; length silvery pollinose. Sternites 1-4 covered of first tarsomere about 1.3-1.4 times with many long, white, apically-crinkled, length of second and third combined. All hairlike setae; sternites 5-7 with shorter, claws black with brown base. All pulvilli white, hairlike setae. Stemite 8 shining pale yellow. brown, with narrow posterior band of 220 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Distribution of Efirio rzenrornlis by wunty and parish

long, slender, black bristles directed clarification and to ensure stabilization posteriorly. in nomenclature. It is labeled "New Epandrium with deep apical notch, as Roads, La, July 14-18'05/Jas. S Hine, in Fig. 4B. Gonocoxite with posteroven- Collector/COTYPE" [red label]lErax ne- tral lobe narrow, more or less parallel- moralis Hine [handwritten, with black sided; apex surpassing that of anterodor- borderlllectotype Male [red type], Dip- sal lobe. Gonostylus long, surpassing tera: Asilidae, Efferia nemoralis (Hine)/ gonocoxite by distance approximating designated by, J. K. Barnes [red bor- length of apical gonocoxite lobes der]." New Roads, Louisiana, is located (Fig. 4C). Aedeagus sharply hooked in Point Coupee Parish. The lectotype and somewhat sigmoid basally (Fig. 4D). abdomen is mounted on a paper point, Type material.-None of the female and the dissected genitalia are in a micro- syntypes were found. One specimen from vial, both attached to the same pin that a syntype series of four males located at holds the main body of the lectotype. the Ohio State University Insect Collec- Paralectotypes, 3 males, United States: tion is hereby designated lectotype for "New Roads, La, July 1418'05/Jas. S VOLUME 109, NUMBER 1

Hine, CollectorICOTYPE [red label]/ Madison Co.: 21 June 1931, Bibby & Paralectotype Male [red type], Diptera: Tate (1 8, TAMU); S. Bromley (1 8, Asilidae, Efferria nemoralis (Hine)ldet. J. USNM). McLennan Co.: Waco, 22 June K. Barnes [red border]." The lectotype 1948, P. A. Glick (1 6, USNM). Rusk and paralectotypes range in length from Co.: Tatum, 8 June 1949, D. J. & J. N. 24.4 to 25.0 mm; their wings range from Knull (1 8, OSUC). Co. unknown: 15.1 to 16.5 mm long. The lectotype Liberty, 10 June 1934 (8 8, TAMU). wing is 16.5 mm long. Distribution (Fig. 5).-Nearctic: Unit- Other specimens examined.-AR- ed States (Arkansas, Louisiana, Missis- KANSAS. Conway Co.: Pointe Remove, sippi, Oklahoma, Texas). 27 July 2003, H. Raney (1 8, UAAM). Note.-The mystax and the long Craighead Co.: Hatchie Coon, 18 July ventral hairs of the femora and tibiae 2003, N. Lavers (1 6, UAAM), 24 are usually straw yellow, although in August 2003, N. Lavers (1 8, UAAM). some specimens they appear to be sordid Dallas Co.: sweepings, 15 August 1960, P. white. They are usually distinctly paler Deema (1 8, UAAM). Faulkner Co.: than in specimens of E. plena. Camp Robinson, 21 July 2003, B. Bald- win (1 8, UAAM); Camp Robinson Wildlife Development Area, 26 Jnly I am grateful to Norm Lavers of 2003, (1 8,UAAM). Hempstead Co.: Joneboro, Arkansas, for drawing my cotton, 1 July 1959, (1 8, UAAM); attention to the problem that is the sweepings, 10 August 1960, P. Deema subject of this paper and to Eric Fisher, (1 8, UAAM); Rick Evans Grandview California Department of Food and Prairie, 400 ft., 25 May-2 June 2004, J. K. Agriculture, Sacramento, for discussions Barnes (1 6, UAAM), 2-14 June 2004, J. concerning the genus Efferia. Eric Fisher, K. Barnes (I 8, UAAM), 18 June-2 July Torsten Dikow (Cornell University and 2004, J. K. Barnes (3 8, UAAM), 26 American Museum of Natural History), August-7 September 2004, J. K. Barnes (1 and an anonymous reviewer provided 8, UAAM). Little River Co.: weeds, 4 thoughtful suggestions that materially July 1961, (2 6,UAAM). Perry Co.: improved the manuscript. The following Harris Brake Wildlife Management Area, individuals donated specimens used in 19 Jnly 2003, H. Raney (I 8, UAAM). this study: Norm Lavers, Herschel Raney LOUISIANA. Bossier Parish: Barks- (Conway, Arkansas), and Aubrey Scar- dale Air Force Base, 3 June 1998, D. M. brough (Towson University, Maryland). Pollock (3 8, MEMU). St. Tammany The following individuals provided loans Parish: Madisonville, 11 June 1951, P. 8, of specimens: Boris Kondratieff (CSUC); Beamers (1 SEMC). Webster Parish: Terence Lee Schiefer (MEMU); F. Chris- Lake Bistineau State Park, 22 May 1996, tian Thompson (USNM); Luciana Mu- M. MacGown (1 8, MEMU). setti and Creighton Freeman (OSUC); MISSISSIPPI. Oktibbeha Co.: Stark- Don. C. Arnold (OSEC); Gregory Zol- ville, 22 June 1982, R. L. Brown (1 8, nerowich (KSUC); Edward G. Riley MEMU); State College, July 1948, Sim- (TAMU); Zachary H. Falin (SEMC); mons (I 8, MEMU). Mark F. O'Brien (UMMZ); and Robert OKLAHOMA. Choctaw Co.: Hugo, W. Sites (UMEE). 20 June 1934, A. E. Pritchard (1 8, USNM). TEXAS. Hunt Co.: Greenville, 10 Artigas, J. N. and N. Papavero. 1997. The June 1935, (2 8, TAMU), 13 June American genera of Asilidae (Diptera): keys 1935, (6 8, TAMU; 2 8, USNM). for identification with an atlas of female 222 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

spermathecae and other morphological details. 1965. Distribution patterns and corrected IX.2. Subfamily Leach - Effeeria- identifications of asilid species reported as group, with the proposal of five new genera common to North and South America (Dip- and a catalogue of the Neotropical species. tera: Asilidae). Transactions of the American Arquivos de Zoologia 34(3): 65-95. Entomological Sociely 91: 1-37. Bromley, S. W. 1951. Asilid notes (Diptera), with Martin, C. H, and N. Papavero. 1970. Family descriptions of thirty-two new species. Amer- Asilidae. A catalogue of the Diptera of the ican Museum Novitates 1532. 36 pp. Americas south of the United States, fascicle Bullington, S. W. and R. J. Lavigne. 1984. 35b. Museu de Zoologia, Universidada de Sio Description and habitat of Efferia kondmlicffi Paulo, Brazil. 139 pp. sp. nnv, with notes on Effefferiuaesruuns (L.) Martin, C. H. and J. Wilcox. 1965. Family (Diptera: Asilidae). Annals of the Entomolog- Asilidae, pp. 360-401. In Stone, A., C. W. ical Society of America 77(4): 40013. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coquillett, D. W. 1893. A new asilid genus related Coulson, eds. A Catalog of the Diptera of to Emx. Canadian Entomologist 25: 175-177. Amercia North of Mexico. United States Department of Agriculture, Agriculture Re- . 1910. The tvoe-species.. . of the North American genera of Diptera. Proceedings of search Service, Agriculture Handbook 276. the United States National Museum 37: 1696 pp. 499-647. McAlpine. J. F. 1981. Morphology and terminol- Forbes, G. S. 1987. The status of Effcriu sirnilis ogy -Adults, pp. 943. In McAlpine, J. F., et (Williston), with descriptions of three new al., eds. Manual of Nearctic Diptera, "01- ume I. Research Branch Agriculture Canada Nearctic Effeer-ia species in the all~il~nrhirgroup Monograph 27. 674 pp. (Diptera: Asilidae). Pan-Pacific Enton~ologist Poole, R. W. 1996. Diptera, pp. 15-604. In Poole, 63(3): 292-300. R. W. and P. Gentili, eds. Nominil lnsecta -. -. 1988. Three new species of Eferia from Nearctica: A Check List of the of southern California and Arizona (Diptera: North America, volume 3. Diotera. Leoidov- Asilidae). Annals of the Entomological Society . . tera, Siphonaptera. Entomological Informa- of America 81(4): 554-559. tion Services, Rockville. Maryland. 1143 pv. Geller-Grimm, F. 2003. A world catalogue of thz Scarbrough, A. G. 1972. ~ecordsof robber-flies genera of the family Asilidae (Diptera). Studia from northeastern Arkansas. Proceedings of di~terolopica10(2): 473-526. the Entomological Society of ~ashington Hine, J. S. 1911. New species of Diptera of the 74(4): 375-378. genus Erox. Ohio Naturalist 1 l(6): 307-31 1. Wilcox, J. 1966. Ef/erin Coquillett in America -. 1916. Descriptions of robber flies of the north of Mexico (Diptera: Asilidae). Proceed- eenus Erax. Ohio Journal of Science 17(1):. . inas of the California Acndemv of Sciences 21-22. 342): 85-234. . 1919. Robberflies of the genus Erox. Williston. S. W. 1891. Cataloaue of the described Annals of the Entomological Society of Amer- species of South American Asilidae. Transac- ica 12: 103-154. tions of the American Entomological Society Hull, F. M. 1962. Robber flies of the world: The 18: 67-91. genera of the family Asilidae. Part 2. United Whitcomb, W. H. and K. Bell. 1964. Predaceous States National Museum Bulletin 224(2): insects. , and mites of Arkansas cotton 433-907. fields. University of Arkansas Agricultural Macquart, J. 1838. Dipthres exotiques nouveaux ou Experiment Station Bulletin 690, 84 pp. peu connus, volume 1, part 2. Paris. 207 pp. Wood, G. C. 1981. Asilidae, pp. 549-573. In Martin, C. H. 1961. The misidentification of Ernx McAlpine, J. F.. et al., eds. Manual of Nearctic Scopoli in the Americas. Journal of the Kansas Diptera, volume 1. Research Branch Agricul- Entomological Society 34(1): 14. ture Canada Monograph 27. 674 pp.