VOL. 106 JANUARY 2004 NO. 1 (ISSN 0013-8797) tcl PROCEEDINGS 'F NT of the ENTOMOLOGICAL SOCIETY of WASHINGTON ETT TSO PEBLISHED 4 0 = QUARTERLY CONTENTS ARCE-PEREZ, ROBERTO—Psephenopalpus browni, a new genus and species of Psepheninae (Coleoptera: Byrrhoidea: Psephenidae) tromeMexico) i961 ce nete sae ek ce eee ee eee 90 BUFFINGTON, MATTHEW L.—Taxonomic notes on Nordlandiella Diaz and Ganaspidium Weldi(ivmenoptera: Pisitidae; Bucolimae)y) vaccine. suet ween ce ctae ort ss areata eee ae ate te eee 192 CHAMORRO-LACAYO, MARIA LOURDES and RALPH W. HOLZENTHAL—Seven new species of Polyplectropus Ulmer (Trichoptera: Polycentropodidae) from Costa Rica ........ 202 CHEN, XUEXIN, J. B. WHITFIELD, and JUNHUA HE—Revision of the subfamily Cardiochilinae (Hymenoptera: Braconidae) in China. I. The genera Austerocardiochiles Dangerfield, Austin, and Whitfield, Eurycardiochiles Dangerfield, Austin, and Whitfield and ESTUOTUNISCUS EMGELI CIN iva 2ini Seis oe de ee Ree en eae oa Stee Pe OA eee ee 35 COLEMAN, B. K., J. K. BISSELL, J. CIHA, P. MacKEIGAN, AND J. B. KEIPER—The moths (Lepidoptera) and associated flora of Kelleys Island, Lake Erie ......................00e0 eee 217 FAVRET, COLIN, JOHN F. TOOKER, and LAWRENCE M. HANKS—/Jowana frisoni Hottes (Hemiptera: Aphididae) redescribed, with notes on its biology .....................4-. eM A 26 FOOTE, B. A.—Acalyptrate Diptera associated with stands of Carex lacustris and C. stricta (Gyscracede) immorineaStenm OIG Aes cnr ace Soleus victs shete wie laanie eae bioes claicin kale de lake ah oem ete 166 FLORES-MENDOZA, CARMEN, E. L. PEYTON, RICHARD C. WILKERSON, and RICAR- DO LOURENCO de OLIVEIRA—Anopheles (Nyssorhynchus) konderi Galvao and Damasceno: Neotype designation and resurrection from synonymy with Anopheles (Nyssorhynchus) oswaldoi (Peryassu) (Diptera: Culicidae) ................2. cece cece eee e nese 118 GAGNE, RAYMOND J. and CELIA dR. MEDINA—A new species of Procontarinia (Diptera: Cecidomyiidae), an important new pest of mango in the Philippines ................--.....25. 19 GRANARA de WILLINK, MARIA CRISTINA and DOUGLASS R. MILLER—Two new species of mealybugs (Hemiptera: Coccoidea: Pseudococcidae) from Patagonia, Argentina 140 HEIDEMAA, MIKK and ALEXEY ZINOVJEV—Dolerus anatolii, n. sp., the first Palearctic member of the subgenus Neodolerus Goulet (Hymenoptera: Tenthredinidae) ................ 159 HOLZENTHAL, RALPH W.—Three new species of Chilean caddisflies (Insecta: Trichoptera) .. 110 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2004 E. E. GrissELL, President Jon A. Lewis, Custodian JASON P. W. HALL, President-Elect MICHAEL G. PoGuE, Treasurer Stuart H. McKamey, Recording Secretary | JOHN W. BROWN AND Davin G. Furtu, Program Chairs Ho.tis B. WittiAMs, Membership Secretary JONATHAN R. Mawps ey, Past President Davip R. Smitu, Editor Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY Wayne N. Maruis Honorary President DoNnALD M. ANDERSON Honorary Members WILLIAM E. BICKLEY Kar V. KROMBEIN RONALD W. HopGeEs PAUL J. SPANGLER Manya B. STOETZEL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168. MEETINGS.—Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu- tion, on the first Thursday of each month from October to June, inclusive, at 7:00 P.M. Minutes of meetings are published regularly in the Proceedings. 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PLEASE SEE PP. 247-248 OF THIS ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, “% Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 7 January 2004 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 1-18 A NEW SPECIES OF MOMPHA HUBNER (LEPIDOPTERA: COLEOPHORIDAE: MOMPHINAE) FROM BUTTONBUSH (CEPHALANTHUS OCCIDENTALIS LL.) WITH DESCRIPTIONS OF THE EARLY STAGES DAviD L. WAGNER, DAVID ADAMSKI, AND RICHARD L. BROWN (DLW) Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT 06269, U.S.A. (e-mail: dwagner@uconnvm.uconn.edu); (DA) Department of Systematic Biology, Entomology Section, National Museum of Natural History, Smith- sonian Institution, Washington, DC 20560-0168, U.S.A. (e-mail: dadamski@sel.barc. usda.gov); (RLB) Mississippi Entomological Museum, Box 9775, Mississippi State, MS BOT 02a Abstract.—Mompha solomoni, n. sp., is described from moths reared from buttonbush, Cephalanthus occidentalis L. (Rubiaceae). Spring generations bore in young terminals of this plant species; summer and fall generations mine leaves. The egg, larval, pupal, and adult stages of the new species are described and illustrated. Mompha solomoni appears to be related to other Rubiaceae-feeding momphines, including the previously described and widely distributed Mompha cephalonthiella (Chambers), which also feeds on button- bush. Life history data are provided for both of these Cephalanthus-feeding momphines. Parasites of Mompha solomoni include members of the genera Pholetesor Mason (Bra- conidae), Bracon FE (Braconidae), and Euderus Haliday (Eulophidae). Key Words: In North America the Momphinae con- tain 37 described species and at least as many unrecognized species (Hodges et al. 1983, Hodges 1992, Poole and Gentili 1996, Koster 2002). While all momphines in North America are thought to share the habit of being internal feeders, larvae show a remarkably diverse range of feeding nich- es—tunneling in stems, forming galls, min- ing leaves, and boring through flowers and fruits. Larvae of the species described here feed in young shoots, killing the terminal portion in spring and early summer broods, but in subsequent broods the larvae mine leaves. Because Mompha solomoni appears to be related to a second buttonbush-leaf- miner, Mompha cephalonthiella (Cham- bers), we include considerable information on this species as well. leafmining, stem-mining, flagging, Rubiaceae Preparations of all life stages were ex- amined with dissecting, compound, and scanning electron microscopes. The Methu- en Handbook of Colour (Kornerup and Wanscher 1978) was used as a color stan- dard for the description of the adult. Geni- talia were dissected as described by Clarke (1941), except that mercurochrome and chlorazol black were used as stains. All measurements were made with a calibrated ocular micrometer. Ultrastructural studies were performed with a Hitachi HH-S-2R scanning electron microscope at an accelerating voltage of 20kV. For SEM examination, immature specimens were fixed in 3% glutaraldehyde in 0.1M potassium phosphate buffer (pH 7.3), rinsed in phosphate buffer (pH 7.3), i) and postfixed in 2% osmium tetroxide in 0.1M potassium phosphate (pH 7.3). After dehydration in ethyl alcohol, specimens were critical point dried, mounted on stubs with silver paint and paste, and coated with gold-palladium. Life history data for both Mompha spe- cies were obtained from laboratory rearings at ambient temperatures. James D. Solomon studied a population of the new Mompha species in the Delta Experimental Forest in Stoneville, Washington Co., MS. Casual observations were made on the population beginning in 1980. In 1986, collections of 10—20 shoots were made once or twice weekly in April, May, and June. Half of these were dissected to monitor larval de- velopment; larvae in the other half of the shoots were held in plastic Petri dishes over filter paper in ventilated jars to study pu- pation habits, parasitoids, and adult emer- gence. Additional collections were made by James Solomon in 1987 and 1988, mostly in the Delta National Forest, Issaquena and Sharkey cos., MS, but a few additional samples were secured from Chicot Co., AR, and East Carroll Parish, LA. Leafmines of Mompha_ cephalonthiella were taken at Nicholas along the Feather River, Sutter Co., CA, in September 1981 (DLW Lot: 81J6) and October 1982 (DLW Lot: 82K5); from along the American Riv- er, 4.8 km E of Auburn, El Dorado Co., CA, in September 1982 “(DLW Lot: 82312); Mansfield, Tolland Co., CT (DLW Lot: 89H49) in August 1989; and Sand Bar State Park, (Chittenden Co. V1, ine June 1957 (DLW Lot: 87F59). Collections of Mompha solomoni were made from Tolland and Windham cos., CT, in August, September, and October 1989 (DLW Lots: 89H49, 89J6, 89J14, 89K37, and 89K62); Monroe Co; Fie aim. March s199% “CDIEW or 91C14); Anne Arundel Co., MD, August 2002 (DLW Lot: 2002H28); and from Oak Point, Hammond, Saint Lawrence Co., NY, in August 1988 (DLW Lot: 88H45). In each case, leaves with actively feeding larvae were placed in a plastic bag with lightly PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON moistened paper toweling and were moni- tored daily for adult emergences. Rearing lots were held outdoors during the winter months. Larvae and pupae for subsequent study were killed in boiling water and trans- ferred to 70% ethyl alcohol. Specimen repositories are as follows: BL = private collection of Bernard Landry, Ge- neve, Switzerland; CNC = Canadian Na- tional Collection, Ottawa; ECK = private collection of Edward C. Knudson, Houston, TX; GB = private collection of George Balogh, Portage, MI; INHS = Illinois Nat- ural History Survey, Champaign; JRW = private collection of James R. Wiker, Ath- ens, IL; MEM = Mississippi Entomological Museum, Mississippi State; SK = private collection of Sjaak Koster, Leiden, Nether- lands; UCONN = University of Connecti- cut Insect Collection, Storrs; USNM = Na- tional Museum of Natural History, Smith- sonian Institution, Washington DC. In the material examined a “‘u”’ indicates a spec- imen could not be reliably sexed. Mompha solomoni Wagner, Adamski, and Brown, new species (Figs. 1, 3, 4, 7, 9-18, 20—40, 43—46, 49) Diagnosis.—In the discal area of the forewing, M. solomoni (Fig. 1) possesses an elongate black dash that is usually joined with a white dash along its anterior margin; neither marking is well developed in M. ce- pPhalonthiella (Fig. 2). The forewing of M. cephalonthiella is usually suffused with ap- preciably more coffee-brown and _ steely blue scales than that of M. solomoni. In M. solomoni the dark spot along forewing cos- ta at % is often strongly oblique and di- rected toward the outer edge of the wing or tornus; in M. cephalonthiella this mark is often less oblique and directed toward the tornus or inner margin. Between the dark costal spots at % and %4, M. cephalonthiella has a patch of tan scales; this area is mostly gray or brown in M. solomoni. In M. ce- phalonthiella the 1 or 2 white flagellomeres that cap the antenna are preceded by a se- ries of five dark flagellomeres; in M. solo- VOLUME 106, NUMBER 1 ly Figs. 1-2. moni an additional pale ring interrupts the preapical run of dark flagellomeres. Genitalia of the two species (Figs. 3—8) are very different in both sexes. In male M. solomoni (Fig. 3) the anellus has distolater- al, curved arms (that are absent in M. ce- phalonthiella) (Fig. 5); the ventral margin of the valva is entire (notched in M. ce- Phalonthiella); and the sacculus is narrow and acuminate (wide and erose in M. ce- phalonthiella). In female M. solomoni (Fig. 7), the invaginated pockets at the postero- lateral corners of tergum VII are much less prominent than in M. cephalonthiella (Fig. 8); also, in M. solomoni, the posterior pro- jections of the lamella postvaginalis are subtriangular, separated by a deep, V- shaped cleft, and preceded by a short, spi- nose thickening located medially at the an- terior end of the cleft (projections rounded, separated by a shallow cleft, and preceded by a long, sclerotinized thickening at the anterior end of cleft in M. cephalonthiella); furthermore, both of the sclerotized plates associated with the signa are small and crescent shaped in M. solomoni (one or both of the plates are large and round and completely surround the signum in M. ce- Phalonthiella); and finally, a lamella antev- aginalis is present in M. solomoni, but ab- sent in M. cephalonthiella. The mandibular setae of the larvae differ: in M. solomoni the setae are of unequal length (Fig. 18), whereas those of M. cephalonthiella are of similar length (Fig. 19). Description.—Head: Base of proboscis Adults: 1, Mompha solomoni. 2, M. cephalonthiella. and frons shiny white, vertex yellowish white, becoming gray to brownish gray posteriorly on occiput, a line of sparse brown scales extend from middle of eye an- teriorly to base of antenna, brown scales be- coming dense and covering dorsal surface of scape; distal portion of antenna with six white rings of one flagellomere each, each white ring separated by three brown flagel- lomeres, except terminal two rings separat- ed by two brown flagellomeres; labial pal- pus with first segment and medial surface of second segment white, lateral surface of second segment brown basally, intermixed with variable number of white scales api- cally, third segment brown, intermixed with white scales, with indistinct white bands at middle and apex. Thorax: Mesonotum and tegula dark brownish gray, some scales with gray tips, tegula with apex gray to orange gray; fore- leg with femur laterally brownish gray ba- sally, white to gray apically, medially white except for brownish-gray posterior margin, tibia laterally dark brownish gray, medially dark brownish gray on margins, white in middle, tarsomeres laterally dark brownish gray with white apices, medially white mixed with scattered brownish-gray scales; midleg with femur white except for scat- tered dark brownish-gray scales on postero- lateral margin, tibia laterally dark brownish gray intermixed with white to form diago- nal bands at base, middle, and apex, tarso- meres dark brownish gray laterally, first, second, fourth, and fifth tarsomeres with PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Pins euwapeocten isk ER Figs. 3-8. Male and female genitalia. 3, Mompha solomoni male. 4, M. solomeni aedeagus. 5, M. cephal- onthiella male. 6, M. cephalonthiella aedeagus. 7, M. solomoni temale. 8, M. cephalonthiella female. white apices; hindleg with femur white, tib- gray with tarsomere 5 and apices of tarso- ia dark brownish gray with broad white di- meres 1—4 white. Forewing (Fig. 1): agonal bands of setiform scales near base, Length: 3—3.9 mm (16 males), 2.7—3.8 mm middle, and apex, tarsus dark brownish (13 females); ground color dark gray to VOLUME 106, NUMBER 1 brownish gray, scales with white apices ex- cept on basal % of costa; scale tufts at % and %4, each with 5—6 rows of erect scales, basal tuft dark brown, outer tuft white ba- sally, dark brown apically, line of sparse dark brown scales extending apically for short distance from middle of basal tuft, line of dense dark brown scales extending along Rs from anteroapical corner of each tuft, basal radial line bordered by narrow white to orange white line on costal side; costa with dark brown strigulae at about 4%, *%3, and %4; apical strigula connecting with apical radial line near its midlength, areas between basal radial line and dorsum and on each side of apical radial line suffused with light brown, small white spot posterior to end of apical radial line; dark brown api- cal strigula, radial line, and apical margin of pale brown spot forming acutely angled indistinct patch contrasting with band of gray scales with white tips on apical end of wing; margin of wing membrane with line of dark brown scales, expanded to form a spot opposite apical end of radial line in some specimens; apical fringe scales with long white bases from apex to midwing, pale gray from midwing to tornus. Under- side dark grayish brown, some specimens with white near midwing. Hindwing: Uni- formly grayish brown on both surfaces. Abdomen: Grayish brown dorsally, white to orange white ventrally. Male genitalia (Fig. 3, 4, 9-14): Uncus separated from tegumen by membranous line, dorsally rounded at base, ventrally and apically flattened, apex acuminate, laterally with flattened setae at base (Figs. 9-10); anellus divided into two flat plates with curved apicolateral arms; aedeagus tubular with subparallel sides, cornuti absent (Fig. 4); valva (Figs. 3, 12—14) apically cleft for ¥% length of costa, apex rounded in dorsal division, acute in ventral division (saccu- lus), costa lightly sclerotized, densely se- tose apically, sacculus folded dorsally for % length of valva, fold narrowed apically, api- cal third of ventral division more heavily sclerotized than basal *%, sacculus and api- 5 cal third of ventral division sparsely setose (five preparations examined). Female genitalia (Fig. 7): Tergum VIII weakly sclerotized with line of sparse setae on posterior margin; papillae anales facing laterally, sparsely setose; lamella postvagin- alis with two subtriangular projections sep- arated posteriorly by V-shaped cleft, ante- rior end of the cleft with a small dentate projection, lamella antevaginalis forming projecting rim over ostium bursae; ductus bursae heavily sclerotized near ostium bur- Sae; corpus bursae with two bladelike signa connected to crescent-shaped plates (four preparations examined). Holotype.—d, Miss[issippi]., Washing- ton Co., Stoneville, May 1987, J. D. Solo- mon, ex Cephalanthus occidentalis, twig borer. Data are given as on labels except for bracketed information. Deposited in USNM. Paratypes (n = 157).—CANADA: QUE- BEC: Gatineau, Aylmer, 18 rue Washing- ton, 30-31 May 1994 (1 3), 20-24 Jun. 1994, (iG, 1 2); TO=M1 Al 19 945s), B. Landry, at mercury vapor light (BL, SK). UNITED STATES: CONNECTICUT: Windham Co., Chaplin, 0.3 mi. S jct Hwys 6 and 198, 25 Jul. 1992, D:. L.-8Waener (1 2) (UCONN), Hampton, Rock Springs, mines 3 Sep. 1989, adults issued 18 Sep.— 2 Oct, 1989, DEW Lot. 8916, VR= and DL. Wagener (7 3.9 25 tf aw (UCONN, ECK), Thompson, Quaddick Reservoir St. Pk., mines 4 Sep. 1989, adults issued 5 Mar.—4 May 1990, DLW Lot: 89J14, D. L. Wagner (9 6, 9 9, 1 u) (UCONN). DIS- TRICT of COLUMBIA: Jun. 1902, A. Busck (1 ¢) (USNM). FLORIDA: High- lands Co., Lake Placid, Archbold Biol. Sta- tion, 16—22 May 1964, R. W. Hodges (2 u) (USNM); Monroe Co., Big Cypress Natl. Park, Loop Road, vicinity Interport Center, mines 13 Mar. 1991, adults issued 4—19 Apr. 1991, DLW Lot: 91C14, D. L. Wagner, De Re, Davis) and!) Tu DickeloGrig2 41) 9) (UCONN), mines 11 Apr. 1998, D. R. Da- vis (4 6, 5 2) (USNM), Everglades Natl. Park, Anhinga Trail, mines 12 Apr. 1998, 6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 7 as eee Figs. 9-14. Male genitalia of Mompha solomoni. 9, Uncus, lateral (ca. X550). 10, Squamiform setae of uncus (ca. 4500). 11, Piliform setae of uncus (ca. X 10,000). 12, Plumose setae over medial surface of valva (ca. X1,000). 13, Scale cluster over proximomedial surface of valva (ca. X 1500). 14, Detail of plumose setae in Fig. 12 (ca. X6,000). D. R. Davis (6 ¢, 11.2, 1 u) CUSNM). 24, 6 Jul. 1996, T: Harrison, coll. as adult ILLINOIS: Clark Co., Rocky Branch Pre- at UV light (1 6) (INHS), 21 Jun. 1997, T. serve, 3 July 1995, T. Harrison, coll. as Harrison, coll. as adult at UV light (1 2) adult at UV light (1 2) (INHS); Coles Co., (INHS); Mason Co., 1.5 mi. west of Topeka Lake View Park, T12N, R9E, NW % Sec. along C & IM RR in wetland, T22N, R8W, VOLUME 106, NUMBER 1 D2 D2 \ we \e i p1 / ‘ Y 2 / / D2 Wx Ab Ay Tan nee) uf Pap ai yt = ¢ hl ene a Bt ke a) ya | Pay ty ie \ ae) _ fr ea. fe} ae == : u 6 | iy ° ees iS D1/) oe “7, JP H 12 A ee Z el Ve 2p ot soy” aN Oe svi ee = L3 : E al3 lle = \ ore Tu! Ts | q No aaa ), Ie) 5M ; \ mS a sv1\ ‘sy4\ Se aN ‘i a ee | w~ LAY \ OAS 16 lee . AN fr ; oN \ \ { | bees \ [' } | y Su \ Va 4 sa Bae 18 oy ee 19 Figs. 15-19. Larvae of Mompha solomoni, ventral setae not shown. 15, T1—T3 of M. solomoni. 16, A1—A2 of Mompha solomoni. 17, A6—A10 of M. solomoni. 18, M. solomoni mandibles. 19, M. cephalonthiella man- dibles. Sec. 25, J. R. Wiker, larvae boring in stem tip of Cephalanthus occidentalis, 25 May 1995,,emerced: 11 Jun. 1995.@Q id, :2 2) (JRW); Union Co., LaRue Pine Hills, 15 Jul. 1995, T. Harrison and J. Wiker, Iss. 29 Aug. 1995, coll. as larva in leaf mine on Cephalanthus occidentalis (2. 2) (INHS). LOUISIANA: Bossier Parish, Barksdale A.FB., 32°31'13"N 93°35'46"W, 21 May 1996, R. L. Brown and D. Pollock, black- light in calcareous prairie (1 6) (MEM), same data except 32°29'19"N 93°35'20’"W, R. L. Brown, blacklight in calcareous forest (1 2) (MEM). MARYLAND: “5/8 84,” (1 u) (USNM), “°30/[?]7 84” (1 6, genitalia slide, SBA, USNM 5325) (USNM). MAS- SACHUSETTS: Barnstable Co., Barnsta- ble, 26) Jun. 1972, .€. P Kimball, CG dc) (USNM). MICHIGAN: Allegan Co., Alle- gan State Game Area, 2 Jul. 1992, J.-E and B. Landry, at mercury vapor light (1 ¢, 2 2) (CNC); Clinton Co., Bath, 9 May 1957, R. W. Hodges (1 6) (USNM), 1 May 1957, R. W. Hodges (1 2, 1 u) (USNM); Kala- mazoo Co., Portage, 10 Sep. 1995, G. Bal- ogh, coll. as larva in leaf mine on Cephal- anthus occidentalis, Iss. 2 Mar. 1996 (7 6, Peay “(ECK GB: INHS):” MISSISSIPPE Franklin Co., Trib. of McGehee Ck., TON, R4E, Sec. 26SW, 1 Jun. 1992, T. Schiefer, Re Fontenot @l iG. 2), 27 Ful. 1992, ROE: Brown (1 2, genitalia slide TH-MEM 17) (MEM). Porter @reek, ISN; R4E. Sec. 8NW, 1 Jun. 1992, T. Schiefer and R. Fon- tenot (1 6) (MEM); Lowndes Co., 33°18'01"N 88°36'38"W, Black Belt Prairie, 31 May 1994, D. M. Pollock, blacklight in oak-hickory forest (1 ¢) (MEM), same data except T17N, R16E, Sec. 34, 24 Aug. 1993, R. L. Brown and D. Pollock (1 2, genitalia slide TH-MEM 16) (MEM); Smith Co., 1 mi N Raleigh, 14-15 Jun. 1985, R. L. Brown and G. Burrows (1 2) (MEM); Washington Co., same data as holotype (10 3, 8 &, genitalia slides male MEM 491, female MEM 492) (MEM), same data ex- cept 5 May 1986 (1 2) (MEM), 6 May 1986 (3 6, genitalia slide MEM 473) (MEM), 8 May 1986 (1 3), 9 May 1986 (1 2, genitalia slide MEM 320) (MEM), 10 May 1986 (2 3) (MEM), 11 May 1986 (1 2) (MEM), 14 May 1986 (1 3) (MEM), 15 May 1986 (1 6) (MEM); Winston Co., Tombigbee Nat. Forest, 33°19'20"N 89°03'55”W, 25 May 1999, R. L. Brown (1 dj) 1S.Jul 1999) Re Le Brown and J. MacGown (1 2?) (MEM), 13 Sep. 1999, J. A. MacGown (1 2) (MEM). NEW YORK: St. Lawrence Co., Hammond, Oak Point, mines 12-17 Aug. 1988, adults issued 7— 11 Apr. 1989, DLW Lot:, 88H45, D. L. Wapener ‘(2..6,°2 2) (UCONN): SOUTH ~ 8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON CAROLINA: Charleston® Co., Mc- Clellanville, The Wedge, 22 Mar. 1968, R. W. Hodges (1 2) (USNM). VIRGINIA: Falls Church, 1924 (1 &, genitalia slide, SBA, USNM 5302) (USNM). TEXAS: Jas- perm Co. Martin Dies™ State Park, E. C- Knudson 8 May 1986 (2 6, one genitalia slide TH-ECK-18), 14 Nov. 1985 (1 4, genitalia slide ECK-1275); San Jacinto Co., Sam Houston National Forest, Coldspring, 1 May 1988 (1 6, genitalia slide TH-ECK- 1a). Other material examined.—UNITED STATES: OHIO: Hamilton Co., Cincinnati, Ferris Woods, mines 14 Sep. 1912, B. 134, A. Braun 3 6,1 2, 1 vu) CUSNM). Distribution.—Mompha solomoni has been found from Illinois, Michigan, south- ern Québec south to Florida and Texas (Fig. Sil): Etymology.—Mompha_ solomoni is named for James D. Solomon, an entomol- ogist retired from the US Forest Service Southern Hardwoods Laboratory in Stone- ville, MS, who first brought our attention to this species. Egg (Figs. 45, 46).—Size 0.13-0.17 xX 0.22—0.29 mm (n = 5). Glassy, iridescent, oval to round and somewhat flattened, with small nipplelike projection to one side from which numerous small ridges radiate. Larva (Figs. 15—18, 20—31).—Length 3.8-5.6 mm (n = 8). Body cream, with slightly raised dome-shaped spinules; tho- racic legs and pinacula not strongly pig- mented. Head (Figs. 20—26): Hypognath- ous; generally smooth, with some shallow ridges; frons wide, AFI not observed, F1 and AF2 short, Cl much longer than C2 long; labrum shallowly notched; Pl, A3, and S2 about equal in length, longest setae on head; A2 slightly longer than adfrontal setae, Al intermediate in length; L1 poste- rior to A3, short; three mid-dorsal setae very reduced; six stemmata arranged in a C-shaped configuration (Fig. 23); antenna as in Fig. 24; mandible (Fig. 18) broad, with several rounded dentitions; two un- equal setae on outer surface; labium (Figs. 21, 26) with two-segmented palpus, spin- neret parallel sided; maxilla prominent (Fig. 25). Prothorax (Figs. 15, 22): L-group bis- etose, L2 and LI in straight line parallel to median longitudinal axis; LI about twice length of L2; shield with SDI 3—4 times longer than XD2 and SD2; SD2 posterior to SDI; XD1 about % length of SD1; D2 about 4 times length of D1; T2 and T3 with D2 about 2—3 times longer than D1, D2 in vertical line with SD2; SD1 about 2 % times length of SD2; L2 and L3 subequal in length, LI about two times longer than L2 and L3; SV1 slightly posterior to L3; all thoracic legs with a pair of broad bladelike setae ventrolaterad of terminal claw (Fig. 2/1): Abdomen (Figs; 16; 17) 28-3303 3): Prolegs on A3—A6 and A10 of equal size; crochets in a circle, uniserial and uniordinal (Fig. 29); Al-A7 with D2 about 2-—2.5 times longer than D1; SD1 slightly anterior to DI, SDI above spiracle; L2 posteriad of SD1, L1 about 2.5 times length of L2; L3 subequal to Ll; A3—A6 with SV-group tri- setose; SV-group unisetose on Al—A2 and A7—A10; A8& with L-group trisetose, L1, L3, SV1, and V1 roughly in line; A9 with 6 setae, all setae in line; A1O with SDI about twice the length of SD2 and about three times the length of D1 and D2; L- group trisetose. Pupa (Figs. 32—40).—Elongate oval in dorsal and ventral view, widest and slightly depressed dorsally in thoracic region; epi- cranial suture present; anterior tentorial pits visible (Figs. 33, 35); frontoclypeal suture absent and mandibular. and/or piliferal area demarcated as a subovate disk (Figs. 33, 35); caudal portion of antennae adjacent on meson, not separating distally to expose metathoracic legs as in most other Gele- chioidea (Mosher 1916); axillary tubercle (= mesothoracic spiracle) covered with honeycomb latticework (Fig. 36); midab- dominal segments with mediolateral exca- vations (Figs. 32, 37, 38); cremaster with one pair of dorsal setae and three pairs of ventrolateral setae (Figs. 39, 40). VOLUME 106, NUMBER 1 9 Figs. 20-25. Larval head and prothorax of Mompha solomoni. 20, Head, dorsofrontal (ca. *200). 21, Head, ventrofrontal (ca. X 200). 22, Head and prothorax, lateral (ca. * 150). 23, Stemmatal area (ca. X550). 24, Antenna (ca. X 1700). 25, Maxilla (ca. 6500). BIOLOGY Although the new species of Mompha was initially identified as M. cephalonthiel- la, differences in the biologies between these two Cephalanthus-feeding momphi- nes suggested otherwise. Mompha solomoni is a borer in young shoots of Cephalanthus occidentalis early in the growing season but later switches to leafmining. M. cephalon- thiella is believed only to mine leaves of C. occidentalis. The life histories of the two species, as far as known, are given below. 10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 26-31. 650). 28, A3—A6 (ca. X95). 29, Abdominal proleg (ca. X500). 30, A8—A10 (ca. X150). 31, A1O (ca. X200). Shoot-mining broods of Mompha_ solo- moni.—Early instar larvae tunnel in new shoots below the shoot apex in the first week of April in MS. Only tender, succu- lent new growth is utilized. The larval tun- nel meanders about the shoot axis, eventu- ally killing the new growth. Initially, the Larva of Mompha solomoni. 26, Maxillolabial complex (ca. * 1500). 27, Thoracic claw (ca. larva may proceed upward to the meristem, but eventually the tunnel is directed toward the base of the new shoot, where the stem is hollowed out and filled with dark brown frass (Fig. 43). Gallery lengths range from 15 to 55 mm (nm = 60). Of 100+ dissected shoots, about 90% were occupied by a sin- VOLUME 106, NUMBER 1 Figs. 32-34. gle larva, the remaining shoots by two lar- vae; exit holes (Fig. 42) on a few shoots suggest that as many as three larvae mature in a single twig. Prepupal larvae exited mined shoots from 16 April to 7 May in MS. Infested shoots wilt, droop, and within a few days begin to blacken (Fig. 41). Slight swellings, just below the discolored portion of the shoot, may be evident (Fig. 41). The majority of the damaged shoots wither and Pupa of Mompha solomoni (montages at ca. X65). 32, Dorsal. 33, Ventral. 34, Lateral. break away from the plant, although a few swell and split about the gallery and remain on the plant as evidence of previous attack. Wilting and dying shoots were most con- spicuous from 17 April to 15 May (MS). Damage ranged from none to severe, and on some plants, larvae killed every new shoot. A few infested shoots were found in June, indicating that at least a partial second stem-mining generation is produced. Be- 12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 35-40. Pupa of Mompha solomoni. 35, Mandibles and labrum (ca. X300). 36, Mesothoracic spiracle (ca. X300). 37, Al—A2, dorsolateral (ca. * 160). 38, Al—A2, posterodorsal (ca. * 150). 39, A9—A10, dorsal (ca. * 150). 40, Cremaster (ca. 550). cause succulent, thick-stemmed new growth of Cephalanthus is much reduced after the spring flush, only a small percentage of the population is able to locate suitable stem tissue for larval development during the summer and fall months; most individuals develop as leafminers through the rest of the growing season (see below). Natural enemies issued from about 10% of the field-collected stem-mining larvae; these included three braconids (Pholetesor Mason, Bracon E and an undescribed spe- VOLUME 106, NUMBER 1 cies of microgastrine) and an eulophid of the genus Euderus Haliday. Previously re- corded hosts for Pholetesor were all leaf- mining or shelter-feeding Lepidoptera (Whitfield and Wagner 1988). Predaceous thrips were recovered from five dissected shoots that contained dead Mompha larvae. Leafmining broods of Mompha_ solo- monit.—Oviposition on leaves occurs along- side the midrib or a lateral vein on the leaf undersurface, often under tufts of axillary hairs (n = 35). The egg texture is remark- ably similar to that of the leaf surface (Figs. 45, 46). The first instar tunnels from the egg directly into the leaf blade and, in so doing, fills the eggshell with brown or black frass. The mine begins as a hair-thin sinuous track that is extended to the upper leaf surface, such that the mine is visible from both above and below (Figs. 47, 48). Some early instar mines are tortuously contorted and appear blotch-like. Early mines start adja- cent to the midrib (30/54) or a secondary vein (14/54)—none were noted in the lam- ina away from larger veins. The larva may exit the first mine, after tunneling for 5—10 mm, and begin a new serpentine mine else- where, again alongside the midrib or a lat- eral vein. Secondary mines occasionally en- ter vascular tissue, staining it dark brown or black. Mine exit and reentry occurs through the lower leaf surface. Larvae feed ventral side up. After the larva has tunneled for a dis- tance of 10—20 mm, either a new mine is formed or the initial track is enlarged abruptly into a full depth blotch (Fig. 48). All green tissue is removed from within the blotch; in contrast to early mines, the blotch mine is often made along the leaf edge or apex (Fig. 48). The blotch mine is mostly free of frass; the sparse frass is scattered about the mine, staining both leaf surfaces, with most accumulating toward the base or one side of the mine. The mature mine is oval to round with an irregular outline and measures 10—15 X 22—31 mm (n = 5). The pale yellow larva turns smoky red prior to 13 exiting the mine (Fig. 49) to pupate in leaf litter MM = 20). The white cocoon (Fig. 50) is spun be- tween overlapping leaves or in litter below the hostplant. The pupal stage during spring and summer generations lasts 12 to 14 days (n = 30). The majority of fall-collected lar- vae in New England yielded pupae that held over through the winter months, although fall-emerging adults sometimes issued from our collections made in August and as late as early September. An August lot of larvae from Hammond, NY, failed to hatch by late October, when they were placed outdoors; this same collection produced adults in May 1989. A large collection made at Thomp- son, CT, in early September yielded no adults that fall but more than 50 adults the following spring. Yet, another collection made the previous day nearby in Hampton, CT, produced nearly 20 adults over the en- suing five weeks (into mid October). Our observations indicate that M. solo- moni is multivoltine, reaching greatest abundance in late summer and early fall. Larvae and mines can be exceedingly com- mon from August onwards, with a dozen or more mines occurring in a single leaf. Mompha cephalonthiella.—Details of the life history are mostly unstudied. So far the species is only known to mine leaves. Five later instar mines are all full-depth mines, formed along a leaf margin (6—12 * 12-16 mm). In appearance the mines resemble those of M. solomoni (e.g., the two mines on the left hand side of Fig. 48). As in M. solomoni the mine is mostly free of frass. Both Frank Hsu (pers. comm.) and DLW have found abandoned Mompha mines in Cephalanthus in California, where Mompha cephalonthiella is common, with a shield- like oval cut-out at one end of the mine, reminiscent of a mine of Antispila Hiibner (Heliozelidae). Presumably these ovals are fashioned by the prepupal larva and are used to construct a cocoon, just as they are in the Heliozeli- dae. In Florida and the Neotropics an un- described Mompha mines in Hamelia pat- 14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 41-44. at bases of mined shoots. 43, Larval gallery with loosely packed frass. 44, Adult resting posture, note raised scale tufts. ens Jacq. (Rubiaceae). It too may cut-out an oval shield at the mine terminus. Adults of this moth are similar in appearance to those of M. cephalonthiella and M. solo- mont. Chambers (1871: 222) stated that the mine of M. cephalonthiella resembles that of an Antispila, but he elaborated no further. In both Mexico and Costa Rica, DLW has reared additional momphines from melas- tomes whose prepupal larvae also remove oval leaf sections from the mine terminus, although their cut-outs are often more jag- ged than those of the Rubiaceae-feeding momphines. Busck’s (1912) account of Moriloma_ pardella Busck, a momphine leafminer in Conostegia sp. (Melastomata- Life history of Mompha solomoni. 41, Flagging leaves on mined shoots. 42, Larval exit holes ceae), also makes mention of this rather un- usual behavior. We have collected active mines of Mom- pha cephalonthiella from mid June (VT) through early October (CA), contradicting reports by both Chambers (1871) and Forbes (1923) that it is single-brooded. Cer- tainly, populations reach their greatest abundance in late summer (as do those of Mompha solomoni). In our collections adults issued two to four weeks after the larvae were secured—with no pupae hold- ing over to the following season, supporting the suggestion of Chambers (1871) and Forbes (1923) that this species overwinters as an adult. A March record of a flown VOLUME 106, NUMBER 1 Figs. 45-50. Early instar mine. 48, Late instar full depth blotch mines [mines without dense frass packing (left and below) represent secondary mines, primary mines (right) contain considerable frass]. 49, (Reddened) prepupal larva. 50, Cocoons. adult of Mompha cephalonthiella collected by Edward Knudson from Houston, Texas further supports the notion that adults over- winter. DISCUSSION Leafmining moths are usually highly specific in the tissues that they mine (Need- ham et al. 1928, Hering 1951), but mom- Life history of Mompha solomoni. 45, Egg (scale = 100 ppm). 46, Egg (scale = 200 pm). 47, phines provide numerous exceptions. The Palearctic momphines include species that switch between mining leaves and flowers or seeds, e.g., Mompha miscella (Dennis and Schiffermtiller) (Lhomme [1948— 1949]); stems and seed pods, e.g., Mompha sturnipennella (Treitschke) (Emmet 1982); and stems and leaves, e.g., Mompha ochra- ceella (Curtis) and M. propinquella (Stain- 16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 51. ton) (Grabe 1955, Koster 2002). In some species the changes occur during the course of development: The early instars of M. ochraceella mine in the bast of a stem, but later tunnel into the roots to overwinter and then return above ground to mine leaves in the spring (Koster 2002). In other Mompha, changes in feeding habits occur between broods, e.g., M. miscella was reported by Lhomme [1948—1949] to mine leaves in the spring and flowers or seeds in the second generation. The mining habits of Mompha solomoni, involving a switch from shoot mining in the spring to leaf mining in the summer and fall, initially seemed excep- tional to us; however, this change is not so improbable given the variation displayed by Palearctic species. The similarity of scaling characters and genitalia of the shoot boring and leafmining generations suggests that the two represent but a single species. The new species appears to be a member of a species group that includes Mompha cephalonthiella and additional undescribed species, at least one of which is a leafminer ( cal Distributions of Mompha solomoni (triangles) and M. cephalonthiella (circles). in another member of the Rubiaceae, Ha- melia patens. DLW has made collections of this moth in both southern Florida and Cos- ta Rica. Other undescribed taxa that may be members of this species cluster were found among unsorted Floridian momphines in the USNM. Given the phenotypic similarity of the imagoes, the group’s members exhib- it marked biological differences, e.g., Mom- pha cephalonthiella is thought to overwin- ter as an adult and M. solomoni as a pupa. M. cephalonthiella and the Hamelia-feeder may fashion an oval cut-out or shield from the mine terminus, in the same manner as is done by heliozelids; M. solomoni simply drops into leaf litter to make its cocoon. (Curiously, the shield-making behavior ap- pears to be facultative in both M. cephal- onthiella and the new species from- Hamelia.) While many authors have recognized the importance of Onagraceae in temperate ra- diations of the Momphinae (Forbes 1923, Powell 1980, Stehr 1987, Scoble 1992, Hodges 1998, Powell et al. 1998), few have VOLUME 106, NUMBER 1 mentioned Rubiaceae as a host family for momphines. We know of five North Amer- ican species associated with Rubiacae (the two on Cephalanthus treated here and three undescribed species: on Galium, Hamelia, and Spermacoce). Given our present under- standing of the biologies of the North American momphines, which is admittedly incomplete, the Rubiaceae rank third in their importance as larval hosts for Mom- phinae, behind the Onagraceae and Cista- ceae (DLW unpublished data; T. Harrison in litt.). Given the presence of momphines in Central America and the great diversity of Rubiaceae in low latitudes, we think it likely that many other (undescribed) species will be found to be associated with this family. L group is bisetose on T1 (Fig. 22)—a condition unique to the momphines among all Gelechioidea (Stehr 1987, Hodges 1998). Larvae of Mompha solomoni pos- sess a pair of flattened setae below each tar- sal claw (Fig. 27), a feature present in many but not all momphines (Hodges 1998). Mosher (1916) claimed that the frontocyl- peal suture was present in momphids; our preparation of the pupa of M. solomoni (Fig. 35) lacked the suture. One of the most curious features separating the two button- bush-feeding species is the development of the lateral oval pockets that open toward the caudal end of A7 in females of Mompha solomoni (Fig. 7) and M. cephalonthiella (Fig. 8). These pockets are more pro- nounced in M. cephalonthiella than in M. solomoni. Their function is unknown; per- haps they receive the valvae during copu- lation or are in some way involved in pher- omone release. Evidently, these pockets are widespread among momphines (John Wil- terding, personal communication). Additional taxonomic and _ biological work is needed. It would be helpful to doc- ument that the spring-active, stem-mining individuals do, in fact, give rise to the sum- mer-active leafmining generations. Stated differently, confirmation is needed that Mompha solomoni is a single species, and M7 not two closely allied cryptic species, one of which mines only in new shoots. Are Mompha solomoni and M. cephalonthiella sister taxa or do they represent different sections of a Rubiaceae-feeding clade of momphines? Is either species capable of overwintering as both pupae and adults, or is the Overwintering stage a defining trait for each of these moths? ACKNOWLEDGMENTS James Solomon provided numerous col- lections of Mompha solomoni, as well as biological notes and photographs (Figs. 41— 44). Ron Hodges, Jadranka Rota, and John Wilterding made numerous suggestions that improved the manuscript. Terry Harrison shared his personal observations with us and offered numerous helpful suggestions on an earlier version of the manuscript. Specimens were loaned by George Balogh, Terry Harrison, Edward Knudson, Bernard Landry, Jean-Francois Landry, and James Wiker. Shawn Kennedy assisted with the preparation and labeling of the figures. John Steiner, Office for Imaging and Photograph- ic Services, Smithsonian Institution, pro- vided the images of adult Mompha_ solo- mont and M. cephalonthiella. Virge Kask helped with the distribution map and Figure 1. Support for this research was provided by NSF (BSR-9007671) to DLW, the De- partment of Systematic Biology to DA, and NSF (DEB-9200856), Mississippi Agricul- tural and Forestry Station, and Mississippi Natural Heritage Program to RLB. LITERATURE CITED Busck, A. 1912. Descriptions of new genera and spe- cies of microlepidoptera from Panama. Smithson- ian Miscellaneous Collections 59: 1—10. Chambers, V. T. 1871. Micro-lepidoptera. Canadian Entomologist 3: 221—224. Clarke, J. EK G. 1941. The preparation of slides of the genitalia of Lepidoptera. Bulletin of the Brooklyn Entomological Society 36: 149-161. Emmet, A. M. 1982. Notes on Mompha nodicolella Fuchs (Lep.: Momphidae). Entomologist’s Record and Journal of Variation 94: 21—24. Forbes, W. T. M. 1923. The Lepidoptera of New York and Neighboring States. Primitive Forms, Micro- 18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON lepidoptera, Pyraloids, Bombyces. Cornell Uni- versity Agricultural Experiment Station Memoir 68. 729 pp. Grabe, A. 1955. Kleinschmetterlinge des Ruhrgebietes. Mitteilungen aus des Ruhrlandmuseums der Stadt Essen 177: 1-159. Hering, E. M. 1951. Biology of the Leaf Miners. Dr. W. Junk, s’Gravenhage, Netherlands, 420 pp. Hodges, R. W. 1992. Two new species of Mompha from California (Lepidoptera: Momphidae). Jour- nal of the New York Entomological Society 100: 203-208. 1998. The Gelechioidea. pp. 131-158. In Kristensen, N. P., ed. Lepidoptera, Moths, and Butterflies. Volume |: Evolution, Systematics, and Biogeography. Handbook for Zoology. Volume IV. Arthropoda: Insecta. Walter de Gruyter, Berlin, Germany, 491 pp. Hodges, R. W. et al., eds. 1983. Checklist of the Lep- idoptera of America North of Mexico. E.W. Clas- sey and Wedge Entomological Research Founda- tion, London, England, 284 pp. Kornerup, A. and J. H. Wanscher. 1978. Methuen Handbook of Colour. Third Ed. Methuen and Company, Ltd., London, 252 pp. Koster, J. C. 2002. Momphidae. pp. 224-254. In Em- met, A. M. and J. R. Langmaid, eds. The Moths and Butterflies of Great Britain and Ireland, Har- ley, Martins, England. Lhomme, L., [1948-1949]. Cosmopterygidae. pp. 677-699. In Lhomme, L., 1935—-[1963], Cata- logue des Lépidopteéres de France et de Belgique. 2. Microlépidopteres. Le Carriol, Douelle (Lot), France, 1253 pp. Mosher, E. 1916. A classification of the Lepidoptera based on characters of the pupa. Bulletin of the Illinois State Laboratory of Natural History 12: 15-159. Needham iG2aS, WW. Erost, and Be Heri S28: Leaf-mining Insects. Williams and Wilkins, Bal- timore, Maryland, 351 pp. Poole, R. W. and P. Gentili, eds. 1996. Nomina Insecta Nearctica. Volume 3. Diptera, Lepidoptera, Si- phonaptera. Entomological Information Services, Rockville, Maryland, 1143 pp. Powell, J. A. 1980. Evolution of larval food prefer- ences in microlepidoptera. Annual Review of En- tomology 25: 133-159. Powell, J. A., C. Mitter, and B. Farrell. 1998. Evolu- tion of larval food preferences in Lepidoptera, pp. 403—422. In Kristensen, N. P., ed. Lepidoptera, Moths, and Butterflies. Volume |: Evolution, Sys- tematics, and Biogeography. Handbook for Zool- ogy. Volume IV. Arthropoda: Insecta. Walter de Gruyter, Berlin, 491 pp. Scoble, M. J. 1992. The Lepidoptera. Form, Function, and Diversity. Oxford University Press, Oxford, 404 pp. Stehr, E W. 1987. Momphidae. p. 389. Jn Stehr, E W., ed. Immature Insects. Kendall-Hunt, Dubuque, lowa, 754 pp. Whitfield, J. B. and D. L. Wagner. 1988. Patterns and host ranges within the parasitoid genus Pholetesor Mason (Hymenoptera: Braconidae). Environmen- tal Entomology 17: 608-615. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 19-25 A NEW SPECIES OF PROCONTARINIA (DIPTERA: CECIDOMYHDAE), AN IMPORTANT NEW PEST OF MANGO IN THE PHILIPPINES RAYMOND J. GAGNE AND CELIA DR. MEDINA (RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, Smithsonian Insti- tution, Washington, DC 20560-0168, U.S.A. (e-mail: rgagne @sel.barc.usda.gov); (CdRM) National Crop Protection Center, University of the Philippines, Los Banos College, La- guna 4031, Philippines (e-mail: celia@laguna.net) Abstract.—A new species of cecidomyiid, Procontarinia frugivora Gagné, is reported from mango, Mangifera indica (Anacardiaceae), in Luzon Island, Philippines, where it has become a serious pest. Adults, pupae, and larvae are described, illustrated, and com- pared to other Procontarinia species. Erosomyia is a new junior synonym of Procon- tarinia, so Erosomyia mangiferae Felt is newly combined in Procontarinia. Procontar- inia mangiferae (Felt 1916) becomes a new junior homonym of P. mangiferae (Felt 1911), so is given the new replacement name P. biharana Gagné. Rabdophaga man- giferae Mani is newly referred to Procontarinia where it is made a new synonym of P. mangiferae (Felt 1911). Key Words: In February 2002, gall midge larvae were found exiting from holes of mango fruit (Fig. 1) in Bulacan Province, Luzon Island, Philippines (15.04°N, 121.02°E). Shortly af- ter, adults were reared from these larvae that proved to be a species of Procontarinia Kieffer & Cecconi unlike any of the other 11 species of that genus, all of which make galls on mango leaves. The new species de- scribed here was also referable to Eroso- myia Felt, which was known from a single species, Erosomyia mangiferae Felt, a gall- former on mango shoots. Upon investiga- tion, the two genera were found to share distinguishing characters and so are syn- onymized here. Mango, Mangifera indica L. (Anacardi- aceae), is a tropical species of Indo-Bur- mese origin. It is widely cultivated pantrop- ically and even subtropically, from 36°N to 33°S. In the Philippines, it is the second mango, gall midges, new species most important fruit crop in terms of do- mestic consumption and export value. The damage caused by the newly described ce- cidomyiid seriously affects the quality and yield of mango. Infested fruit initially show small brownish lesions of | mm diameter that grow larger and deeper as the fruit en- large. Most infested fruit fall to the ground before ripening; those that reach maturity are not marketable. The population out- break appears at present to be localized on Luzon Island in the provinces of Bulacan, Cavite, and Bataan. METHODS Infested fruit were collected and kept in containers lined with paper towels until full-grown larvae emerged. Larvae were then placed in individual vials with vermic- ulite, where pupation and eventual adult emergence occurred. Specimens of imma- 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. ture stages and reared adults were preserved in 70% isopropyl alcohol. Samples were mounted on microscope slides using the method outlined in Gagné (1989). Termi- nology for adult morphology follows usage in McAlpine et al. (1981) and for larval morphology that in Gagné (1989). Larvae were obtained and adults reared by C. dR. Medina who is studying the biology and impact of this pest in the Philippines. The taxonomic investigation was the responsi- bility of R. J. Gagné. Procontarinia frugivora Gagné, new species (Figs. 2-13) Adult.—Head: Eyes connate, 7—8 facets long at vertex; facets circular, closely ad- jacent but not abutting, lateralmost facets farther apart, separated by up to one facet diameter. Occiput with dorsal protuberance with 2 apical setae. Frons with 4—5 setae. Labella ellipsoid, each with 5—6 lateral se- tae. Palpus 4-segmented, first segment Young mango fruit with holes made by larvae of Procontarinia frugivora. slightly longer than wide, remaining seg- ments about twice as long as wide, the two distal segments not as wide as first two seg- ments. Antenna: Scape cylindrical, as wide as long, with | seta on outer lateral surface and |—3 setae on inner lateral surface; ped- icel with 5—6 setae on inner lateral and ven- tral surfaces combined; with 12 flagello- meres. Male flagellomeres (Fig. 2) binodal; one circumfilum on each node, loops of the circumfila subequal in length; nodes coy- ered with setulae. Female flagellomeres (Fig. 3) cylindrical, slightly constricted at middle, surrounded by two appressed cir- cumfila connected by two longitudinal bands; necks slightly longer than wide. Thorax: Wing unmarked, 1.2—1.3 mm long in male (n = 5), 1.5—1.6 mm long in female (n = 5), R; curved toward apex, joining C posterior to wing apex, Rs present as stub near base of R;. Mesanepimeron with 3—4 setae, thoracic pleura otherwise bare. Tarsal claws (Fig. 4) toothed, curved VOLUME 106, NUMBER 1 near midlength; empodia attaining bend in claws. Male abdomen: First through sixth ter- gites entire, rectangular, with single poste- rior row of setae, no lateral setae, scattered scales, and 2 anterior trichoid sensilla; sev- enth tergite as for preceding but with fewer scales, unsclerotized posteriorly and lacking the posterior row of setae and scales; eighth tergite less sclerotized than preceding ter- gites, the only vestiture the anterior pair of trichoid sensilla. Second through eighth sternites rectangular, with single posterior row of setae, 2 lateral and one mesal group of setae near midlength, and 2 anterior trichoid sensilla; eighth sternite foreshort- ened, midlength setal groups consequently abutting posterior setae. Genitalia (Figs. 5— 7): cerci nearly completely fused, only a short incision between them present poste- riorly, with several posterior setae; hypo- proct narrower than conjoined cerci, nar- rowing slightly from base to broadly con- vex apex, with a few posterior setae; ae- deagus about as long as gonocoxite, cylindrical, rounded at apex, laterally with longitudinal rows of sensory pits; gonocox- ite elongate-cylindrical with large, conical mesobasal lobe, bearing scattered cuticular spines but devoid of setulae; gonostylus elongate-cylindrical, tapering gradually from basal third to distal tooth, with setulae near base and covered beyond with minute carinae and widely scattered short setae. Female abdomen (Figs. 8—10): First through seventh tergites and second through seventh sternites as for male. Eighth tergite weakly sclerotized (Fig. 8, arrow), separation from seventh tergite sub- equal to length of eighth, with single row of short posterior setae and anterior pair of trichoid sensilla the only vestiture. Eighth sternite not evident. Ovipositor slightly pro- trusible, venter of eighth segment and dor- sum of ninth and tenth segments without vestiture, venter of ninth segment with se- tae; cerci entirely connate, short, convex apically, with 2 pairs apical sensory setae 21 and scattered setae elsewhere; hypoproct short, narrow, with 2 posterior setae. Pupa.—Unknown, not preserved. Third larval instar (Figs. 11—13).— Length, 1.6-1.9 mm. White. Integument smooth except for several horizontal rows of tiny spicules on anteroventral surfaces of the first through seventh abdominal seg- ments. Head with posterior apodemes lon- ger than head capsule. Antenna about twice as long as wide. Spatula (Fig. 12) clove- shaped with 2 acutely triangular anterior teeth. Lateral thoracic papillae on each side of central line (Fig. 12) in 2 groups, a triplet and a singlet, 2 papillae in each triplet each with tiny seta, remaining lateral setae with- out. Dorsal and pleural papillae with setae no longer than wide. Terminal segment (Fig. 13) narrowed abruptly from eighth segment, blunt posteriorly, with 8 papillae as follows: 1 mesoposterior pair large, cor- niform; | pair between the 2 corniform pa- pillae each with a seta no longer than wide; and 2 pairs of papillae laterally, each with short seta several times longer than wide. Area between terminal corniform papillae not pigmented. Holotype.—Male, emerged 20-II-2002 from larvae fallen from mango fruit, Ala- gao, Bulacan, Philippines, collected by C. dR. Medina, deposited in the University of Philippines Museum of Natural History, Los Banos, Laguna, Philippines. Other material examined.—Same data as holotype, 4d, 52, 5 larvae, all with same relevant data as holotype, deposited in the University of Philippines Museum of Nat- ural History and the National Museum of Natural History, Washington, DC, USA. Etymology.—The specific name, /rugi- vora, 1S an adjective that means “fruit eat- ing. Discussion.—The genus Procontarinia Kieffer & Cecconi (1906) was described for Procontarinia matteiana Kieffer & Cecconi (1906), reared from leaf galls found on mango grown in the Botanical Gardens in Palermo, Sicily. Felt (1911) described Ero- somyia tor Erosomyia mangiferae Felt gall- 22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 5 Figs. 2-7. Procontarinia frugivora. 2, Male third antennal flagellomere. 3, Female third antennal flagello- mere. 4, Tarsal claw and empodium. 5, Male genitalia, only left gonopod shown (dorsal). 6, Aedeagus flanked by mesal lobes of gonocoxites (dorsal). 7, Nearer mesal lobe of gonocoxite, aedeagus, and farther gonopod (mesal). ing shoots of mango on St. Vincent. Felt gave no indication that he took Procontar- inia into account when describing his ge- nus. When one considers these two genera together one finds that they not only lack differentiating characters but share several apomorphies. The 11 species previously in Procontarinia, including Erosomyia man- gicola Shi (1990) recently transferred to Procontarinia by Uechi et al. (2002) and Erosomyia mangiferae share the following derived characters: male flagellomeres each have two circumfila per flagellomere (Fig. 2) instead of three; male cerci are more or less fused (Fig. 5) instead of separated from the base; the gonocoxite has a definite me- sobasal lobe (Fig. 5), which may be low and rounded or conical, depending on the species; the female eighth tergite is weakly developed and, in addition, separated from the seventh tergite by more than the length of the eighth tergite (Fig. 8); the female cer- ci are short and partially or completely fused into a single lobe (Fig. 9); and the larvae have only four lateral setae, a triplet and a singlet on either side of the spatula VOLUME 106, NUMBER 1 lj y f Ea -“ y cS S) Po 2 Figs. 8-13. Procontarinia frugivora. 8, Female postabdomen, seventh segment to fused cerci (dorsolateral): arrow points to eighth tergite. 9, Female fused cerci, detail (dorsal). 10, Female ninth segment and fused cerci, detail (lateral). 11, Outline of third instar larva (dorsal). 12, Larval spatula and associated papillae (ventral). 13, Posterior larval segments, detail (dorsal). i) i>) 24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Fig. 10), instead of two triplets. Other characters shared by these species are the toothed claws (Fig. 4), probably a plesiom- orphy, and the fact that one pair of larval terminal papillae are enlarged and recurved (Fig. 11). Because of the above shared char- acters between Erosomyia and Procontari- nia, the two genera are considered syno- nyms here (new synonymy). The larvae are similar to those of Contarinia, which may indicate a relationship between the two gen- era. All of the previously described species now in Procontarinia are from mango and all make leaf galls, except P. mangiferae Felt, which causes swellings on shoots. Pre- viously described species are as follows: allahabadensis Grover 1962: 312, Amradi- plosis (as Amraeomyia in error), from India. amraeomyia Rao 1950: 37, Amraeomyia, from India. biharana Gagné, new name for /ndodi- plosis mangiferae Felt, from India. mangiferae Felt 1916: 403, Indodiplosis, (new junior secondary homonym of Erosomyia mangiferae Felt 1911). brunneigallicola Rao 1950: 39, Amraeo- myia, from India. echinogalliperda Mani 1947: 443, Amra- diplosis, from India. keshopurensis Rao 1952: 52, Amraeomyia, from India. ; mangicola Shi 1980: 131, Erosomyia, from China, Guam, and Japan, where it is evidently a recent introduction (Uechi etiale 2002): schreineri Harris, in Harris & Schreiner 1992: 42, Procontarinia. mangiferae Felt 1911: 49, Erosomyia, new combination, from India, Réunion: immigr.: West Indies (Guadeloupe, St. Lucia, St. Vincent, and Trinidad) and Brazil. mangiferae Tavares 1918: 48, Mangodi- plosis, junior secondary homonym of mangiferae Felt. mangiferae Mani 1938: 331, Rhabdopha- ga, new junior secondary homonym of mangiferae Felt. Mani evidently placed this species in Rabdophaga be- cause of the fused cerci of the female. That character and the fact that the specimens were reared from mango shoot galls indicate that the species is properly placed here. New synonym. indica Grover & Prasad 1966: 7, Eroso- myld. mangtifoliae Grover 1965: 115, Indodiplo- sis, from India. matteiana Kieffer & Cecconi 1906: 135, Procontarinia, from India, Kenya, Mauritius, Réunion, and Java. tenuispatha Kieffer 1909: 150, Oligotro- phus, from India. viridigallicola Rao 1950: 34, Amraeomyia, from India. Illustrations of male genitalia of all of the above species accompany the original de- scriptions, except for the Felt species, which I have been able to study. All Pro- contarinia species differ from the new spe- cies, P. frugivora, in having setulae on the mesobasal lobes of the male genitalia in- stead of being devoid of setulae and bearing cuticular spines as in the new species. Fur- ther, only the new species and P. mangifer- ae have completely fused female cerci, al- though at least P. mangicola has almost completely fused cerci. Larvae of all Pro- contarinia species except the new species appear to have a sclerotized and pigmented area between the corniform papillae of the terminal segment. ACKNOWLEDGMENTS We thank P. Malikul for making the slide preparations, Lucrecia Rodriquez for com- puter assistance in making the plates, the Bureau of Agricultural Research, Depart- ment of Agriculture, Philippines for finan- cial support of CdRM’s research, Aile S. Alba for assistance in collecting specimens, and, for their comments on drafts of the manuscript: K. M. Harris, international In- stitute of Entomology, CAB, London; S. VOLUME 106, NUMBER 1 McKamey and A. L. Norrbom, Systematic Entomology Laboratory, USDA, Washing- ton, DC, USA; and J. Yukawa, Kagoshima University, Fukuoka, Japan. LITERATURE CITED Felt, E. P. 1911. A generic synopsis of the Itonidae. Journal of the New York Entomological Society 19: 31-62. . 1916. New Indian gall midges. The Canadian Entomologist 48: 400—406. Gagné, R. J. 1989. The Plant-Feeding Gall Midges of North America. Cornell University Press, Ithaca, New York, xili & 355 pp. & 4 pls. Grover, P. 1962. Studies on gall midges (Itonididae: Cecidomyiidae: Nematocera: Diptera) from India, VI. Proceedings of the National Academy of Sci- ences, India (B) 32: 312-318. . 1965. Studies of Indian gall midges XV: One new genus and one new species of Bifilini (Ce- cidomyiidae: Diptera). Marcellia 32: 111—126. Grover, P. and S. N. Prasad. 1966. Studies on Indian gall midges XVI: Four species of gall midges (Ce- cidomyiidae: Diptera) affecting inflorescence of mango. Cecidologia Indica 1(1): 1-19. Harris, K. M. 1992. Pp. 41—44. Jn Harris, K. M. and I. H. Schreiner. A new species of gall midge (Dip- tera: Cecidomytlidae) attacking mango foliage in Guam, with observations on its pest status and biology. Bulletin of Entomological Research 82: 41-48. Kieffer, J.-J. 1909. Description de galles et d’insectes gallicoles d’ Asie. Marcellia 7: 149-167, pls. HII- IV. Kieffer, J.-J. and G. Cecconi. 1906. Un nuovo dittero i) Nn galligeno su foglie di Mangifera indica. Marcellia 5: 135-136. McAlpine, J. E, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds. 1981. Manual of Nearctic Diptera. Vol. 1. Re- search Branch, Agriculture Canada, Monograph 27. vi & 674 pp. Mani, M. S. 1938. Studies on Indian Itonididae (Ce- cidomytidae: Diptera). V. Descriptions and records of midges in the Pusa Collection. Records of the Indian Museum 40: 331-336. . 1947. Some new and imperfectly known gall midges (Itonididae: Diptera) from India. Bulletin of Entomological Research 38: 439-448. Rao, S. N. 1950. Descriptions of one new genus and three new species of gall midges (Itonididae: Dip- tera), causing galls on mango in India. The Indian Journal of Entomology 9(1947): 33—40. . 1952. Some new species of gall midges (Iton- ididae: Diptera) from India. Proceedings of the Royal Entomological Society of London 21: 49— 35) D. 1980. A new species of gall midge affecting young leaf of mango (Diptera: Cecidomyiidae). Entomotaxonomia I1: 131—134. Tavares, J. S. 1918. Cecidologia brazileira: cecidias que se criam em plantas das familias das Verben- Shi, aceae, Euphorbiaceae, Malvaceae, Anacardiaceae, Labiatae, Rosaceae, Anonaceae, Ampelidaceae, Bignoniaceae, Aristolochiaceae e Solanaceae. Brotéria, Série Zoologica 16(1): 21—48. Uechi, N., EF Kawamura, M. Tokuda, and J. Yukawa. 2002. A mango pest, Procontarinia mangicola (Shi) comb. nov. (Diptera: Cecidomyiidae), re- cently found in Okinawa, Japan. Applied Ento- mology and Zoology 37: 589-593. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 26-34 IOWANA FRISONI HOTTES (HEMIPTERA: APHIDIDAE) REDESCRIBED, WITH NOTES ON ITS BIOLOGY COLIN FAVRET, JOHN EF TOOKER, AND LAWRENCE M. HANKS (CF) Hlinois Natural History Survey, Center for Biodiversity, 607 E. Peabody Dr., Champaign, IL 61820, U.S.A. (e-mail: crf@uiuc.edu); (JFT, LMH) University of Illinois at Urbana-Champaign, Department of Entomology, 505 S. Goodwin Av., Urbana, IL 61801, WeSAc Abstract.—lowana frisoni Hottes was collected and studied for the first time since its original collection in 1925. It was found in remnant prairies where it fed on the basal stem of the endemic plants Si/phium laciniatum L. and S. terebinthinaceum Jacquin (As- teraceae). Later in the season it fed on the stem or leaf stipules within the cavity created by the leaf axil. The aphid was always tended by ants, Lasius flavus (FE), L. neoniger Emery, and two Crematogaster species, which housed the aphids by building soil collars around the bases of plants and sealing leaf axils with soil and dead plant material. The ants also carried /. frisoni underground when the aphid colony was disturbed. We rede- scribe the apterous vivipara morph of /. frisoni and describe for the first time the fundatrix, alate vivipara, male, ovipara, and egg. Key Words: In 1925, G. Hendrickson collected a few aphids in a leaf axil of Si/phium laciniatum L. (compass plant, Asteraceae) in Ames, Iowa. Twenty-nine years later, EK C. Hottes described a new genus and species based on two specimens (Hottes 1954). As far as we can surmise, the aphid, Jowana frisoni Hottes (Hemiptera: Aphididae), had not been seen in the 75 years since its original collection. In the summer of 2000, we discovered J. frisoni feeding on flowering stems of S. la- ciniatum and S. terebinthinaceum Jacquin (prairie dock) at Loda Cemetery Prairie Na- ture Preserve (~1.4 ha in area; Iroquois Co., IL, 40°31.62’N X 88°4.57'W), a prairie remnant (White 1988). Both S. laciniatum and S. terebinthinaceum are common pe- rennials in prairies of central Illinois and produce as many as 12 flowering stems per plant that can reach heights of 2—4.5 m (Weaver 1954). Silphium, Lasius, endemic species, prairie remnant, symbiosis Here, we redescribe the /. frisoni apter- ous vivipara and describe for the first time the fundatrix, alata, male, ovipara, and egg. We also discuss the biology of the aphid with regard to host plants and attendant ants, Lasius flavus (EF), L. neoniger Emery, and two unidentified Crematogaster spe- G1es. MATERIALS AND METHODS To evaluate the distribution of /. frisoni in east central Illinois, we surveyed three other prairie remnants and three prairie res- torations where both species of Si/phium were present, thoroughly canvassing these sites for aphids on plants of both species. Remnants were Prospect Cemetery Prairie Nature’ Preserve: (~2” has 402671 Nw 88°5.87'W) and Paxton Railroad Prairie (~2 ha; 40°26.17’N X 88°06.36'’W) in Ford County and Grant Creek Prairie Nature Pre- VOLUME 106, NUMBER 1 serve (~31 ha; 41°22.10'’N X 88°11.55'’W) in Will County. The restorations were all in Champaign Co.: Red Bison Prairie Corridor (~1.7 ha; 40°4.81’'N X 88°14.83’W), Tre- lease, Prac \(~7.3. tha; <40°7.76'N: x 88°8.59’'W), and Meadowbrook Prairie (~24 ha; 40°4.72’N X 88°12.41’W). See Tooker et al. (2002) and Tooker and Hanks (in press), for more information on sites. To estimate the abundance of /. frisoni and its attendant ant species at Loda Cem- etery Prairie, we established five east-west transects across the long axis of the prairie (~200 m). We walked transects on 12, 19, and 26 July, 27 August, and 6 September 2002, examining plant stems within | m of transects for J. frisoni and ants (n = 75 S. laciniatum stems, 106 S. terebinthinaceum stems). We also assessed the prevalence of I. frisoni at Prospect Cemetery Prairie in August 2002 by walking a single transect (~50 m long) through a patch of S. fere- binthinaceum and recording the presence of I. frisoni and ants. Because flowering stems of S$. laciniatum were not abundant at that site, we examined all plants of that species. We collected aphids (n = 42) and ants (n > 30) during the growing seasons in 2001 and 2002. On 6 October 2002, a colony was brought back to the lab and reared on a cut stem of S. terebinthinaceum. This colony produced males (n = 6), oviparae (n = 8), and eggs (n = 11), which were laid ran- domly along the stem under a leaf stipule. Collected aphids were cleared with a KOH and chloral phenol procedure and _slide- mounted in Canada balsam. Species iden- tity was confirmed with the holotype (the single paratype is a nymph) and associated ants were identified. Aphid and ant speci- mens are deposited at the Illinois Natural History Survey insect collection and ex- emplars of each aphid morph were sent to the National Museum of Natural History (aphid collection, Beltsville, MD). Fifty- seven aphids were measured with an ocular micrometer on a compound microscope. Specimen photos were taken using a Nikon Eclipse E600 microscope, Spot Insight dig- 27 ital camera model 3.2.0, and Spot Ad- vanced software (Diagnostic Instruments, Inc., Sterling Heights, MI). Because no obvious morphological dif- ferences distinguished three putative fun- datrices, we performed canonical discrimi- nant function analysis to confirm that they were indeed morphometrically distinct from the other apterous viviparae. Discriminant factor analysis of aphids has been used to discriminate between species (Brown and Blackman 1994) and populations of the same species (Foottit and Mackauer 1980). Hand (1986) used this method to discrimi- nate holocyclic and anholocyclic alate aphids of the same species. To confirm that aphids collected as solitary adults early in the season were fundatrices and not other Viviparae, discriminant function analysis was conducted (SYSTAT® 10 software, SPSS Inc. 2000) using 10 characters to con- firm morphological distinctness (Albrecht 1980): lengths of the whole body, antennal segment II, antennal segment IV, the base of antennal segment V, the processus ter- minalis of antennal segment V, rostral seg- ment III, siphunculus, metafemur, metati- bia, and counts of rhinaria on antennal seg- ment II. To maintain a large sample size, some typical measurements (ultimate rostral segment, for instance) were omitted if they were not available for all three putative fun- datrices. The groupings in the analysis were ap- terous viviparae, alate viviparae, Oviparae, and males; we also added the three putative fundatrices and the holotype (apterous vi- Vipara) to determine their association with the other groups. If putative fundatrices did not group with apterous viviparae, for ex- ample, we could conclude that they were indeed a distinct morph. To control for pos- sible seasonal variation in aphid morpho- metrics (Debaraj and Singh 2000), we used apterous viviparae from throughout the growing season, including early-season col- lections made soon after the putative fun- datrices had been collected. We tested for 28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Second Discriminant Function Oo Fundatrices © ie) Apterous viviparae Oviparae Holotype First Discriminant Function Fig. 1. Graph of first and second discriminant factors showing the fundatrices as distinctly different from the other apterous viviparae. Ovals are 95% confidence intervals. distinctness using the first through fifth dis- criminant factors. RESULTS AND DISCUSSION Populations of /. frisoni were present in all four prairie remnants but were not ob- served in any of the three restorations, sug- gesting it may be a remnant dependent spe- cies (Panzer et al. 1995). We did not search for or collect the ant species in the resto- rations. The aphid fed on both S. laciniatum and S. terebinthinaceum at all four sites, but not on Silphium integrifolium Michaux, which was also present at all four sites. Nei- ther have we seen /. frisoni on Silphium perfoliatum L. at Meadowbrook Prairie or several railroad rights-of-way in east central Illinois. Because all colonies of J. frisoni ob- served were tended in summer by the ants L. flavus, L. neoniger, or one of two Cre- matogaster species, it seems likely that the symbiosis is obligate, as is true for some other aphid species (Eastop 1953, Carter 1983). In October, ants were absent in the several declining aphid colonies, the ants having retreated below ground to overwin- ter. The ant species never co-occurred on the same plant, but occasionally tended aphids on adjacent plants. Lasius flavus is a common species that ranges across North America and Eurasia (Wilson 1955), may be completely subterranean, and tends a di- versity of aphid species (Wilson 1955, Zwo6lfer 1958). This ant is known to carry aphid eggs into its nest for overwintering (Pontin 1960), but it also may be an aphid predator (Pontin 1958, 1978). Lasius neo- niger is a dominant ant species in open, grassy habitats of eastern North America and commonly tends aphids, including the corn root aphid, Aphis maidiradicis Forbes (Wilson 1955, Traniello and Levings 1986), which also may be entirely dependent on VOLUME 106, NUMBER 1 ants (Hottes and Frison 1931). The identity of the two Crematogaster species remains uncertain because their colonies appear rare and we have collected very few individuals, even after further searches during the sum- mer of 2003. Nevertheless, the genus is cos- mopolitan (H6lldobler and Wilson 1990) and tends hemipterans, including other aphid species (Dejean et al. 1991). Ant species appeared to nest at the base of individual plants and built a collar of soil (~4-8 cm in height) encircling the bases of a few to several flowering stems of S. la- ciniatum and S. terebinthinaceum (we did not observe the aphid on leaf stems). Jo- wana frisoni fed within this ant-built ref- uge. Some soil collars had tunnels isolated from the stem through which ants could move around the base of the plant and down into the nest. When /. frisoni colonies were disturbed, the ants of all species car- ried the aphids underground, moving the entire colony within the course of a few minutes. Aphids also occurred in leaf axils feeding on either the stem or clasping leaf stipules, where they were also tended by ants (Fig. 2). Lasius neoniger usually sealed these leaf axils with soil and/or dead plant material, housing the aphids, but did not transport aphids from these feeding sites. Late in the season (September), a few plants of both species had ants and aphids present in every leaf axil and even in axils of branches and modified leaves in proximity to flowers (>2 m above the ground). In our transects at Loda Cemetery Prai- rie, /. frisoni was present on only one of 75 S. laciniatum plants (1.3% of all plants) and this one aphid colony was tended by L. neo- niger. lowana frisoni was present on 34 of 106 (32%) S. terebinthinaceum plants and the majority of these were tended by L. neo- niger. At Prospect Cemetery Prairie, /. fri- soni Was present on three of five S. lacinia- tum (40%) and 10 of 12 S. terebinthina- ceum (83%), and L. neoniger was the dom- inant ant species tending these aphid colonies. The first discriminant factor did not dis- 29 tinguish the fundatrices as separate from the other apterous viviparae. This was not un- expected because the first discriminant fac- tor may be a function of size (Foottit 1992) and the greatest size variation was between alatae and apterae, and not between funda- trices and other apterae. The second dis- criminant factor did separate the putative fundatrices from the other apterous vivipa- rae (Fig. 1), but the third, fourth and fifth factors did not discriminate any of them. Based on these results we believe that our three specimens were indeed fundatrices and we describe them as such below. DESCRIPTION Iowana frisoni Hottes 1954 (Figs. 2-19) Apterous vivipara (Table 1, Figs. 3, 7, 10, 15).—Color (alive): Body pale green. Adults with head and prothorax light red- dish brown. Eyes dark brown to red. All of ant. I, ant. II, and proximal % of ant. II segments pale green; distal *% of ant. III, and all of ant. I1V and ant. V brown. Legs pale green except distal-most portion of tib- iae and both tarsal segments brown. Si- phunculus brown. Morphology of cleared, slide-mounted specimens: See Table 1 for anatomical measurements. Body entirely pale, neither sclerites nor scleroites present (Fig. 3). Head: Frons flat, smooth with a few scattered minute hairs. Rostrum reaches abdominal segment I. Rostral segments III to V dark, each darker than one before (Fig. 7). Rostral segment IV with 2 small acces- sory setae (Fig. 9), these may not be visible in some specimens. Antenna 5 segmented, dark and imbricated beyond base of ant. III (Fig. 10). Ant. IT with O0—21 apical and sub- apical rhinaria. Ant. [IV with a single apical primary rhinarium. Thorax: Large protho- racic tubercles, usually small lateral tuber- cles on thoracic segments 2 and 3 which may be hard to see on some specimens. Furculum broadly V-shaped (Fig. 7). Legs short. Metatibia darkened distally only, with darkening continuing into tarsus; no 30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 5 oe 6 Figs. 2-6. Jowana frisoni. 2, Feeding on the inside of a leaf stipule of Silphium laciniatum, tended by Crematogaster sp. 3, Apterous vivipara. 4, Alate vivipara. 5, Male. 6, Ovipara. (ao) VOLUME 106, NUMBER 1 (prO'0-Fr0'0) FrO'O (CL0'0-£90'0) 8900 (760'0-690'0) 6L0'°0 (¢90'0-7r0'0) OSO'0 (LF0'0-8£0'0) Tr0'0 (LOI°0-C80'0) €60°0 (F60°0-880'0) 160°0 (1¢0°0-$T0'0) 870°0 (Iv'0-9€'0) 6£°0 (C7 O-17'0) €70 (CEL O-611'O) 9TT0 (6110-1010) 8010 (971 0-¥60'0) 8010 (€T1'0-C80'0) 1600 (880°0-SL0'0) 8L0°0 (O9LT'O-StI1'0) 19TO (T81O-1ST'O) OLT0 (So TelOa viet (9 = u) Jeu snoiajdy (OSO'0-1£0'0) £700 (880°0-690'0) 8Z0'0 (SEI O-LOT'O) ZITO (LOL 0-P60'0) £010 (690'0-0S0'0) 190°0 (OCT O-LST 0) O80 (6110-1010) 8010 (rr0'0-1¢£0'0) 6£0°0 (S9'0-0S'0) SSO (8¢'0-0£'0) €£°0 (681 O0-€91'0) 9LT 0 CUS Ome IO) MVE TO (COT O-61 10) TFI'O (971 0-690'0) TOTO (6110-1010) 6010 (p9T'0-T81'0) SIT O (977 O-S61'0) LITO (ZO'T-OS'T) OL'T (g = u) eiediAo snoijdy (680'0-SS0'0) L900 (801 0-LL0'0) 960°0 (8Z1'0-€60'0) OIT 0 (OTI-0-T80'0) 9010 (S80°0-9¢0'0) 790°0 (€9T 0-881 0) SET O (CEL O-ZIT'O) E710 (6F0°0-6£0'0) S00 (06°0-0L'0) €8°0 (TS 0-OF'0) Lr'0 (TLTO-9F1'0) LST O (LST O-CCI'0) EvI'0 (COT O-£9T'0O) 810 (-71°0-€60'0) LOTO (PST O-ST1O) Trl 0 (LE'0-P0E'0) THEO (S€T0-C6I 0) FOTO (S6 I-81) COI (T] = U) erediata aepy (680°0-6S0'0) FZ0°0 (FI 1°0-6L0'0) 960°0 (OSTO-STI'0) 8ET0 (€T10-£60'0) 8010 (760°0-9S0'0) 9L0°0 (10¢0-S9T'0) F870 (F71 0-700) FIT 0 (6¢0'0-St0'0) LOO (18'0-€L'0) LL'0 (OS O-St'0) Lr'0 (O91 0-091'0) O9T'0 (F91'O-brl 0) ISTO (O61°0-6S 10) SLT-0 (SIT T'O0-160'0) 9010 (8E1 0-810) pel O (€8e0-S9E 0) PLEO NECCUncec 0) Cec CEGOW OC) eric (¢ = U) xXEMepuNny snoijdy (LOT'O-8£0'0) TL0°0 (8E1T°0-690'0) OOT'O (SPl'O-880'0) 6110 (O€ 1 0-600) OTTO (811 0-LS0'0) S80°0 (8Tr'0-9LI'0) 6670 (Stl 0-£90'0) €T1°0 (¢SO0'0-8£0'0) 9F0°0 (LO'I1-9S'0) 6L'0 (69'0-€£'0) 6F'0 (LOT O-CS1'O) 6LT0 (LLT'O-TeT 0) 9ST 0 (9170-1010) TLIO (8E1L'0-880'0) E110 (c6l0-T80'0) 6E 10 (Cer 0-S6I 0) LTEO (OLC O-V1I7 0) Ive 0 (90'€-SZ 1) CET (97 = U) viediaAta snoiady S900 €llO eu Pu £60'0 6c 0 ccl O Lr0'0 L9'0 tv0 O9T'O LST O rS1O 860°0 odMojoy IYSloy spoiaqny s19eK1OYyIO.Ag USUI] spo1aqn} d19RI1OyIOIg aseq 18 yIpIM epnea yysueay] epneD aseq 78 YIPIM snynounydis yisua] snjnounydis I] susieieiapy [P-NUDA ‘T snsie}RIa\ BIGneIOI INWIAJBIII\ A] winnsoy I] winnsoy Ld A Wy aseq A “Uy A] JV I] Wy (s]Jayoos [RUUaJUR UdaaMJaq) SUOI{ (epnes 0} suo1y) Apog eg Apogsydiop ‘adAjojoy ay) pure iosif pupMoy Jo sydiow JAY OF ‘UT UT ‘aSuRI PUR UvdUT ‘s}UdUTAINSeoU [RoTSO;OYydIoyY “{ ATqQRL 32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-9. ment and thoracic furculum of apterous vivipara. 8—9, Metatibia and ultimate rostral segment of ovipara. lTowana frisoni. 7, Ultimate rostral seg- darkening of the knees. Abdomen: Tergum smooth with a few scattered minute hairs, <0.01 mm long. Venter with scattered mi- nute hairs and faint imbrications. Large lat- eral tubercles on abdominal segments I-V and VII. A small lateral tubercle on either, neither, or both sides of segment VI. Sub- genital plate with short hairs profuse only on hind margin, sparse elsewhere. Cauda short, roughly as broad at base as it is long. Three rudimentary gonapophyses. Siphun- culus dark, imbricated, with rounded flange and slight constriction just below distal end (Fig. 15). Collection dates: 8 June, 8 Au- gust 2001, 27 August 2002. Apterous fundatrix (Table 1, Figs. 11, 16).—As apterous vivipara except for fol- lowing. Head: Ant. II with 6—11 rhinaria (Fig. 11). Collection date: 9 May 2002. Alate vivipara (Table 1, Figs. 4, 12, 17).—As apterous vivipara except for fol- lowing. Head: Ant. HI with 12—22 rhinaria (Fig. 12). Thorax: Media once (Fig. 4) or twice forked, if twice, second branch arises right before apex of wing. Abdomen: Si- phunculus parallel-sided and straight, with or without slight flange (Fig. 17). Collec- tion dates: 22 May, 8 June, 8 August 2001. Apterous: male (lable 1, Figs: 35) 13: 18).—As apterous vivipara except for fol- lowing. Head: Rostrum reaches abdominal segment V. Ant. HI with 5—11 rhinaria. Ant. IV with 0—2 secondary rhinaria, 1 primary rhinarium (Fig. 13). Abdomen: Siphunculus only about twice as long as wide, lightly pigmented (Fig. 18). Male genitalia with 2 setose claspers anteriorly, aedeagus central- ly, and 2 sclerotized apophyses posteriorly. Collection date: 6 October 2002. Apterous ovipara (Table 1, Figs. 6, 14, 19).—As apterous vivipara except for fol- lowing. Head: Rostrum reaches abdominal segment V. Ant. HI with 1—9 rhinaria (Fig. 14). Thorax: Metatibia with 10—18 senso- ria. Collection date: 6 October 2002. Egg.—Oval, smooth, shiny and_ black, 0.34—0.36 mm wide by 0.78—0.93 mm long (means of 11 measurements are 0.351 and 0.853 mm, respectively). Collection date: 6 October 2002. Diagnosis.—The viviparae (apterae and alatae) of /. frisoni are similar to Aphis spe- cies in having the same lateral tubercle placement (ventral to the line joining the first two abdominal spiracles), the siphun- culus longer than the cauda, and the pro- cessus terminalis longer than the base of terminal antennal segment. Jowana frisoni is unique with its enlarged lateral protho- VOLUME 106, NUMBER 1 Figs. 10-19. Towana frisoni. 10-14, Antenna. 15—19, Siphunculus. 10, 15, Apterous vivipara. 11, 16, Fun- datrix. 12, 17, Alate vivipara. 13, 18, Male. 14, 19, Ovipara. racic and abdominal tubercles, five-seg- mented antenna, and sparse and minute se- tae. The alate morph is also unusual in hav- ing the medial vein either forked once, or the second fork arising only at the wing apex. ACKNOWLEDGMENTS Our grateful appreciation to D. J. Voeg- tlin (Center for Economic Entomology, Il- linois Natural History Survey) for assis- tance with basic issues of aphid biology, photomicroscopy, and for helpful com- ments on the manuscript. Thanks to S. Cov- er (Museum of Comparative Zoology, Har- vard University) and S. Beshers (Depart- ment of Entomology, University of Illinois at Urbana-Champaign [UIUC]) for identi- fying the two Lasius species; to M. B. Sto- etzel (Systematic Entomology Laboratory, 34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON U.S. Department of Agriculture) for loan of the type specimen of /. frisoni from the Na- tional Aphid Collection, Beltsville, MD, and for a helpful review of the manuscript; and to J. B. Nardi (Department of Ento- mology, UIUC) for use of his photomicros- copy equipment. We appreciate the assis- tance provided by J. A. Mohler, M. W. Tooker, and A. V. Weaver in the field. We also thank the Illinois Nature Preserves Commission, Red Bison (a UIUC Regis- tered Student Organization), the Urbana Park District, and Steve Buck and the Com- mittee of Natural Areas of the School of Integrative Biology, UIUC, for access to re- search sites. LITERATURE CITED Albrecht, G. H. 1980. Multivariate analysis and the study of form, with special reference to canonical variate analysis. American Zoologist 20: 679— 693. Brown, P. A. and R. L. Blackman. 1994. Morphometric variation in the Geoica utricularia (Homoptera: Aphididae) species group on Pistacia (Anacarda- ceae), with descriptions of new species and a key to emigrant alatae. Systematic Entomology 19: 119-132. Carter, C. I. 1983. Some new aphid arrivals to Britain’s forests. Proceedings and Transactions of the Brit- ish Entomological and Natural History Society 16: 81-87. Debaraj, Y. and T. K. Singh. 2000. Morphometric study of different stages of cabbage aphid, Brev- icoryne brassicae (L.) (Homoptera: Aphididae). Russian Entomological Journal 9: 315-319. Dejean, A., B. Nkongmeneck, B. Corbara, and C. Djie- to-Lordonl. 1991. Impact des fourmis arboricoles sur une pullulation d’Achaea catocaloides (Lepi- doptera, Noctuidae) dans des plantations de ca- caoyers du Cameroun, et étude de leurs Homop- teres associés. Acta Oecologica 12: 471—488. Eastop, V. E 1953. A study of the Tramini. Transac- tions of the Royal Entomological Society of Lon- don 104: 385-413. Foottit, R. G. 1992. The use of ordination methods to resolve problems of species discrimination in the genus Cinara Curtis (Homoptera: Aphidoidea: Lachnidae), pp. 193—221. Jn Sorenson, J. T. and R. G. Foottit, eds. Ordination in the Study of Mor- phology, Evolution and Systematics of Insects: Applications and Quantitative Genetic Rationals. Elsevier, Amsterdam. Foottit, R. G. and M. Mackauer. 1980. Morphometric variation within and between populations of the pine aphid, Cinara nigra (Wilson) (Homoptera: Aphidoidea: Lachnidae), in western North Amer- ica. Canadian Journal of Zoology 68: 1410-1419. Hand, S. C. 1986. The use of multivariate morpho- metric methods in the separation of alate morphs of the rose-grain aphid, Metapolophium dirhodum. Annals of Applied Biology 109: 19-31. H6lldobler, B. and E. O. Wilson. 1990. The Ants. Har- vard University Press, Cambridge, Massachusetts. Hottes, KE C. 1954. Description of a new genus and species of Aphidae. Proceedings of the Biological Society of Washington 67: 99-102. Hottes, E C. and T. H. Frison. 1931. The plant lice, or Aphiidae, of Illinois. Bulletin of the Illinois Nat- ural History Survey 19: 121—447. Panzer, R., D. Stillwaugh, R. Gnaedinger, and G. Der- kovitz. 1995. Prevalence of remnant dependence among the prairie- and savanna-inhabiting insects of the Chicago region. Natural Areas Journal 15: 101-116. Pontin, A. J. 1958. A preliminary note on the eating of aphids by ants of the genus Lasius. Entomol- ogists’ Monthly Magazine 95: 154—155. . 1960. Observations of the keeping of aphid eggs by ants of the genus Lasius. Entomologists’ Monthly Magazine 96: 198-199. . 1978. The numbers and distribution of sub- terranean aphids and their exploitation by the ant Lasius flavus (Fabr.). Ecological Entomology 3: 203-207. SPSS. 2000. SYSTAT 10 User’s Guide. Chicago, Illi- nois. Tooker, J. EK and L. M. Hanks. In press. Impact of Prescribed Burning on Endophytic Insect Com- munities of Prairie Perennials (Asteraceae: Si/- phium spp.) Biodiversity and Conservation. Tooker, J. E, W. A. Koenig, and L. M. Hanks. 2002. Altered host plant volatiles are proxies for sex pheromones in the gall wasp Antistrophus rufus. Proceedings of the National Academy of Science 99: 15486-15491. Traniello, J. EF A. and S. C. Levings. 1986. Intra- and inter colony patterns of nest dispersion in the ant Lasius neoniger: Correlations with territoriality and foraging ecology. Oecologia 69: 413—419. Weaver, J. E. 1954. North American Prairie. Johnsen Publishing Co., Lincoln, Nebraska. White, J. 1988. Protection of Pine Ridge Cemetery Prairie: A story of persistence and cooperation. Natural Areas Journal 8: 100—106. Wilson, E. O. 1955. A monographic revision of the ant genus Lasius. Bulletin of the Museum of Com- parative Zoology 113: 1-199. Zwolfer, H. 1958. Zur Systematik, Biologie und Oek- ologie unterirdisch lebender Aphiden. Zeitschrift fiir Andewandte Entomologie 42: 129-172. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 35-51 REVISION OF THE SUBFAMILY CARDIOCHILINAE (HYMENOPTERA: BRACONIDAE) IN CHINA. I. THE GENERA AUSTEROCARDIOCHILES DANGERFIELD, AUSTIN, AND WHITFIELD, EURYCARDIOCHILES DANGERFIELD, AUSTIN, AND WHITFIELD AND PSILOMMISCUS ENDERLEIN XUEXIN CHEN, J. B. WHITFIELD, AND JUNHUA HE (XC, JH) Institute of Applied Entomology, Zhejiang University, Hangzhou 310029, China; (JBW) Department of Entomology, University of Illinois, Urbana, IL 61801, U.S.A. (e-mail: jwhitfie @life.uiuc.edu) Abstract.—An illustrated key to the six genera of the Cardiochilinae known to occur in China is presented, with three genera, Austerocardiochiles Dangerfield, Austin, and Whitfield, Eurycardiochiles Dangerfield, Austin, and Whitfield and Psilommiscus Ender- lein, reported for the first time. Three new species of Austerocardiochiles are described: A. tujiazu Chen, Whitfield and He, A. xibozu Chen, Whitfield and He, and A. zhe- jJiangensis Chen, Whitfield and He. Four species of Eurycardiochiles are reported: E. occidentalis Dangerfield and Austin, and the new species FE. dongzu Chen, Whitfield and He, E. shezu Chen, Whitfield and He, and E. jiulong Chen, Whitfield and He. Psilommiscus 1s new to China, represented only by P. sumatranus Enderlein. Austero- cardiochiles japonicus (Watanabe), new combination, and A. turga (Belokobylskij), new combination, are transferred from Cardiochiles Nees. Key Words: Braconidae, Cardiochilinae, Austerocardiochiles, Eurycardiochiles, Psilom- miscus, new species, new records, China The subfamily Cardiochilinae is a small, well-established subfamily in the family Braconidae (Quicke and van Achterberg 1990, Whitfield and Mason 1994, Whitfield and Dangerfield 1997, Dangerfield et al. 1999, Mercado and Wharton 2003). The knowledge of its biology is mostly restrict- ed to a few commonly studied species of Cardiochiles Nees s.s. and Toxoneuron Say, particularly 7. nigriceps (Viereck). Cardi- ochilinae are endoparasitoids of lepidopter- an larvae of the families Apatelodidae, Cos- mopterigidae, Gelechiiidae, Pyralidae, Noc- tuidae and Uraniidae, some of which are major pests of agricultural crops and forests (Huddleston and Walker 1988). The species appear to be most diverse and abundant in temperate grasslands and tropical forests, especially seasonally dry forests. The Cardiochilinae contain 16 extant genera worldwide according to the new ge- neric Classification of this subfamily based on a phylogenetic analysis of qualitative and morphometric characters carried out by Dangerfield et al. (1999). The subfamily is cosmopolitan in distribution, while individ- ual genera are sometimes restricted to in- dividual hemispheres or continents (Table 1). Several of the sixteen genera have been redefined recently (Mercado and Wharton 2003), and it is likely that the generic total will continue to change for some time. Dol- phin and Quicke (2001) have estimated a total diversity of from 187 to 283 world PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 36 ON C661 NOYD -906[ “Ule[tepuy ON Jaded sty, ON ON ON ON (8661) BIN Pur 2H “UeYD ‘0661 “UNsNYy 2 playsesueqd ON ON Jaded styy C661 “NoYyD ‘QO61 “Ula[IapUY :ZO6T “Hesydezg ‘Cog “Aefnoyoy ON Jaded styy ON suOIsay [RoIdOMOANY ‘oDIvANY ‘[eotdo.noipy uvypodowsop UOIBdY ONOIe[OH, UOISAY [PIUILIO SUOISAY URITRISNY ‘oNdIvAR[ed UuOIsoY UPITeNSNYy UOIsoy UPITR.QSNY uOISayY [RoIdO.NOADN UOIBAY [VIUIIO uoIsey [eordo.noaN UOIsOyY UPITeNSNY SUOISOY IjoIvaL[e ‘[eIUIUIOC, uevyyodouwsog jeotdomnoay 1da0xe suOIseal [TV uevlyensny ‘yeordonoiyy ‘[eUsUIO ‘onoI1eae[ed SUOISOy ULITeNSNY ‘oNoIvaRled ARG UOANIUOXO J, UOIOWILD VjJapubjua2oYyIS ‘PIA pue “jsny “sued /alspsisnjay Uldp1apuy SNIsMUUOpIS J SIMPoH{ SN/1YIOIpAVIPNAS “PINAA pue “sny “Ssuegq wnssajjvoT ‘pPIUM pue “jsny “sued siimawdy g[nig uosajdosalayy UOIOWD VIMNMALID Pleysosueq viuosuvy ‘IY AA pue “ysny “Sued vIUuanyH ‘PUY pure ‘|sny “sued saj1yIo01IpsAVINAN| SOON Sa71YyIOIpPADD frys[Aqoyojeg Yjjatpyog ‘HIYA pue “ysny “sued sap1yoo1psvz0s1a1sSny BSUDA], SAIYIOIPADIVISY BUIYD WIOIy pop10zay uonnqmsiq uMouy snuen ‘BUIYD Ul ddUaLIND9O Ilay] puke SUONNQLNSIp UMOUY IY) YUM IBUT[IYSOIPIeD JO PIBUDS PLIOM “T IQVL VOLUME 106, NUMBER 1 cardiochiline species, using different esti- mation methods. There are just under 200 described species at present (Dangerfield et al. 1999, Mercado and Wharton 2003); however, there are large numbers of unde- scribed species, particularly from south- eastern Asia, Africa, and Central and South America where the subfamily has not been studied comprehensively. For example, Mercado and Wharton (2003) recognize 45 species from Mexico where only 28 are cur- rently described, even after their study. Thus the Dolphin and Quicke (2001) esti- mates are likely somewhat low. The Chinese fauna of Cardicochilinae is among the most sparsely studied in the world, only the species of Hartemita Cam- eron having been revised (Chen et al. 1998). This paper is the first to more broad- ly revise the Chinese Cardiochilinae. Pre- viously, three genera, i.e., Cardiochiles Nees s. str., Hartemita Cameron, and Schoenlandella Cameron were recorded from China, but revision of the large genera Cardiochiles s.s. and Schoenlandella Cam- eron from China still remains. Austerocar- diochiles Dangerfield, Austin, and Whit- field, Eurycardiochiles Dangerfield, Austin, and Whitfield and Psilommiscus Enderlein are reported and described here for the first time from China. Based on their presently recorded distributions, Asiacardiochiles Te- lenga, Bohayella Belolobylskij and Pseud- cardiochilus Hedwig are also likely to oc- cur in China, but we have not seen Chinese specimens of them. The number of specimens available for the taxa treated here is low. XC and JH have examined the following collections: Zoological Research Institute (Beijing); En- tomological Research Institute (Shanghai) of the Chinese Academy of Sciences; Fu- jian Agricultural University (Fuzhou); Shaanxi Agricultural University (Xi'an, one of the largest insect collections in China); China Agricultural University (Beijing); Nanjing Agricultural University (Nanjiang); Huanan Agricultural University (Guang- zhou); Guizhou University (Guiyang); and 37 Zhejiang University (the largest parasitic wasp collection in China). They found about 2,200 specimens of Cardiochilinae, the vast majority of them belonging to Car- diochiles, Schoenlandella, and Hartemita. For example, as many as 125 specimens of one species, Cardiochiles fuscipennis Sze- pligeti, were found in the Zhejiang Univer- sity collection. Thus, the small numbers of specimens of the taxa treated here indicate that these genera and their species are not very abundant in China, at least compared to the three large genera, rather than that almost no material of Cardiochilinae has been sampled. For the identification of extralimital gen- era of Cardiochilinae and most of the mor- phological terminology used in this paper, see Dangerfield et al. (1999). POL:OD: OOL refers to the three-way ratio of the distance between the inner edges of the pos- terior ocelli (POL) to the diameter of a pos- terior ocellus (OD) and to the distance be- tween the outer edge of a posterior ocellus and its corresponding compound eye (OOL). Wing vein terminology follows Achterberg (1993). Voucher and type spec- imens are deposited in the Hymenoptera Collection, Zhejiang University, Hangzhou, China. KEY TO THE CHINESE GENERA OF THE SUBFAMILY CARDIOCHILINAE 1. Ovipositor and sheaths very short, less than 0.2 times hind tibia, stout and sharply down- turned; propodeal areola reduced anteriorly; hind basitarsus laminate Hartemita Cameron — Ovipositor and sheaths much longer, not as stout, only weakly down-turned to straight at apex; propodeal areola complete; hind basitar- sus rounded or oval in cross-section (some- times distinctly compressed but never strongly Laminate) © of 3 ok toe se Oe. eee cee re 2, . Tl and latero-tergites with lateral suture clearly defined throughout; hypopygium pointed at apex with medial longitudinal area descleroti- zZedvand foldedsimwatdsyve sen oe ee . Austerocardiochiles Dangerfield, Austin, and Whitfield — T1 with lateral suture reduced, poorly defined particularly in apical half; hypopygium vari- able i) 38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 3. 3r in fore wing virtually always present as a spectral trace, if apparently absent then mouth- parts elongate; glossa usually moderately elon- gate and deeply bilobed at apex; galea mostly long, narrow and bladelike YS he Fan eee Schoenlandella Cameron — 3r in fore wing absent; glossa variable; galea never bladelike, usually short and, if moderate- lvelonicesthenibtoad ae ees cee 4 4. Hypopygium with medial longitudinal descler- otized and membranous area; galea long and broad, most clearly visible past mandible (glos- sa sometimes moderately long) Prey OSS Cae Say Os fey Cardiochiles Nees — Hypopygium evenly sclerotized throughout, sometimes partially desclerotized and folded inwards, but never membranous; galea short and either narrow or broad 5. Eye setae reduced to minute interommatidial spines; scutellum with apical cuplike pit .... eR i ec ere Psilommiscus Enderlein — Eye with conspicuous pilosity; apex of scutel- hime wath outc mp like ipiter ens ieee ee Eurycardiochiles Dangerfield, Austin, and Whit- field Austerocardiochiles Dangerfield, Austin, and Whitfield, 1999 Austerocardiochiles Dangerfield, Austin, and Whitfield 1999: 929. Type species (by original designation): Cardiochiles pollinator Dangerfield and Austin. The genus was proposed by Dangerfield et al. (1999) to include eight described spe- cies. Two additional described species, Car- diochiles japonicus (Watanabe), n. comb. (Japan) and C. turga (Belokobylskij), n. comb. (Russian Far East) are transferred into this genus (following Belokobylskij, personal communication). The number of species in this genus accounts for about 5% of the described cardiochiline species. It is widely distributed in the Old World, being recorded from Australia, Philippines, Laos, Japan, Russia, Africa, and now China, but most species occur in the Southern Hemi- sphere. The species known from Philip- pines and Laos are currently undescribed. The biology of this genus is unknown, but A. enderleini (Szepligeti) has been collected from crops of sweet potato (Jpomaea sp.) in Kenya. Austerocardiochiles species have the oc- cipital carina present in the postgenal re- gion, a longitudinal carina set in a furrow on the medial scutum, and epicnemial ca- rina developed (except for A. tujiazu, n. sp.), while the medial hypopygium shows variable stages of longitudinal descleroti- zation among species. Austerocardiochiles is, in the analyses of Dangerfield et al. (1999), the sister group to Cardiochiles s. str. on the basis of the putative synapo- morphy of the hypopygium being medially desclerotized (see Austin 1990 and Danger- field et al. 1999 for a discussion of the po- larity of this feature). It also has the lateral sutures of Tl defined and percurrent, a character also found, apparently indepen- dently, in Hansonia and Heteropteron (Dangerfield et al. 1999). KEY TO THE CHINESE SPECIES OF AUSTEROCARDIOCHILES 1. T1 short, distinctly widened apically, 0.9 times as long as broad (Fig. 3); T2 punctate; wing membrane largely distinctly brown; clypeal tu- bercles obsolescent (Fig. 2); epicnemial carina absent; posterior cuplike pit of scutellum ob- vious; scutellar sulcus wide, 3.2 times as broad as long; hind basitarsus almost cylindrical; Rs bent in basal 0.20, straight to subapex, then curved downwards to apex (Fig. 1); hind wing with six hamuli; body large, approx. 9-10 mm 5 ES Cie hades, Eyl roe tee, heel CL ene A. tujiazu n. sp. — T1 longer, hardly widened apically, 1.4—2.0 times longer than broad (Figs. 5, 8); T2 smooth; wing membrane distinctly brown only at apical 4; clypeal tubercles distinct (Figs. 6, 9); epicnemial carina present; posterior cup- like pit of scutellum absent; scutellar sulcus narrow, 4.4—4.5 times as broad as long; hind basitarsus distinctly compressed; Rs bent in basal 0.24—0.25, then straight to apex (Figs. 4, 7); hind wing with five hamuli; body smaller, approx 5—7 mm . Rs of fore wing evenly curved basally (Fig. 4); discal cell of fore wing 2.1 times longer than wide; T1! 2.0 times longer than broad, bulb slightly convex and less rugose (Fig. 5); middle lobe of T1 with a fine longitudinal carina in its longitudinal groove .......... A. xibozu, N. sp. — Rs of fore wing angularly bent basally (Fig. 7): discal cell of fore wing 1.7 times longer than wide; T1 1.4 times longer than broad, bulb dis- tinctly convex and distinctly rugose (Fig. 6); i) VOLUME 106, NUMBER 1 Figs. 1-3. middle lobe of Tl with a more prominent lon- gitudinal carina in its longitudinal groove; Rite: RG din tee OR oe OLA A. zhejiangensis, n. sp. Austerocardiochiles tujiazu Chen, Whitfield, and He, new species (Figs. 1-3) Female.—Body length 9.8 mm, fore wing length 8.8 mm. Head: Eyes with mod- erately dense fine pilosity; head distinctly sculptured with dense long pilosity; labio- maxillary complex just visible below man- dible; malar suture present; epistomal su- ture reduced but visible; clypeal margin convex laterally, with straight medio-lateral part and convex medial part with two ob- solescent tubercles; clypeus convex and dis- tinctly rugo-punctate, broad, 2.6 times as broad as high; face 2.0 times as broad as high, distinctly convex, transversely rugose, with medial dorsal node and irregular lon- Austerocardiochiles tujiazu. 1, Wings. 2, Head, dorsal view. 3, First to third metasomal tergites. gitudinal carina indicated; frons distinctly concave, transversely carinate, with smooth medial carina; length of eye equal to length of temple in dorsal view; POL:OD:OOL = 7:6:21; vertex not raised behind ocelli, nar- row, transversely rugose; depth of occiput in dorsal view 0.37 times length of head; antenna with 41(+) segments, short, robust, scape 1.4 times as long as broad; distance between antennal sockets equal to width of antennal socket. Mesosoma: Side of pronotum distinctly longitudinally rugo-striate ventrally, rest rugo-striate, dorso-medial longitudinal fur- row crenulate; notauli present, distinctly crenulate, of even width, meeting posteri- orly; scutum distinctly punctate, middle lobe with a fine longitudinal carina in its longitudinal groove; parapsidal grooves present; scutellum sparsely pilose, coarsely 40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rugose, 1.3 times as broad as long, lateral carinae distinct, posterior cuplike pit obvi- ous; scutellar sulcus wide, 3.2 times as broad as long, with four carinae; dorsellum with medial longitudinal carina weakly de- fined; propodeal areola lens-shaped, 0.51 times as broad as long, propodeal spiracles elliptical, 2.2 times as long as broad; pro- podeum short, distinctly rugose; epicnemial carina absent; sternaulus distinct, broad, an- teriorly crenulate, posteroirly nearly smooth; mesopleuron glabrous medially above sternaulus, below it foveate-rugose; metapleuron coarsely rugose. Legs: Fore tibial spur 0.85 times as long as basitarsus; hind tibia expanded slightly at apex, 3.0 times as wide as at base; hind tibia and tarsus with brown spines among pilosity; inner hind tibial spur 1.4 times as long as outer spur, 0.7 times as long as bas- itarsus; hind basitarsus almost cylindrical with flattened dorsal ridge at base; tarsal claws pectinate, hind claw with six teeth. Wings: Fore wing pilosity sparse at base, becoming thicker towards apex; stigma 4.0 times as long as wide; Rs bent in basal 0.2, straight to subapex, then curved downwards to apex; 1-SR+M 3.3 times as long as m- cu; 2-SR+M _ 1.7 times as long as m-cu; 1- M straight; m-cu 0.4 times as long as 1-M; 1-M 0.74 times as long as 1-SR+M;; apical % of 2-M and 3-M spectral; discal cell 1.5 times as long as wide; 2cu-a represented by pigmented area; la present, spectral. in whole length; 1-2A+3A spectral, reaching to la; 1-CUa 0.27 times as long as 1-CUb; 2r 0.54 times inside height of second sub- marginal cell, arising 0.6 times length along stigma; second submarginal cell 2.4 times as long as wide; 2+3-SR slightly arched near 2-M; hind wing with six evenly spaced hamuli; 2-1A and 2A absent; 2-SC+R ver- tical; r present, spectral. Figs. 4-9. Metasoma: T1 short, distinctly widened apically, 0.9 times as long as broad, stem 0.3 times as long as bulb; bulb convex but medio-longitudinally concave, distinctly coarsely rugose; suture between T2 and T3 represented by well-defined groove; median field of T2 0.35 times as long as wide; T2 punctate; ovipositor sheaths broad and long, 0.55 times as long as hind tibia, 0.27 times as broad as long, densely hairy along length, broadening to and rounded at apex with dorsal notch; hypopygium with sparse fine hairs, hairless at apex, rounded and acutely angled at apex, largely sclerotised except medially slightly desclerotized with sclerotized bridge apically and basally. Color: Black, fore tibia, tarsus and apex of fore femur orange; wing membrane dis- tinctly brown, basally paler, stigma and veins dark brown. Male.—Unknown. Material examined.—Holotype °, China: Hubei, Shiniandachuan, 1986.vili.2, Mao Xiaoyuan, no.870217. Note.—This species is similar to A. xib- ozu and A. zhejiangensis, but can be sepa- rated from them by the characters listed in the key. From the type species, this species (as do the other two Chinese species below) has a broader and more apically rounded first metasomal tergite. Etymology.—This species is named after one of the Chinese minorities, Tujia Zu, who live mainly in Hubei and Hunan prov- inces. Austerocardiochiles xibozu Chen, Whitfield, and He, new species (Figs. 4—6) Female.—Body length 5.6 mm, fore wing length 5.4 mm. Head: Eyes with mod- erately dense fine pilosity; head distinctly sculptured with dense long pilosity; labio- > 4—6, Austerocardiochiles xibozu. 4, Fore wing. 5, First to third metasomal tergites, dorsal view. 6, Head, dorsal view. 7-9, A. zhejiangensis. 7, Fore wing. 8, First to third metasomal tergites, dorsal view. 9, Head, dorsal view. Al VOLUME 106, NUMBER 1 42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON maxillary complex just visible below man- dible; malar suture present; epistomal su- ture reduced medially; clypeal margin con- vex laterally, with straight medio-lateral part and convex medial part with two dis- tinct tubercles; clypeus convex and distinct- ly rugo-punctate, broad, 2.1 times as broad as high; face 1.7 times as broad as high, distinctly convex, transversely rugose, with an obvious longitudinal carina; frons dis- tinctly concave, transversely finely rugose, with a medial carina; length of eye 1.3 times as long as length of temple in dorsal view; POL:OD:OOL = 9:6:17; vertex not raised behind ocelli, narrow, transversely rugose; depth of occiput in dorsal view 0.52 times length of head; antenna with 39 seg- ments, short, robust, scape distinctly punc- tate, 1.8 times as long as broad; distance between antennal sockets about equal to width of antennal socket. Mesosoma: Side of pronotum coarsely longitudinally rugo-striate ventrally, rest rugo-striate, dorso-medial longitudinal fur- row crenulate anteriorly; notauli present, distinctly crenulate, of even width, meeting posteriorly; scutum distinctly rugose, mid- dle lobe with a fine longitudinal carina in its longitudinal groove; parapsidal grooves present; scutellum sparsely pilose, coarsely rugose, 1.5 times as broad as long, lateral carinae distinct, without posterior cuplike pit; scutellar sulcus narrow, 4.4 times as broad as long, with four strong carinae; pro- podeal areola lens-shaped, 0.55 times as broad as long, propodeal spiracles elliptical, 2.5 times as long as broad; propodeum short, distinctly rugose; epicnemial carina present, distinctly ventrally; sternaulus dis- tinct, broad, anteriorly crenulate, posteroirly nearly smooth; mesopleuron glabrous me- dially above sternaulus, below sternaulus striate-rugose; metapleuron coarsely ru- gose. Legs: Fore tibial spur 0.9 times as long as basitarsus; hind tibia expanded slightly at apex, 3.0 times as wide as at base; hind tibia and tarsus with brown spines among pilosity; inner hind tibial spur 1.6 times as long as outer spur, 0.66 times as long as basitarsus; hind basitarsus compressed with flattened dorsal ridge at base; tarsal claws pectinate, hind claw with seven teeth. Wings: Fore wing pilosity sparse at base, becoming thicker towards apex; stigma 3.4 times as long as wide; Rs bent in basal 0.25, straight to near apex, then slightly curved downwards to apex; 1-SR+M 3.6 times as long as m-cu; 2-SR+M _ 1.5 times as long as m-cu; 1-M straight; m-cu 0.4 times as long as 1-M; I-M 0.7 times as long as 1I- SR+M; apical %4 of 2-M and 3-M spectral; discal cell 2.1 times as long as wide; 2cu-a represented by pigmented area; la present, spectral in whole length; 1-2A+3A _ spec- tral, reaching la; 1-CUa 0.32 times as long as 1-CUb; 2r 0.67 times inside height of second submarginal cell, arising 0.6 times length along stigma; second submarginal cell 2.6 times as long as wide; 2+3-SR slightly arched near 2-M; hind wing with five evenly spaced hamuli; 2-1A and 2A absent; 2-SC+R vertical; r present, spectral. Metasoma: T1 long, hardly widened api- cally, 2.0 times as long as broad, stem about 0.33 times as long as bulb; tergite beside stem finely striate, bulb slightly convex, in- distinctly rugose, but smooth apically; su- ture between T2 and T3 represented by well-defined groove; median field of T2 0.31 times as long as wide; T2 smooth; ovi- positor sheath broad and long, 0.52 times as long as hind tibia, 0.29 times as broad as long, densely hairy along length, broad- ening to and rounded at apex with dorsal notch; hypopygium with sparse fine hairs, hairless at apex, rounded and acutely angled at apex, largely sclerotized except medially slightly desclerotized and folded inwards. Color: Black; antenna and palpi dark brown; fore tibia, tarsus and apex of fore femur orange; tibia and basitarsus of middle leg brown, 2nd—Sth tarsi of middle leg brownish yellow, middle and hind tibiae with a yellowish white basal ring; latero- tergite of Tl dark reddish brown; wing membrane only brownish on apical 4%, rest VOLUME 106, NUMBER 1 hyaline, stigma dark brown, veins brown to dark brown. Male.—Unknown. Material examined.—Holotype °, China: Jilin, Antu, 1994.viii.5—6, Lou Juxian, no.977083. Note.—This species is closely similar to A. rugosus (Telenga) but can be separated from the latter by T1 longer, 2 times as long as wide, rugose, but smooth apically; eyes with moderately dense and fine pilosity; POL 1.5 times OD; and latero-tergite of first tergite ventrally dark reddish brown. Etymology.—This species is named after one of the Chinese minorities, Xibo Zu, who live mainly in Jilin and Liaoning prov- inces. Austerocardiochiles zhejiangensis Chen, Whitfield, and He, new species (Figs. 7-9) Female.—Body length 6.5 mm, fore wing length 6.1 mm. Head: Eyes with dense fine pilosity; head distinctly sculp- tured with dense long pilosity; labio-max- illary complex just visible below mandible; malar suture present; epistomal suture vis- ible; clypeal margin convex laterally, with straight medio-lateral part and convex me- dial part with two tubercles; clypeus convex and distinctly rugose, broad, 2.1 times as broad as high; face 1.7 times as broad as high, distinctly convex, coarsely rugose, with strong but fine medio-longitudinal ca- rina; frons distinctly concave, transversely carinate, with smooth medial carina; length of eye 1.3 times as long as temple in dorsal view; POL:OD:OOL = 10:6.5:18; vertex slightly raised behind ocelli, narrow, coarsely rugose; depth of occiput in dorsal view 0.6 times length of head; antenna with 41 segments, short, robust, sacape 1.7 times as long as broad; distance between antennal sockets equal to width of antennal socket. Mesosoma: Side of pronotum coarsely longitudinally striate ventrally, dorso-me- dial longitudinal furrow coarsely crenulate; notauli present, distinctly crenulate, of even width, meeting posteriorly; scutum distinct- 43 ly foveate-rugose, middle lobe with a dis- tinct longitudinal carina in its longitudinal groove; parapsidal grooves present; scutel- lum moderately densely pilose, coarsely ru- gose, 1.3 times as broad as long, without posterior cuplike pit; scutellar sulcus nar- row, 4.5 times as broad as long, with two strong carinae; dorsellum without medial longitudinal carina; propodeal areola lens- shaped, 0.57 times as broad as long, pro- podeal spiracles elliptical, 2.3 times as long as broad; propodeum short, distinctly ru- gose; epicnemial carina present, distinctly ventrally; sternaulus distinct, broad, anteri- orly crenulate, posteroirly nearly smooth; mesopleuron glabrous medially above ster- naulus, below it foveate-rugose; metapleu- ron coarsely rugose. Legs: Fore tibial spur 0.9 times as long as basitarsus; hind tibia expanded at apex, 3.1 times as wide as at base, with a longi- tudinal groove on its outside at apical %; hind tibia and tarsus with brown spines among pilosity; inner hind tibial spur 1.4 times as long as outer spur, 0.7 times as long as basitarsus; hind basitarsus com- pressed with flattened dorsal ridge at base; tarsal claws pectinate, hind claw with seven teeth. Wings: Fore wing pilosity sparse at base, becoming thicker towards apex; stigma 3.7 times as long as wide; Rs angularly bent in basal 0.24, straight to apex; 1-SR+M 4.0 times as long as m-cu; 2-SR+M _ 1.7 times as long as m-cu; 1-M straight; m-cu 0.4 times as long as 1-M; 1-M 0.63 times as long as 1-SR+M;; apical %4 of 2-M and 3- M spectral; discal cell 1.7 times as long as wide; 2cu-a represented by pigmented area; la present, spectral in whole length; 1I- 2A+3A spectral, reaching la; 1-CUa 0.37 times as long as 1-CUb; 2r 0.64 times in- side height of second submarginal cell, aris- ing 0.68 times length along stigma; second submarginal cell 2.5 times as long as wide; 2+3-SR_ slightly arched near 2-M; hind wing with five evenly spaced hamuli; 2-1A and 2A absent; 2-SC+R vertical; r present, spectral. 44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Metasoma: T\ long, hardly widened api- cally, 1.4 times as long as broad, stem 0.28 times as long as bulb; tergite beside stem longitudinally striate; bulb distinctly con- vex but medio-longitudinally concave, dis- tinctly coarsely rugose, apically smooth; su- ture between T2 and T3 represented by well-defined groove; median field of T2 0.33 times as long as wide; T2 smooth; ovi- positor sheath broad and long, 0.56 times as long as hind tibia, 0.28 times as broad as long, densely hairy along length, broad- ening to and rounded at apex with dorsal notch; hypopygium with sparse fine hairs, hairless at apex, rounded and acutely angled at apex, largely sclerotized except medially slightly desclerotized and folded inwards. Color: Black; antenna and palpi dark brown; fore tibia, tarsus and apex of fore femur orange, middle tarsus brown to dark brown, middle and hind tibiae with a yel- lowish white basal ring; latero-tergite of T1 dark reddish brown; wing membrane dis- tinctly brown at apical %, basally hyaline, stigma and veins dark brown. Variation.—Body length 6.5—7.0 mm, fore wing length 6.1—6.5 mm. Bulb of first tergite convex but medio-longitudinally not concave. Male.—Unknown. Material examined.—Holotype @, China: Zhejiang, Anjie, Mt. Longwang, 1995.vii.21, Wu Hong, no.970862. Paratype: 1 2, China: Zhejiang, Mt. W. Tianmu, 1987.ix.4, Fan Jinjiang, No.876253. Note.—This species is similar to A. ru- gosus (Telenga), but can be separated from the latter species by having the Rs of fore wing angularly bent in basal 0.24, then straight to apex; bulb of first tergite convex, distinctly coarsely rugose, apically smooth, and sometimes medio-longitudinally con- cave; and POL longer than OD. It is also similar to A. xibozu n. sp., but can be dis- tinguished from the latter by the characters listed in the key. Etymology.—This species is named after Zhejiang province in China, on the east bor- der of the Palaearctic and Oriental regions, where this new species occurs. Eurycardiochiles Dangerfield, Austin, and Whitfield, 1999 Eurycardiochiles Dangerfield, Austin, and Whitfield 1999: 938. Type species (by original designation): Cardiochiles occi- dentalis Dangerfield and Austin. This genus can be recognized by having an apically strongly broadened ovipositor sheath and a broadened clypeus. It is the sister group to Schoenlandella + Austero- cardiochiles + Cardiochiles s. str., all of which have antennae of more than 33 seg- ments, and two apical clypeal tubercles. It is known previously from Australia with the type species and Sri Lanka with an un- described species. The biology of the genus is unknown. This genus is recorded here for the first time in China and three new species are de- scribed below. The Chinese species de- scribed here are quite different from the type species in general appearance. The type species has the head and mesosoma largely smooth, and the hypopygium evenly sclerotized while all Chinese species have the head and mesosoma largely sculptured and the hypopygium medio-longitudinally less strongly sclerotized and often folded inwards. For the moment the Chinese spe- cies appear to fit best within Eurycardi- ochiles, but eventually may be found to rep- resent a distinct group. KEY TO SPECIES OF EURYCARDIOCHILES 1. Head and mesosoma largely smooth; median field of T2 0.5 times as long as wide; hypo- py siunmpevenilyasclerotize diy nee eee E. occidentalis Dangerfield and Austin — Head and mesosoma largely sculptured; me- dian field of T2 0.28—0.32 times as long as wide; hypopygium medio-longitudinally less sclerotized and folded inwards ........... 2 . Body smaller, about 5.5 mm; face 1.9 times as broad as high; vein 1-CUa of fore wing 0.33 times vein 1-CUb; first metasomal tergite lon- ger, 1.2 times as long as broad; antenna with BVESESIMENIS es mae cee E. dongzu, n. sp. — Body larger, about 7.8—8.1 mm; face 1.5 times i) VOLUME 106, NUMBER 1 as broad as high; vein 1-CUa of fore wing 0.42—0.46 times vein 1-CUb; first metasomal tergite short, as long as broad; antenna with ANIAGYSEPMMENES 2 seeleina, is Gr Sane = eel a 3 3. Clypeal margin medially with two distinct tu- bercles; face with a distinct medio-longitudinal carina; wing membrane entirely evenly brown- ish; vein m-cu of fore wing 0.4 times vein 1- M; hind claw with eight teeth; hind wing with eight hamuli; and antenna with 46 segments RPMs) ae Ss Ge ENS a ceeee t E. shezu, n. sp. — Clypeal tubercles indistinct; medio-longitudi- nal carina on face much less distinct; wing membrane much less brownish basally; vein m-cu of fore wing 0.5 times vein 1-M; hind claw with nine teeth, hind wing with six ham- uli; and antenna with 41 segments E. jiulong, n. sp. Eurycardiochiles dongzu Chen, Whitfield, and He, new species (Figs. 10-12) Female.—Body length 5.5 mm, wing length 5.3 mm. Head: Eyes with dense fine pilosity; head sculptured with moderately dense long pilosity; labio-max- illary complex just visible below mandible; malar suture present; epistomal suture vis- ible; clypeal margin convex laterally, with straight medio-lateral part and convex me- dial part with two distinct tubercles (Fig. 12); clypeus broad, convex and distinctly rugose, 2.2 times as broad as high; face 1.9 times as broad as high, coarsely transverse- ly rugose, with a weak medio-longitudinal carina; frons transversely striate, with a me- dial carina reaching near anterior ocellus; vertex not raised behind ocelli, narrow, coarsely transversely striate; depth of occi- put in dorsal view 0.53 times length of head; length of eye 1.4 times length of tem- ple in dorsal view; POL:OD:OOL = 9:6: 15; antenna with 37 segments, short, thick, scape 1.7 times as long as broad; antennal sockets separated by 0.83 times width of antennal socket. Mesosoma: Side of pronotum coarsely longitudinally striate ventrally, dorso-me- dial longitudinal furrow coarsely crenulate, rest striate; notauli present, deep, crenualte, of even width, meeting posteriorly; scutum distinctly rugose, a medio-longitudinal ca- fore 45 rina present in medio-longitudinal groove; parapsidal grooves present; scutellum with dense long pilosity, coarsely rugose, 1.4 times as broad as long, lateral carinae dis- tinct; scutellar sulcus very narrow, 5.1 times as broad as long, with four carinae; dorsellum with a weak medial longitudinal carina; propodeum short, rugose, with cren- ulae branching from areola; propodeal are- ola lens-shaped, 0.57 times as broad as long, propodeal spiracles elliptical, 2.2 times as long as broad; epicnemial carina present; sternaulus broad, coarsely crenu- late; mesopleuron smooth and shiny above sternaulus, below sternaulus distinctly retic- ulate-rugose. Legs: Fore tibial spur as long as basitar- sus; hind tibia expanded at apex, 2.6 times as wide as at base; hind tibia and tarsus with brown spines among pilosity; inner hind tibial spur 1.6 times as long as outer spur, 0.62 times as long as basitarsus; hind basitarsus moderately compressed with flat- tened dorsal ridge at base; tarsal claws pec- tinate, hind claw with six teeth. Wings: Fore wing (Fig. 10) with dense pilosity, much thicker towards apex; stigma 3.2 times as long as wide; 2r 0.43 times inside height of second submarginal cell, arising 0.62 times length along stigma; Rs bent at basal 0.20, then nearly straight to apex; second submarginal cell 2.4 times as long as wide; 2+3-SR slightly arched near 2-M; 1-SR+M 3.7 times as long as m-cu; 2-SR+M 1.8 times as long as m-cu; 1-M straight; m-cu 0.39 times as long as 1-M; 1-M 0.70 times as long as 1-SR+M;; discal cell 1.55 times as long as wide; 2cu-a pre- sented by pigmented area; la present, spec- tral in apical two-thirds; 1-2A+3A spectral, reaching to la; 1-CUa 0.33 times as long as 1-CUb; hind wing with five evenly spaced hamuli; 2-1A and 2A absent; 1-SRa present but short. Metasoma: T1 widened apically, 1.2 times as long as broad, stem 0.26 times as long as bulb, tergite beside stem nearly smooth, bulb convex and smooth (Fig. 11); suture between T2 and T3 represented by 46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 10-15. 10-12, Eurycardiochiles dongzu. 1, Fore wing. 2, First to third metasomal tergites, dorsal view. 3, Head, dorsal view. 13-15, E. jiulong. 13, Fore wing. 14, First to third metasomal tergites, dorsal view. 15, Head, dorsal view. VOLUME 106, NUMBER 1 well-defined groove; median field of T2 0.28 times as long as wide; T2 smooth; ovi- positor sheaths broad and long, 0.56 times as long as hind tibia, 0.28 times as broad as long, densely hairy along length, broad- ening to and rounded at apex with dorsal notch; hypopygium with moderately dense hair laterally, hairless at apex, rounded and acutely angled at apex, evenly sclerotized except for medio-longitudinally less scler- otized and folded inwards. Color: Black; fore tibia, tarsus and apex of femur, middle tarsus and basal 0.2 of middle tibia, basal 0.2 of hind tibia brown- ish yellow to orange, first tergite except for stem and bulb dark red; wing membrane brown, paler basally, stigma and veins brown to dark brown. Male.—Unknown. Material examined.—Holotype ° , China: Hunan, Liuyang, 1984.vii.31, collector un- known, no.846373. Note.—This new species can be easily separated from the type species, E. occiden- talis by the largely sculptured head and me- sosoma; hypopygium medio-longitudinally less sclerotized and folded inwards; median field of T2 more transverse, 0.28 times as long as wide; and hind claw with six teeth. Etymology.—This species is named after one of the Chinese minorities, Dong Zu, who mainly live in Hunan and Guizhou provinces, and in the Guangxi Autonomous Administration Region. Eurycardiochiles jiulong Chen, Whitfield, and He, new species (Figs. 13-15) Female.—Body length 7.8 mm, fore wing length 7.8 mm. Head: Eyes with dense fine pilosity; head sculptured with moderately dense long pilosity; labio-max- illary complex just visible below mandible; malar suture present; epistomal suture vis- ible; clypeal margin convex laterally, then evenly convex medially with two very faint tubercles (Fig. 15); clypeus broad, convex and distinctly rugose, 2.2 times as broad as high; face 1.5 times as broad as _ high, 47 coarsely transversely rugose, with a fine medio-longitudinal carina; frons transverse- ly striate, with a medial carina reaching to anterior ocellus; vertex not raised behind ocelli, narrow, coarsely transversely striate; depth of occiput in dorsal view 0.58 times length of head; length of eye 1.3 times length of temple in dorsal view; POL:OD: OOL = 10:6.5:21; antenna with 41 seg- ments, short, thick, scape 2.0 times as long as broad; antennal sockets separated by 0.9 times width of antennal socket. Mesosoma: Side of pronotum coarsely longitudinally striate ventrally, dorso-me- dial longitudinal furrow coarsely rugose, rest striate; notauli present, deep, crenulate, of even width, meeting posteriorly; scutum distinctly rugose, a medio-longitudinal ca- rina present in medio-longitudinal groove; parapsidal grooves present; scutellum with dense long pilosity, coarsely rugose, 1.3 times as broad as long, lateral carinae dis- tinct; scutellar sulcus narrow, 3.6 times as broad as long, with three carinae; dorsellum with a fine medial longitudinal carina; pro- podeum short, rugose, with crenulae branching from areola; propodeal areola lens-shaped, 0.47 times as broad as long, propodeal spiracles elliptical, 2.2 times as long as broad; epicnemial carina present ventrally; sternaulus shallow, broad, coarse- ly crenulate; mesopleuron smooth and shiny above sternaulus, below sternaulus distinct- ly reticulate-rugose. Legs: Fore tibial spur as long as basitar- sus; hind tibia expanded at apex, 2.9 times as wide as at base; hind tibia and tarsus with brown spines among pilosity; inner hind tibial spur 1.6 times as long as outer spur, 0.75 times as long as basitarsus; hind basitarsus moderately compressed with flat- tened dorsal ridge at base; tarsal claws pec- tinate, hind claw with nine brown teeth. Wings: Fore wing (Fig. 13) with dense pilosity, much thicker towards apex; stigma 3.7 times as long as wide; 2r 0.46 times inside height of second submarginal cell, arising 0.62 times length along stigma; Rs bent at basal 0.15, then nearly straight to 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON apex; second submarginal cell 2.9 times as long as wide; 2+3-SR slightly arched near 2-M; 1-SR+M 3.0 times as long as m-cu; 2-SR+M 1.3 times as long as m-cu; 1-M straight; m-cu 0.5 times as long as 1-M; I- M 0.67 times as long as 1-SR+M; discal cell 1.5 times as long as wide; 2cu-a rep- resented by pigmented area; la present, spectral in apical two-thirds; 1-2A+3A spectral, reaching to la; 1-CUa 0.46 times as long as 1-CUb; hind wing with six even- ly spaced hamuli; 2-1A and 2A absent; I- SRa shortly present. Metasoma: T1 short, widened apically, as long as broad, stem 0.3 times as long as bulb, tergite beside stem finely longitudi- nally striate, bulb largely smooth, but ba- sally rugulose, medio-longitudinally con- cave (Fig. 14); suture between T2 and T3 represented by well-defined groove; median field of T2 0.32 times as long as wide; T2 punctulate; ovipositor sheaths broad and long, 0.48 times as long as hind tibia, 0.34 times as broad as long, densely hairy along length, broadening to and rounded at apex with dorsal notch; hypopygium with mod- erately dense hair laterally, hairless at apex, rounded and acutely angled at apex, evenly sclerotized except for medio-longitudinally less sclerotized and folded inwards. Color: Black; palpi, fore tibia, tarsus and apex of femur, middle tarsus and basal Y; of middle tibia, basal /; of hind tibia brownish yellow to orange, first tergite except for stem and bulb dark red; wing membrane brown, much paler basally, stigma and veins dark brown. Male.—Unknown. Material examined.—Holotype @ , China: Zhejiang, Suichang, Mt. Jiulong, 1994.viii. 18, Chen Xuexin, No.944703. Note.—This new species is similar to E. shezu, but can be separated from the latter by the indistinct clypeal tubercles; medio- longitudinal carina on face much less dis- tinct; wing membrane much less brownish, hind claw with nine teeth, hind wing with six evenly spaced hamuli; and antenna with 41 segments. Etymology.—This species is named after the mountain where the holotype was col- lected. The Chinese word “‘Jiulong’’ means nine dragons. There is an old legend about this mountain. Ancient Chinese people be- lieved that there were once nine dragons living in the mountain before people lived nearby. Eurycardiochiles shezu Chen, Whitfield, and He, new species (Figs. 16-18) Female.—Body length 7.9 mm, fore wing length 7.9 mm. Head: Eyes with dense fine pilosity; head sculptured with moderately dense long pilosity; labio-max- illary complex just visible below mandible; malar suture present; epistomal suture vis- ible; clypeal margin convex laterally, with straight medio-lateral part and convex me- dial part with two small tubercles (Fig. 18); clypeus broad, convex and distinctly ru- gose, 2.3 times as broad as high; face 1.5 times as broad as high, coarsely transverse- ly rugose, with a distinct medio-longitudi- nal carina; frons transversely striate, with a medial carina reaching to anterior ocellus; vertex not raised behind ocelli, narrow, coarsely transversely striate; depth of occi- put in dorsal view 0.47 times length of head; length of eye 1.2 times length of tem- ple in dorsal view; POL:OD:OOL = 8:6: 18; antenna with 46 segments, short, thick, scape 1.9 times as long as broad; antennal sockets separated by 0.8 times width of an- tennal socket. Mesosoma: Side of pronotum coarsely longitudinally striate ventrally, dorso-me- dial longitudinal furrow coarsely crenulate, rest striate; notauli present, deep, crenualte, of even width, meeting posteriorly; scutum distinctly rugose, a medio-longitudinal ca- rina present in medio-longitudinal groove; parapsidal grooves present; scutellum with dense long pilosity, coarsely rugose, 1.4 times as broad as long, lateral carinae dis- tinct; scutellar sulcus narrow, 3.4 times as broad as long, with four carinae; dorsellum without medial longitudinal carina; propo- VOLUME 106, NUMBER 1 49 Figs 16-18. view. 18, Head, dorsal view. deum short, rugose, with crenulae branch- ing from areola; propodeal areola lens- shaped, 0.5 times as broad as long, propo- deal spiracle elliptical, 2.3 times as long as broad; epicnemial carina present ventrally; sternaulus shallow, broad, coarsely crenu- late; mesopleuron smooth and shiny above sternaulus, below sternaulus distinctly retic- ulate-rugose. Legs: Fore tibial spur 0.89 times as long as basitarsus; hind tibia expanded at apex, 2.7 times as wide as at base, medio-longi- tudinally slightly concave on outer side at apical %; hind tibia and tarsus with brown spines among pilosity; inner hind tibial spur 1.6 times as long as outer spur, 0.65 times as long as basitarsus; hind basitarsus mod- erately compressed with flattened dorsal ridge at base; tarsal claws pectinate, hind claw with eight teeth. Wings: Fore wing (Fig. 16) with dense pilosity, much thicker towards apex; stigma 3.0 times as long as wide; 2r 0.48 times Eurycardiochiles shezu, holotype. 16, Fore wing. 17, First to third metasomal tergites, dorsal inside height of second submarginal cell, arising 0.63 times length along stigma; Rs bent at basal 0.23, then nearly straight to apex; second submarginal cell 2.6 times as long as wide; 2+3-SR slightly arched near 2-M; 1-SR+M 3.3 times as long as m-cu; 2-SR+M 1.6 times as long as m-cu; 1-M straight; m-cu 0.40 times as long as 1-M; 1-M 0.72 times as long as 1-SR+M;; discal cell 1.7 times as long as wide; 2cu-a pre- sented by pigmented area; la present, spec- tral in apical two-thirds; 1-2A+3A spectral, reaching to la; 1-CUa 0.42 times as long as 1-CUb; hind wing with eight evenly spaced hamuli; 2-1A and 2A absent; 1-SRa present. Metasoma: T1 short, widened apically, as long as broad, stem 0.25 times as long as bulb, bulb distinctly punctate-rugose (Fig. 17); suture between T2 and T3 rep- resented by well-defined groove; median field of T2 0.29 times as long as wide; T2 punctate; ovipositor sheaths broad and long, 50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0.48 times as long as hind tibia, 0.34 times as broad as long, densely hairy along length, broadening to and rounded at apex with dorsal notch; hypopygium with mod- erately dense hair laterally, hairless at apex, rounded and acutely angled at apex, evenly sclerotized except for medio-longitudinally less sclerotized and folded inwards. Color: Black; fore tibia, tarsus and apex of femur, middle tarsus and basal % of mid- dle tibia, basal 0.6 of hind tibia brownish yellow to orange, first tergite except for stem and bulb dark red; wing membrane en- tirely evenly brown, stigma and veins brown to dark brown. Male.—Body length 7.8—8.1 mm, fore wing length 8.0—8.2 mm. Antenna slender, 46 segments. Material examined.—Holotype 2, China: Zhejiang, Longquan, Mt. Fengyang, 1,400— 1,600 m, 1982.viii.16—27, Xu Junhuan, No.826636. Paratypes: 1 ¢, China: Zhe- jiang, Wuyanling, 1983.viii.1, Cai Zhenbin, no.833380; 1 ¢6, China: Zhejiang, Lishui, 1989.vi.27—29, Xu Deling, no.896560. Note.—This new species is similar to E. dongzu, but can be separated from the latter by the larger body; shorter first tergite (as long as wide); Tl and T2 punctate; wing membrane evenly entirely brownish; hind claw with eight teeth; hind wing with eight evenly spaced hamuli; and antenna with 46 segments. Etymology.—This species is named after one of the Chinese minorities, She Zu, who mainly live in Zhejiang, Jiangxi, Anhui and Fujian provinces. Psilommiscus Enderlein, 1912 Psilommiscus Enderlein 1912: 98. Type species (by monotypy): Psilommiscus su- matranus Enderlein. This genus was brought out of synonymy with Cardiochiles s. l. as the sister group of Hansonia Dangerfield based on the phylo- genetic analysis of Dangerfield et al. (1999). Mercado and Wharton (2003) cast doubt upon the sister-group relationship with Hansonia, so currently the relation- ships of this genus are uncertain. It shares with Hansonia the presence of two clypeal tubercles and the reduction of eye setae, but can be separated from the latter by the pres- ence of the occipital carina in the malar re- gion, the apical scutellum with a cup- shaped medial pit, Rs curved in the fore wing, and the broad form of T1 and T3, which are laterally compressed and elon- gate in Hansonia. The genus is known only from the type species, P. sumatranus En- derlein in Sumatra, and three undescribed species from Brunei, Peninsula Malaysia, and eastern Malaysia (Dangerfield et al. 1999). This genus is reported here for the first time in China and the type species is re- corded below from the northeastern part of the Oriental Region in China. Psilommiscus sumatranus Enderlein Psilommiscus sumatranus Enderlein 1912: 98. Material examined.—1 2, China: Fujian, Shaowu, 1945.x.1, Zhou Xiufu (= Chao Hsiufu); 1 @, China: Fujian, Guadun, 1963.vii.12, Chen Jiahua. Distribution.—China (Fujian); Sumatra. Note.—This species is new to China. ACKNOWLEDGMENTS We thank Dr. S. A. Belokobyski (St. Pe- tersburg, Russia) for checking the holotype of Cardiochiles rugosus Telenga and for providing the information on the transfer of Cardiochiles japonicus Watanabe (Japan) and C. turga Belokobylskij (Russia Far East) into Austerocardiochiles. The project was partly supported by a National Scien- tific Foundation of China (NSFC number: 39970099) to the first author. LITERATURE CITED Achterberg, C. van. 1993. Illustrated keys to the sub- families of the Braconidae (Hymenoptera: Ichneu- monoidea). Zoologische Verhandelingen, Leiden 283: 1-189. Austin, A. D. 1990. Revision of the enignmatic Aus- VOLUME 106, NUMBER 1 tralasian genus Miropotes Nixon (Hymenoptera: Braconidae: Microgastrinae), with comments on the phylogenetic importance of the female ovo- positor system. Systematic Entomology 15: 43-— 68. Belokobylskij, S. A. 1996. Nine new species of Bra- conidae (Hymenoptera) from the Russian Far East. Journal of Natural History 30: 1661—1681. Chen, X., J. He, J.. and Y. Ma. 1998. Revision of the genus Hartemita Cameron (Hymenoptera: Bra- conidae: Cardiochilinae) from China. Entomotax- onomia 19: 208-218. Chou, L. Y. 1995. The Braconidae (Hymenoptera) of Taiwan 5. Cardiochilinae and Orgilinae. Journal of Agricultural Research China 44: 174—220. Dangerfield, P. C. and A. D. Austin. 1995. Revision of the Australian species of Cardiochilinae (Hyme- noptera: Braconidae). Invertebrate Taxonomy 9: 387-445. Dangerfield, P. C., A. D. Austin, and J. B. Whitfield. 1999. Systematics of the world genera of Cardi- ochilinae (Hymenoptera: Braconidae). Inverte- brate Taxonomy 13: 917-976. Dolphin, K. P. and D. L. J. Quicke. 2001. Estimating the global species-richness of incompletely de- scribed taxa: an example using parasitoid wasps (Hymenoptera: Braconidae). Biological Journal of the Linnean Society 73: 279-286. Enderlein, G. 1906. Die Indo-australischen und Afri- canischen Cardiochilinen. Stettiner Entomologis- che Zeitschrift 67: 245-252. . 1912. Neue Gattungen und Arten von Bra- coniden. Archiv fuer Naturgeschichte 78: 94-100. ail Huddleston, T. and A. K. Walker. 1988. Cardiochiles (Hymenoptera, Braconidae), a parasitoid of lepi- dopterous larvae, in the Sahel of Africa, with a review of the biology and host relationships of the genus. Bulletin of Entomological Research 78: 435-461. Kokoujev, N. 1895. Fragments braconologiques. Ho- rae Societatis Entomologicae Rossicae 29: 363— 392 Mercado, I. and R. A. Wharton. 2003. Mexican car- diochiline genera (Hymenoptera: Braconidae), in- cluding a preliminary assessment of species- groups in Toxoneuron Say and Retusigaster Dan- gerfield, Austin and Whitfield. Journal of Natural History 37: 845-902. Quicke, D. L. J. and C. van Achterberg. 1990. Phy- logeny of the subfamilies of the family Braconi- dae (Hymenoptera: Ichneumonoidea). Zoologis- che Verhandelingen Leiden 258: 1—95. Szepligeti, G. V. 1902. Tropische Cenocoeliden und Braconiden aus der Sammlung des Ungarischen National-Museums. Termeszetrajzi Fiizetek 25: 39-84. Whitfield, J. B. and P. C. Dangerfield. 1997. Subfamily Cardiochilinae, pp. 176-183. Jn Wharton, R. A., P. M. Marsh, and M. J. Sharkey, eds. Manual of the New World Genera of the Family Braconidae (Hymenoptera). Special Publications of the Inter- national Society of Hymenopterists 1. 439 pp. Whitfield, J. B. and W. R. M. Mason. 1994. Mende- sellinae, a new subfamily of braconid wasps (Hy- menoptera: Braconidae), with a review of rela- tionships within the microgastroid assemblage. Systematic Entomology 19: 61-76. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 52-73 THE WORLD SUBGENERA OF GLOSSOSOMA CURTIS (TRICHOPTERA: GLOSSOSOMATIDAE), WITH A REVISION OF THE CHINESE SPECIES OF GLOSSOSOMA SUBGENERA SYNAFOPHORA MARTYNOV AND PROTOGLOSSA ROSS JOHN C. MORSE AND LIANFANG YANG (JCM) Department of Entomology, Clemson University, Clemson, South Carolina 29634-0365, U.S.A. (e-mail: jmorse@clemson.edu); (LY) Department of Entomology, Nanjing Agricultural University, Nanjing, Jiangsu Province 210095, PR.C. (e-mail: Ifyang @njau.edu.cn) Abstract.—Newly discovered morphological evidence suggests that there are six prin- cipal monophyletic lineages of Glossosoma globally. We designate these as subgenera and infer the relationships among them using PAUP (Ripaeglossa ((Protoglossa + Lipoglossa) (Muroglossa (Glossosoma + Synafophora)))). Species of Glossosoma subgenus Synafo- phora are reported from China for the first ttme. Subgenus Synafophora is represented in Palearctic China by five species: Glossosoma (Synafophora) altaicum (Martynov), G. (S.) intermedium (Klapalek), G. (S.) minutum (Martynov), G. (S.) nylanderi (McLachlan), and G. (S.) ussuricum (Martynov). Glossosoma (Protoglossa) currently includes two species, both of which are known only from Oriental China. We transfer the Indian (Sikkim) species Glossosoma (Lipoglossa) kchinam Schmid 1971, to subgenus Protoglossa and review the Chinese species of subgenera Synafophora and Protoglossa. Key Words: logeny, male genitalia The genus Glossosoma Curtis 1834, is widely distributed in the Oriental and Hol- arctic Biogeographic Regions and contains 115 species and 2 subspecies (Morse 2001). Ross (1956) inferred nine subgenera for the genus but Morse and Yang (1993) inferred only six subgenera. Schmid (1980, 1998) said that the genus “is of Oriental origin, with its most primitive subgenera, Lipo- glossa Martynov and Muroglossa Ross, be- ing located there.”” A review of these opin- ions is warranted, requiring careful exami- nation of representatives of all purported groups and reinterpretation of several char- acteristics of male genitalia that were thought by Ross (1956) to be novelties in the genus. Trichoptera, Glossosomatidae, Glossosoma, Synafophora, Protoglossa, phy- Thirty-two species of Glossosoma have been reported from China to date. Major cooperative expeditions in southeastern, south central, and north central China in 1990, 1996, and 1998 have increased our knowledge of Chinese Glossosoma species considerably. Yang and Morse (2002) re- ported on the species of Glossosoma sub- genus Lipoglossa. In this paper, we are able to report five Glossosoma species of sub- genus Synafophora Martynov that are new records for China and to review the two Chinese species of Protoglossa Ross. All specimens used in this research are pre- served in the collections of Nanjing Agri- cultural University (NAU) unless otherwise indicated (see Acknowledgments). VOLUME 106, NUMBER 1 Nn Oo Ripaeglossa NA 17 world species Muroglossa OL 9 (3 Chinese spp.) Glossosoma EPNAOL WP 34( 8) (4), 6, 11, Synafophora) EP NA WP OS) (4), 7, 8, 10, 17, (21) Protoglossa OL 31( 2) 14, 20, (21) Lipoglossa EPO 30 (14) 13, 19 Tree length = 24 species incertae sedis 3 Consistency Index (CI) = 0.917 Homoplasy Index (HI) = 0.083 1S (32) Retention Index (RI) = 0.800 Rescaled Consistency Index (RC) = 0.733 Eigse: Glossosoma Subgenera The family Glossosomatidae was revised at the family-group and genus-group levels by Ross (1956) and Morse and Yang (1993). In the course of redescribing the species herein and reviewing the structure of additional species, our knowledge of the synapomorphic homologues at the generic and subgeneric levels for Glossosoma was much improved. Besides the consolidation of several of his subgenera, our phylogeny (Fig. 1; inferred with PAUP version 3.1.1.; CI = 0.917, HI = 0.083, RI = 0.800, RC = (0.733) differs from that of Ross (1956) because of new understanding of structures of sternum IX, the inferior appendages, and the phallus. By examination of specimens from a broader range of species than were available to Ross (the genus has more than doubled in number of species since 1956), we have come to realize that the structures he called ‘“‘claspers” (= inferior append- ages) are, in fact, a ventral projection of sternum IX and that his new so-called “fixed spiny lobes of the aedeagus” are, in fact, the inferior appendages, much modi- fied in this genus. Our interpretations of these and other synapomorphies and rein- terpretations of our earlier (1993) conclu- sions, relative to outgroups Agapetus (Glos- sosomatidae) and Rhyacophila (Rhyaco- Phylogeny, distributions, and species numbers of subgenera of Glossosoma. EP = East Palearctic, NA = Nearctic, OL = Oriental, WP = West Palearctic. philidae) are numbered below to correspond with the numbers in Fig. 1. As a result of this analysis, we still rec- ognize six subgenera, although the justifi- cation for them now is more firmly estab- lished by this investigation. The six sub- genera and their synonyms are as follows (Morse and Yang 1993): Glossosoma Curtis 1834 Anseriglossa Ross 1956 Lipoglossa Martynov 1930 Muroglossa Ross 1956 Protoglossa Ross 1956 Sinoglossa Ross 1956 Ripaeglossa Ross 1956 Synafophora Martynov 1927 Diploglossa Martynov 1934a Eomystra Martynov 1934a Klapalekia Botosaneanu 1955 Mystroglossa Kobayashi 1982 Mystrophora Klapalek 1892 Mystrophorella Kloet and Hincks 1944 MORPHOLOGY AND SYNAPOMORPHIES Unguitractor plate-——For his new genus Synafophora, Martynoyv (1927) described a new character, an enlarged inner unguitrac- tor lobe present apically on each front leg in all males of the Synafophora. Later, he used this character as a major diagnostic 54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON feature for Synafophora in his key to his Glossosomatidae genera (Martynov 1934a). We now submit that this character is found in all major species groups of Glossosoma and probably is a synapomorphy (Synapo- morphy #1) for the genus (Figs. 2—6). Male forewing callosity.—It has long been noted that the males of many Glos- sosoma species have a “‘callosity”’ in the anal region of each forewing (McLachlan 1879), with the venation of females normal. However, the details of this peculiarity have not previously been compared widely throughout the genus. There appear to be three general patterns that seem not to be part of the same transformation series: Pattern #1: Male forewing vein 2A is elongated (Synapomorphy #2; Figs. 8-12, 60, 66) in subgenera Lipoglossa and Pro- toglossa, and in many species it extends nearly to the arculus (Figs. 8, 12, 60, 66). Vein 2A (Fig. 8) and sometimes also 3A (Figs. 8, 10—12, 60, 66) are thickened in a variable pattern (Synapomorphy #3). When both veins are thickened, the two are some- times fused with each other over much or all of their length, forming a broad diffu- sion area which is only slightly thickened, flat, and translucent. Pattern #2: Male forewing vein 2A is shortened and strongly curved toward 3A, often weakened apically, usually accompa- nied by a rounded, thickened, translucent diffusion area on the posterior side or both sides of the vein (Synapomorphy #4; Figs. 16—25) in subgenera Ripaeglossa and Syn- afophora and some Glossosoma. We are unable to distinguish consistent patterns among these callosities, although other characters suggest that Ripaeglossa is dis- tantly related to subgenera Glossosoma and Synafophora, suggesting a homoplasious parallelism in this ““synapomorphy.” Some species have a dorsally protruding flap (Figs. 15, 16, 21), but this flap is not ho- mologous in the three subgenera: The flap seen in some species of subgenus Glosso- soma (Figs. 15a, 15b) entails a highly mod- ified anastomosis of 1A and 3A; the flap seen in some species of Synafophora (Fig. 21) involves a modified vein 3A at mid- length; and the flap seen in some species of Ripaeglossa involves no vein modifications (Fig. 16). Pattern #3: Male forewing vein 2A is shortened and extremely swollen in some species of subgenus Glossosoma, forming an ovate area typified by very well-defined margins and very thick, leather-like, opaque appearance (Figs. 13—14). It is quite con- sistent and is remarkably different from the callosity of those species of subgenus Glos- sosoma that have the fan-like fold men- tioned above (Figs. 15a, 15b); there are no intermediate forms between these two types of callosity in subgenus Glossosoma. For now, we consider these two types of cal- losity as possible synapomorphies that may have relevance for recognizing species groups within subgenus Glossosoma. Sternum IX.—The ventral process of segment IX [= unpaired ventral appendage of the 9th sternite (Martynov 1935); ventral process of IX (Kimmins 1953); tongue of IX (Ross 1956)] is evident in the lineage of Muroglossa + Synafophora + Glossosoma (Figs: 26; 28). “v.p:EX’”). Thisc structure (Synapomorphy #5) clearly arises from seg- ment IX without articulation, although its base is sometimes quite slender; it is sym- metrical in Glossosoma (Muroglossa), asymmetrical in Glossosoma (Glossosoma) (Ross 1956, figs. 327B, 328B; Synapomor- phy #6), and deeply divided (and some- times asymmetrical) in Glossosoma (Syna- fophora) (Figs. 28, 35, 40, 45, 51; Syna- pomorphy #7). In Synafophora, the sclerotization of the ventral process of sternum IX continues into the phallocrypt as a high keel (Figs. 26, 30; Ross 1956) from which the inferior appendages arise (Synapomorphy #8). Inferior appendages.—In subgenera Mu- roglossa + Glossosoma + Synafophora, the inferior appendages are vertical basally, with the bases parallel with the caudal edg- es of segment IX and attached by narrow membranous strips laterally with segment Nn Nn VOLUME 106, NUMBER 1 10 Figs. 2-12. Characters of Glossosoma species. 2—6, Distotarsus, claws, and unguitractor lobes on Glosso- soma male front legs; a. = dorsal, b. = ventral, c. = mesal. 2, G. (G.) aequale Banks, 1940, right front leg. 3, G. (Ripaeglossa) califica Denning, right front leg. 4, G. (Synafophora) minutum, left front leg. 5, G. (Synafo- Phora) altaicum, left front leg. 6, G. (Lipoglossa) adunatum Yang and Morse, left front leg. 7, G. (G.) aequale, female right front leg, ventral. 8, G. (L.) mirabile Yang and Morse, male right forewing, dorsal. 9-12, Male right forewing anal region, dorsal. 9, G. (L.) adunatum. 10, G. (L.) tortum Yang and Morse. 11, G. (L.) kelleyi Ross. 12, G. (L.) phyllon Yang and Morse. 1A, 2A, 3A = independent first, second, and third anal veins; 1A+2A+3A = fused anal veins; I, II, III, 1V, V = first, second, third, fourth, and fifth apical Forks; arculus = junction of 1A+2A+3A and Cu, on hind margin; Cu, = second cubital vein. 56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 18 Figs. 13-18. Forewings of male Glossosoma (G.) and G. (Ripaeglossa) species. 13, G. (G.) subaequale Schmid, male right forewing, dorsal. 14—18, Male right forewing anal cell, dorsal, a. = fold of callosity in normal position, b. = fold of callosity lifted. 14, G. (G.) n. sp. A. 15, G. (G.) n. sp. B. 16, G. (R.) calificum. 17, G. (R.) velonum Ross. 18, G. (R.) brunum Denning. IX, such that only the apex of each inferior appendage, characteristically setose, is free from “the smembrane> (Fics, 26,730; “inf.app.”’; Synapomorphy #9; Schmid 1971, figs. 33, 38). For subgenus Glosso- soma, JRosswCl95G6; elo: ties. SSAG lt,” and 327A, “‘tf’’) referred to these inferior appendages as the “‘lateral tendons” and their setose, free ends as “‘tendon flaps.” The inferior appendages of subgenus Synafophora are deeply imbedded in the phallocrypt, intimately appressed to the phallus, embracing it laterally (Figs. 26, 33, 38, 43, 49; Synapomorphy #10). Ross (1956) referred to this form of the inferior appendages in Synafophora as the ‘‘dorsal lobes of the aedeagus.” The apices of the inferior appendages of subgenus Muroglossa are approximate dor- sally above the phallus, anteriorly forming a bridge above the phallus that is nearly or completely fused (Synapomorphy #12). Inferior appendages are elongate in Li- poglossa (Morse and Yang 1993, #39, as “ventral lobes of inferior appendages”’; Synapomorphy #13 herein). Inferior appendages are short and trian- gular in lateral view in Protoglossa (Figs. 56, 62; Synapomorphy #14). Tergum X.—Tergum X is reduced in size and retracted beneath tergum IX and pre- anal appendages in Ripaeglossa (Morse and Yang 1993, #31 as “‘tergum IX hood-like”’; Synapomorphy #15 herein). Paired preanal appendages and tergum X are fused with each other and generally in- distinguishable in Muroglossa + Glosso- soma + Synafophora + Protoglossa + Li- VOLUME 106, NUMBER 1 poglossa(Pigs:, 26, Zig ~~ X< + prapp.”: Morse and Yang 1993, #32; Synapomorphy #16 herein). Phallocrypt sclerotizations.—A pair of sclerotized strips is seen dorsally in the phallocrypt of subgenus Glossosoma. These dorsolateral strips are long, broad, folia- ceous, and often setose, connecting the dor- solateral edge of the phallobase with the re- spective halves of the combined tergum X and preanal appendages (Synapomorphy Hild). Ross (1956; -e:s., figs: 327A, 329A, dt’) referred to these structures as “‘dorsal tendons.” In subgenus Synafophora, a unique pair of well-developed lateral strips of the phal- locrypt extend on each side from the api- colateral edge of the keel-bearing plate of sternum IX to the mesal bases of the re- spective halves of the combined tergum X and preanal appendages (Figs. 26, 33, 38, 43, 49, “‘lat.str..’; Synapomorphy #17). These may be narrow (e.g., Fig. 26) or very broad (Fig. 49). Ross (1956, figs. 314A, 315A, 318E, 320A, “‘t’’) referred to these structures as simply the “‘tendons.”’ The phallic shield is sclerotized in vari- ous degrees laterally and ventrally in Pro- toglossa and Lipoglossa, fusing with the basomesal surfaces of the inferior append- ages (Fig. 59; Synapomorphy #18). This represents a new understanding of Morse and Yang’s (1993, #33) “‘basal plate cup- ping the phallobase.” In Lipoglossa, this sclerotization 1s ex- tensive, causing the ventral phallocrypt to become a deep cup beneath the phallus (Yang and Morse 2002, fig. 5; Synapomor- phy #19). Phallus.—The phallobase and phallicata are fused in Protoglossa (Figs. 59, 65; Syn- apomorphy #20). Paramere spines are absent in Synafopho- ra and Protoglossa (Figs. 29, 59; Morse and Yang 1993, #34 and 36; Synapomorphy #21 herein), apparently having been lost in- dependently in these lineages. The paramere spine is single, on a long erectile lobe, and terminating in a conical 57 tuft of bristles in subgenus Glossosoma (Ross 1956, fig. 331E; Synapomorphy HZ): The phallicata is very long and com- pressed, bladelike, and also the phallobase is elongate in Muroglossa (Ross 1956, fig. 325C; Synapomorphy #23). MONOPHYLY OF EACH SUBGENUS Ripaeglossa is a monophyletic group as evidenced by Synapomorphies #4 and 15. Synapomorphy #4 occurs in parallel in Glossosoma and Synafophora. Muroglossa is a monophyletic group as evidenced by Synapomorphies #12 and 23. Morse and Yang (1993) inferred its mono- phyly based on ‘“‘ventral lobes of inferior appendages broad apically,” but we now understand that this is a plesiomorphic, un- divided, and symmetrical condition of the ventral process of sternum IX. The subgenus Glossosoma is a monophy- letic group as evidenced by Synapomor- phies #4, 6, 11, and 22. Character #4 ap- pears similar in Ripaeglossa and Synafo- phora. Synapomorphy #6 is a reinterpreta- tion of Morse and Yang’s (1993, #42) “ventral lobe of left inferior appendage ab- sent.”” Synapomorphies #11 and 22 are newly observed. Morse and Yang (1993) inferred that Synafophora is monophyletic based on a single synapomorphy: the absence of para- mere spines (Synapomorphy #21 herein), although Synafophora is similar in this re- gard to Protoglossa. Character #4 of Syn- afophora is similarly expressed in Glosso- soma and Ripaeglossa. However, with four additional synapomorphies (#7, 8, 10, 17), the monophyly for Synafophora is corrob- orated. The characters observed for Protoglossa that suggested its monophyly for Morse and Yang (1993) were “‘basal plate cupping phallobase” and “‘paramere spines absent,” respectively characters #18 and 21 herein. The former is now shown to be a synapo- morphy for Protoglossa + Lipoglossa and 58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 19-25. Forewings of male Glossosoma (Synafophora) species. 19, G. (S.) ussuricum, male right fore- wing, dorsal. 20-24, Base of male right forewings, dorsal. 20, G. (S.) dulkejti. 21, G. (S.) nichinkata. 22, G. (S.) nigrior. 23, G. (S.) nylanderi. 24, G. (S.) minutum. 25, G. (S.) intermedium, male right forewing, dorsal. the latter is also seen in Synafophora. In this work, we observed additional Synapo- morphies #14 and 20 in support of the monophyly of this subgenus. Morse and Yang (1993) observed for Li- poglossa “superior appendages with basov- entral hook” (their #38) and “‘ventral lobes of inferior appendages long” (their #39, Synapomorphy #13 herein). We now con- sider the hooks and other projections of the preanal appendages too variable to provide reliable evidence for monophyly. We add VOLUME 106, NUMBER 1 here Synapomorphy #19 in support of the monophyly of this subgenus. RELATIONSHIPS OF SUBGENERA Morse and Yang (1993) inferred that Synafophora + Lippoglossa + Muroglossa + Glossosoma are a monophyletic group based on “‘dorsal projections of basal plate present”? (their #35). We now understand that these “projections” are actually infe- rior appendages and that they do not have the same configuration in Lipoglossa as in the other three subgenera. The subgenera Muroglossa, Glossosoma, and Synafophora are an unresolved trichotomy. Their sister- group relationship is evidenced by Syna- pomorphies #5 and 9, of which #5 is newly observed and #9 is a reinterpretation of Morse and Yang’s (1993) character #35. Synapomorphies #2 and 3 are new ob- servations and Synapomorphy #18 is a re- interpretation of Morse and Yang’s (1993) character #33, supporting a sister-group re- lationship for Protoglossa and Lipoglossa. These two clusters of genera are proba- bly sister groups as suggested by Synapo- morphy #16 (Morse and Yang 1993, #32), leaving Ripaeglossa as the basal lineage of the genus. Ross (1956) inferred the same monophyletic group based on similar evi- dence [‘“‘cerci (?) absent”’ ]. Glossosoma (Synafophora Martynov 1927) Synafophora Martynov 1927: 165. Type species by original designation: Synafo- phora minutum Martynov. Mystrophora Klapalek 1892: 459 [preoc- cupied by Kayser, 1871, in Brachiopoda]. Type species by monotypy: Mystrophora intermedia Klapalek. Synonym of Eomystra according to Ross 1956: 154. Eomystra Martynoy 1934a: 84; 1934b: 327. Type species by monotypy: Eomystra dulkejti Martynov. Synonymy with Syn- afophora implied by Schmid 1971: 619; synonym of Synafophora by placement of type species in that subgenus by Lev- anidova 1982: 156. Diploglossa Martynov 1934a: 85; 1934b: 3) 327-328. Type species by monotypy: Di- ploglossa nylanderi McLachlan. Syno- nym of Synafophora by placement of type species in that subgenus by Levan- idova 1982: 156. Mystrophorella Kloet and Hincks 1944: 97, nomen novum for Mystrophora Klapalek, 1892, nec Kayser, 1871. Klapalekia Botosaneanu 1955: 792, nomen novum for Mystrophora Klapalek 1892 nec Kayser, 1871. Mystroglossa Kobayashi 1982: 7. Type spe- cies by original designation: Mystrophora altaica Martynov. Synonym of Synafo- phora by placement of type species in that subgenus by Levanidova 1982: 156. Martynov (1927) distinguished his genus Synafophora from its apparent ally Mystro- phora by three diagnostic characters. (1) Third apical fork pedicellate. This is a var- iable character. In our examples of type spe- cies Synafophora minutum, three male specimens have the pedicel of the third api- cal fork very short and one male specimen is without a pedicel. Furthermore, the char- acter is a plesiomorphy, common in Rhy- acophila, Agapetus, and Glossosoma (Mu- roglossa). (2) The fourth fork is shorter than the third. This is true in all members of most genera in this family. (3) “In the male anterior legs inner pulvilli form, each, a thick pale outgrowth inwards, nearly equal in the length to the claws.’” We found that this is true not only in Synafophora, but uniquely so in the males of all species of this genus, as we discussed above. In the male genitalia, the purportedly diagnostic characters Martynovy (1927) provided are no more than autapomorphies of the species. Therefore, we agree with Ross (1956) that Mystrophora and its replacement names Mystrophorella and Klapalekia are syno- nyms of Synafophora. We reexamined Glossosoma (Synafopho- ra) dulkejti, the type species of Eomystra, using specimens from the Kuriles (Figs. 53-55). The species bears all of the syna- pomorphies of Synafophora, such that we 60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON are able to confirm Schmid’s (1971) opin- ion that Eomystra is a synonym of Syna- fophora. Glossosoma (Synafophora) altaicum and G. (S.) nylanderi are redescribed and reil- lustrated below. These type species share most of the synapomorphies of Synafopho- ra, confirming the opinions of Levanidova (1982) that Mystroglossa and Diploglossa, respectively, also are synonyms of Synafo- Phora. Diagnostic characters for the subgenus include Synapomorphies #7, 8, 10, and 17 above. Also, the species of this subgenus usually differ from those of all other sub- genera except Ripaeglossa and some spe- cies of Glossosoma in that male forewing vein 2A is shortened and strongly curved toward 3A, often weakened apically, usu- ally accompanied by a rounded, thickened, translucent diffusion area on the posterior side or both sides of the vein. In addition, paramere spines are absent in only subgen- era Synafophora and Protoglossa. Further- more, males of all species examined by us have the phallocrypt above the phallus in- flated to accommodate the apices of the in- ferior appendages; this possibly synapo- morphous inflation is supported dorsally by a pair of sclerotized tendons in the phallo- crypt wall (Figs. 26, 27, 38, 39, 43, 44), occasionally fused in a single dorsal plate (Figs. 33, 34, 49, 50). The 19 world species of subgenus Syn- afophora are widely distributed in the Hol- arctic Biogeographic Region; its five Chi- nese species, redescribed below, occur only in the Palearctic portion of China. Glossosoma (Synafophora) World Species G. (S.) altaicum Martynov 1914b, Far East Russia, Korea, Japan, northeastern China. G. (S.) angaricum Levanidova 1967, Far East Russia. G. (S.) dulkejti (Martynov 1934a), Far East Russia. G. (S.) hospitum (Tsuda 1940), Japan. G. (S.) inops Tsuda 1940, Japan. G. (S.) intermedium (Klapalek 1892), Hol- arctic Region. G. (S.) lividum (Hagen 1861), northeastern North America. G. (S.) kiritchenkoi (Martynov 1927), cen- tral Asia. G. (S.) minutum (Martynov 1927), central Asia. G. (S.) neffi Arefina 2000, Far East Russia. G. (S.) nichinkata Schmid 1971, Japan. G. (S.) nigrior Banks 1911, eastern North America. G. (S.) nylanderi McLachlan 1879, Palearc- tic Region. G. (S.) sadoense Kobayashi 1982, Japan. G. (S.) spinatum Ruiter 2000, northwestern North America. G. (S.) sumitaense Kobayashi 1982, Japan. G. (S.) uogalanum Kobayashi 1982, Japan. G. (S.) ussuricum (Martynov 1934a), Far East Russia, northeastern China. G. (S.) verdona Ross 1938, western United States. KEY TO MALES OF CHINESE GLOSSOSOMA (SYNAFOPHORA) SPECIES il. Forewing vein 2A resembling other veins. Inner apical spur of each hind leg broad basally, abruptly narrowed and hooked ap- cially (Fig. 31). Tergum X + preanal ap- pendages each with two, simple, large acute projections and without multiple teeth (Fig. 26). Ventral processes of sternum IX with inner branches slender and much shorter than lateral branches (Fig. 28) .. . cae deere pede DR Berra G. (S.) altaicum - Forewing vein 2A callous (Figs. 41, 47). Inner apical spur of each hind leg tapered and curved (Fig. 36). Tergum X + preanal appendages each with 5—15 small acute projections (Figs. 33, 38, 43, 49, 53). Ven- tral processes of sternum IX with inner branches thick (Figs. 35, 40) or longer than lateral branches (Figs. 40, 45, 54) or absent (pty. Sons Sener eee 2 2(1'). Tergum X + preanal appendages each slen- der and with 3—4 long, slender, irregular teeth (Figs. 38, 39). Ventral processes of sternum IX with inner branches much broader than lateral branches (Fig. 40) . . . BOE Oe Es ME a Se AN hes G. (S.) minutum - Tergum X + preanal appendages broad and with 5—15 apical teeth of more nearly uni- VOLUME 106, NUMBER 1 form size and shape (Figs. 33, 43, 49, 53). Ventral processes of sternum IX with inner branches much more slender than lateral branches (Figs. 34, 45, 54) or absent (Fig. SD at Sous is paeseeniyes- ed eee eee eee os 3 . Lateral branches of tergum X + preanal ap- pendages each with about 15 spines of uni- form shape and size (Figs. 33, 34). Ventral processes of sternum LX with inner branch- es shorter than lateral branches (Fig. 35) G. (S.) intermedium - Lateral branches of tergum X + preanal ap- pendages each with about 5 spines that may be uniform in size and shape (Fig. 49) or of different lengths (Fig. 43). Ventral pro- cesses of sternum IX with inner branches longer than outer branches (Fig. 45) or with only one pair of processes (Fig. 51) .... 4 Lateral branches of tergum X + preanal ap- pendages with spines of different lengths (Fig. 43). Ventral processes of sternum IX four in number (Fig. 45) G. (S.) nylanderi _ Lateral branches of tergum X + preanal ap- pendages with spines of uniform lengths (Fig. 49). Ventral processes of sternum IX UNA) tial IMAM Yer (ESS SID) Stowe ble cee e 4(3’). Glossosoma (Synafophora) altaicum (Martynov 1914b) (Figs. 5, 26-32) Mystrophora altaica Martynoyv 1914b; 72— 84; holotype male (Russia, Altai), prob- ably Zoological Institute, St. Petersburg. Mystrophora lauta Tsuda 1940: 191; holo- type male (Japan, Honshu), destroyed (M. Tanida, personal communication). Synonymized by Ross 1956: 155. Glossosoma (Eomystra) altaicum: Ross 1956: 155: Synafophora altaicum: Botosaneanu 1970: 289. Glossosoma (Mystroglossa) altaicum: Ko- bayashi 1982: 7, type species of Mys- troglossa. Glossosoma (Synafophora) altaicum: Lev- anidova 1982: 156; Vshivkova 1986: 63— 64. This species, widely distributed in the Palearctic Region, now is found in China. It is one of a few members in this subgenus with 2A of the male forewing normal, but the inner spur of the male hind legs is high- 61 ly modified (Fig. 31). It somewhat resem- bles G. (S.) nylanderi in that (1) the pair of dorsolateral tendons of the phallocrypt have their anterior ends more widely separated from the phallobase (Figs. 30, 45) and (2) the setose apices of the inferior appendages are not so strongly recurved. These two species, together with G. (S.) dulkejti, all have their ventral process of sternum IX di- vided into four projections, but G. (S.) al- taicum is similar to G. (S.) dulkejti in the shape of the upper branch of each process (Figs. 26, 53). Unlike other species of G. (Synafophora), the inferior appendages are not angled but are more nearly aligned with the phallus throughout their length. Length of forewing: Male 5.1—7.4 mm; female 6.9—8.2 mm. Length of body: male 7.9—9.2 mm. Material examined.—HEI-LONG-JIANG PROVINCE: Tie-li County (N46.9, E128.0): 4 3, Lang-xiang, Xing-lin Bridge, 200 m elevation, 4 Aug. 1993, Coll. Sun Chang- hai; 85 males, 120 2, Lang-xiang, Bai-lan- he Farm, Bai-lan River, 5 Aug. 1993, 160 m elevation, Coll. Sun Chang-hai; 2 6, 2 2, Yi-chun City (N47.7, E128.9), Wu-yi-lin Town, Yong-sheng, Wu-yun River, 160 m el- evation, 31 July 1993, Coll. Sun Chang-hai. Shang-zhi County (N45.2, E127.9): 40 6, 60 2, Wei-he, Yu-ling Forest Farm, Chong- he River, 380 m elevation, 15 July 1993, Coll. Li You-wen and Sun Chang-hai; | 6, 12 2, Wei-he, 1 km N. of Yuling Forest Farm, Da-ling-gou, tributary of Chong-he River, 380 m elevation, 16 July 1993, Coll. Li You-wen and Sun Chang-hai. Distribution.—Previously known from Far East Russia, Korea, and Japan, this spe- cies is now known in northeastern China: NEW CHINESE RECORD. Male 5.8—8.2 mm; fe- Glossosoma (Synafophora) intermedium (Klapalek 1892) (Figs. 25, 33-37) Mystrophora intermedia Klapalek 1892: 461, figs. 1-5; syntypes | female, | male [Bohemia, ‘‘Mumlava 5. VIII. (jedna fe- 62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON X + pr.app. ae eT . ‘ maces Figs. 26-32. Glossosoma (Synafophora) altaicum. 26-30, Male genitalia. 26, Lateral. 27, Dorsal. 28, Ventral. 29, Phallus, lateral. 30, Relationship of phallus to lateral tendons, retracted inferior appendages, and ventro- mesal keel of abdominal segment IX, ventral. 31, Inner spur of right hind leg of male, mesal. 32, Male abdominal sternite VI process, ventral. dor.lat.str. = dorsolateral strip of the phallocrypt (paired), inf.app. = inferior ap- pendage (paired), IX = abdominal segment IX, keel = keel of sternum IX, lat.str. = lateral strip of the phallocrypt (paired), phall. = phallus, v.p.[X = ventral process of abdominal sternum IX (paired), X + pre.app. = abdominal tergum X and preanal appendage (paired). male), Milnice 14. VIII. Geden male)’’], apparently Czech Republic Academy of Sciences, Prague. Many authors have re- ported this species in various scattered lo- cations in Europe and northern North America by this name or in genera Syn- afophora or Glossosoma or Klapalekia or Mystrophorella. Martynov (1914a) first reported it in eastern Asia (Russia: Kam- chatka) by this name. VOLUME 106, NUMBER 1 =* == — | ge pene a ee LN ee s E Rae oes —_ “eL —— — FEN aad “Fes tiny ob f, 34 Figs. 33-37. 63 ZL Uj Lea = Yt 35 Glossosoma (Synafophora) intermedium. 33-35, Male genitalia. 33, Lateral; 34, Dorsal: 35, Ventral. 36, Inner spur of right hind leg of male, mesal. 37, Male abdominal sternite VI process, ventral. Glossosoma (Eomystra) intermedium: Ross 1956: 154-155. Glossosoma (Synafophora) intermedium: Schmid 1980: 24; Levanidova 1982: 156; Vshivkova 1986: 69-70, 72; Schmid 1998: 24-25. Two male specimens collected from northeastern China are very similar to the description and illustration of G. (S.) inter- medium provided by Ross (1956) and Vshivkova (1986) in coloring, the shape of the spurs of the male hind tibiae, and the structure of the male genitalia. Our illustra- tions show clearly that the species has all the synapomorphies, which are shared in this subgenus. Anal cell of each male forewing with small, suboval thickened area bearing hairs; 2A very short and curved, with small, spu- 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rious, curved vein arising posteriorly at mid length (Fig. 25). Tibial spurs 2-4-4, with in- ner apical spur on each male hind tibia highly modified (Fig. 36). Witissspecies asxvery close to: G. GS»)rus- suricum Martynov in that the apicoventral process of sternum IX is divided into two simple broad and flat plates, with only their bases fused with each other (Figs. 35, 51); the mesal keel of sternum IX is very pro- nounced; the inflated phallocrypt above the phallus is supported by a single sclerotized plate; and the lower lateral lobes of tergum X bear several small, regular teeth apico- dorsally, each lobe with a long slender pro- cess arising basomesally. Along with G. (S.) minutum Martynov [and, we predict, with G. (S.) kiritchenkoi Martynov], these two species share an enlarged, partially sclerotized phallocrypt chamber above the phallus to accommodate the apices of the inferior appendages. Length of forewing: Male 6.4—6.6 mm. Length of body: Male 7.2—7.3 mm. Material examined.—2 ¢, HEI-LONG- JIANG PROVINCE: Yi-chun City (N47.7, E128.9) Wu-yi-lin Town, Yong-sheng, Wu- yun River, 160 m elevation, 31 July 1993, Coll. Sun Chang-hai. Distribution.—Holarctic, reported here for the first time from northeastern China: NEW CHINESE RECORD. Glossosoma (Synafophora) minutum Martynov 1927 (Figs. 4, 24, 38-42) Glossosoma (Synafophora) minuta Martyn- ov 1927: 167-168, pl 7 f 8-10; holotype male (Turkestan: “‘Vicinities of Kopal, Semiretchje district; 4-5 VI 16, Coll. W. Snitnikov”), probably Zoological Insti- tute, St. Petersburg. This species was collected for the first time in China (Xin-jiang Province) by Dr. W. Mey (in litt.). Martynov’s original de- scription was very detailed in the color pat- tern, forewing venation, and the general shape of the male genitalia. We can add a few more details. Vein 2A in each male forewing slightly thickened but not curved, with short, water- drop-shaped thickened area (Fig. 24); inner apical spur of each male hind tibia normal. Male genitalia (Figs. 38—42): Segment IX long dorsally, short ventrally, shortened gradually from top to bottom. Apicoventral process of sternum IX deeply divided mes- ally, conforming with those of rest of this subgenus; in this species, forming pair of long, broad lobes, very flat and approxi- mate mesally, each bearing small lateral pa- pillate projection near middle and much longer slender process from near lateral base. In ventral view, retracted mesal por- tion of sternum IX small, its mesal keel pre- sent but not pronounced. In lateral view, se- tose inferior appendages and pair of lateral tendons between phallobase and tergum X very distinctive. Phallocrypt enlarged and sclerotized dorsally as pair of broad plates, permitting movement of tips of inferior ap- pendages within resulting chamber. Side lobes of tergum X long, each composed of two major parts: (1) lower lateral, elongat- ed, finger-shaped outgrowth and (2) upper medial, elevated portion, with its apex pro- duced in long, slender, claw-shaped process and short, papillate mesal process. This species appears very similar to G. (S.) kiritchenkoi Martynov in the hooked apex of the upper part of tergum X. Both of them apparently have a close relationship with G. (S.) ussuricum and G. (S.) inter- medium by sharing the following synapo- morphies: (1) divided ventral process of sternum IX very flat and broad; (2) dorsal phallocrypt very deep and sclerotized, forming spacious chamber for apices of in- ferior appendages (character predicted for G. kiritchenkoi, since we have not yet seen this species); and (3) tergum X lobes each bearing dorsomesal process. Length of forewing: Male 5.1—6.0 mm; female 5.8 mm. Length of body: Male 5.8—6.8 mm; fe- male 6.7 mm. VOLUME 106, NUMBER 1 ‘nif? (pees x Figs. 38-42. 65 Glossosoma (Synafophora) minutum. 38—40, Male genitalia. 38, Lateral. 39, Dorsal. 40, Ventral. 41, Male right forewing, dorsal. 42, Male abdominal sternite VI process, ventral. Material examined.—XIN-JIANG PROV- INCE: 4 6, 1 2 (NAU) and 183 ¢, 71 (MNH), Tian-chi (N44.2, E88.0), Bao-dong Shan, 3-11 Oct. 1991 and 3—15 June 1992, Coll. W. Mey; 1 @, 8 larvae (MNH), U-ru- mu-qi Shi (N43.8, E87.6), Nan Shan, 3—15 junes1992, Colla W2 Mey; and: 3° larvae (MNH), Yining County (N43.9, E81.3), Bo- rohoroshan, N.W. of Yining, 11 X 1991, Coll. W. Mey. Distribution.—Previously known from “Turkestan,” now known also from Xin- jiang Province, northwestern China: NEW CHINESE RECORD. 66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Glossosoma (Synafophora) nylanderi McLachlan 1879 (Figs. 23, 43-48) Glossosoma nylanderi McLachlan 1879: 474, pl 50 f. 1—2; holotype male (Finland: Uleaborg), Helsingfors Museum. Diploglossa nylanderi: Martynov 1934a: 85-86; 1934b: 327-328. Glossosoma (Diploglossa) nylanderi: Ross 1956: 154. Glossosoma (Synafophora) nylanderi: Lev- anidova 1982: 156; Vshivkova 1986: 69, a Levanidova (1982) first assigned this species to subgenus Synafophora, but with no detailed explanation. Our illustrations, based on Chinese specimens, confirm that this species is clearly a member of Syna- fophora. Vein 2A on each male forewing only slightly thickened, nearly straight, and reaching 3A (Fig. 23), not as thickened, strongly curved, nor incomplete as for some other members of this lineage. Male genitalia: Apicoventral process of sternum IX only partly divided, with con- siderable fusion basally. Lower lateral lobes of tergum X each with four deeply and widely separated teeth and dorsomesal lobe with several smaller teeth, resembling some species of Glossosoma (Glossosoma). Spurs 2-4-4, apical spurs on male hind legs nor- mal, not modified. Inferior appendages in this species very peculiar, autapomorphic, extending caudad almost horizontally, then recurved cephalad near apex. Lateral ten- dons of phallocrypt conspicuous. Dorsal tendons of phallocrypt paired. Length of forewing: Male 5.8—7.3 mm;. Length of body: Male 6.7—8.4 mm. Material examined.—HEI-LONG-JIANG PROVINCE: 5 4, Tie-li City, Lang-xiang (N46.9, E128.8), Xing-lin Bridge, 200 m el- evation, 4 Aug. 1993, Coll. Sun Chang-hai. Distribution.—Palearctic Region, now re- ported here for the first time from the Pale- arctic part of eastern China: NEW CHI- NESE RECORD. Glossosoma (Synafophora) ussuricum (Martynov 1934a) (Figs. 19, 49-52) Mystrophora ussurica Martynov 1934a: 79-80; 1934b: 326-327; holotype male (Russia: South-Ussuri region), probably Zoological Institute, St. Petersburg. Glossosoma (Eomystra) ussuricum: Ross 19562" 137-5138. Nl54. 55: Glossosoma (Synafophora) ussuricum: Levanidova 1982: 156; Vshivkova 1986: 72-74. Ross (1956) considered this species most closely related to Glossosoma (Eomystra) inops Kobayashi. Our colleague T. Nozaki (in litt.) considers G. inops a possible syn- onym of G. ussuricum. Vein 2A on each male forewing only slightly curved, with small widened area at its end (Fig. 19). Spurs 2-4-4, inner apical spur on each male hind tibia highly modi- fied in shape, similar to that of G. inter- medium (Fig. 36). This species is closely related to G. intermedium as discussed for that species above. The inferior appendages are quite long and the phallocrypt above the phallus is especially cavernous in this spe- cies, supported by a single, wide, sclero- tized plate dorsally. Length of forewing: Male 6.4—6.9 mm; female 6.9—8.0 mm. Length of body: Male 7.4—8.2 mm; fe- male 7.7—8.9 mm. Material examined.—JI-LIN PROV- INCE: 3 6, 21 @, An-tu County (Song- jiang) (N42.5, E128.3), Chang-bai Shan, Er-dao-ba-he River, 1700 m elevation, 10 Aug. 1993, Coll. Sun Chang-hai. Distribution.—Previously reported from the South Ussuri region of Russia, now known from northeastern China: NEW CHINESE RECORD. Glossosoma (Protoglossa Ross 1956) Protoglossa Ross 1956: 152, 154, as sub- genus of Glossosoma. Type species by original designation: Glossosoma_ taen- latum Ross and Hwang. VOLUME 106, NUMBER 1 67 soe Pe “ ' ' ' ! Figs. 43-48. Glossosoma (Synafophora) nylanderi. 43—46, Male genitalia. 43, Lateral. 44, Dorsal. 45, Ven- tral. 46, Phallus and ventro-mesal keel of segment IX, lateral. 47, Male right forewing, dorsal. 48, Male abdom- inal sternite VI process, ventral. Sinoglossa Ross 1956: 152, 154, as subge- Ross (1956) described his subgenus Pro- nus of Glossosoma. Type species by orig- toglossa with only the type species. Morse inal designation: Glossosoma_ sellatum and Yang (1993) synonymized Ross’ mono- Ross and Hwang. Synonymized by basic subgenus Sinoglossa with Protoglos- Morse and Yang 1993: 140, 144. sa to minimize redundancy after they in- 68 pena ice Sana ce wee e = 51 Figs. 49-52. tral. 52, Male abdominal sternite VI process, ventral. ferred that the two genera and species are sister lineages. Although the present reas- sessment of characters revises and adds to our 1993 inferences, the sister-group rela- tionship, and thus our conclusion for syn- onymy, is confirmed. The type species of each of these two subgenera was described from China (Sichuan). We now add a third species to subgenus PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘ a fol rew~ees ) 1 -. i -—- Glossosoma (Synafophora) ussuricum. 49-51, Male genitalia. 49, Lateral. 50, Dorsal. 51, Ven- Protoglossa: Glossosoma (Protoglossa) kchinam Schmid, 1971, from India (Sik- kim) because it shares Synapomorphies #14, 20, and 21 with G. (P.) taeniatum and G. (P.) sellatum. Although Schmid consid- ered his species to be close to G. (P.) taen- iatum, he placed it in the subgenus Lipo- glossa because he (1958) disagreed with Ross’ subgeneric classification. VOLUME 106, NUMBER 1 53 Figs. 53-55. abdominal sternite VI process, ventral. Besides characters #14, 20, and 21, the subgenus may be recognized by the fact that the bases of the inferior appendages are fused and the sclerotization extends to the venter of the phallobase as a cup or groove for the unusually short phallus. The para- mere spines are absent (Synapomorphy #21) in species of both subgenera Proto- glossa and Synafophora. The three known species of subgenus Protoglossa occur only in the Oriental Bio- geographic Region; the two Chinese species are known only from Oriental Sichuan Province. Glossosoma (Protoglossa) World Species G. (S.) kchinam Schmid 1971, India (Sik- kim). G. (S.) sellatum Ross and Hwang 195 China (Sichuan). G. (S.) taeniatum Ross and Hwang 1953, China (Sichuan). eS) 69 54 Glossosoma (Synafophora) dulkejti. 53—54, Male genitalia. 53, Lateral. 54, Ventral. 55, Male Glossosoma (Protoglossa) sellatum Ross and Hwang 1953 (Figs. 56-61) Glossosoma sellatum Ross and Hwang 1953: 8, f 2A—B; holotype male (China: Sichuan), CFM. Glossosoma (Sinoglossa) sellatum: Ross 1956: 137, 138, 154, figs. 313A—E, chart Dis) The male holotype has been preserved in alcohol for almost 50 years, such that it is very difficult to see any color pattern, but we take this opportunity to provide new fig- ures for the male forewing and genitalia. Vein 2A on each male forewing distinc- tively elongated, with its distal end beyond the cross vein m-cu (Fig. 60); this vein and 3A thickened in pattern similar to those of Glossosoma (Protoglossa) kchinam and G. (P.) taeniatum. Male genitalia (Figs. 56-59, 61): Ster- num VI process very small. Tergum X lat- 70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 56-61. Glossosoma (Protoglossa) sellatum. 56-59, Male genitalia. 56, Lateral. 57, Dorsal. 58, Ventral. 59, Phallus, lateral. 60, Male right forewing, dorsal. 61, Male abdominal sternite VI process, ventral. eral lobes elongated and bearing long, stout setae. Inferior appendages short, triangular in lateral view, fused with each other in basal half and extending along venter of phallobase to its anterior end. Length of forewing: Male 5.8 mm, fe- male 6.4 mm. Material examined.—Holotype, ‘‘Sich- uan, Tu-pa-keo, 2,255 m elevation, 4 Sept. 1929, Coll. Stevens’? (CFM). Diagnosis.—This species is quite differ- ent from the other two members of this sub- genus in the elongated, lateral lobes of ter- gum X with long, stout setae; in the ventral position of the phallobase; and in the long fused bases of the inferior appendages ex- tending along the venter of the phallobase to its anterior end. Distribution.—Known only from the type locality. VOLUME 106, NUMBER 1 Figs. 62-70. 71 : Glossosoma (Protoglossa) taeniatum. 62—65, Male genitalia. 62, Lateral. 63, Dorsal. 64, Ven- tral. 65, Phallus, lateral. 66, Male right forewing, dorsal. 67, Male abdominal sternite VI, ventral. 68, Male abdominal sternite VI (left) and VII (right) sternite processes, lateral. 69—70, Female genitalia. Glossosoma (Protoglossa) taeniatum Ross and Hwang 1953 (Figs. 62—70) Glossosoma taeniatum Ross and Hwang 1953; holotype male (China: Sichuan), CFM. Glossosoma (Protoglossa) taeniatum: Ross 1956: 138, 154, fig. 298, chart 29. Specimens from northern Sichuan were compared with the holotype. New struc- tures are figured here and the female is as- sociated with confidence for the first time. 72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Forewing of male (Fig. 66) with vein 2A both widened and elongated nearly to level of m-cu cross vein, about two-thirds of area between 2A and 3A translucent, slightly thickened and bearing tiny setae. Sternum VI process very small (Fig. 67), as long as sternum VII conical process (Fig. 68). Male genitalia (Figs. 62—65): Inferior appendages short, triangular in lateral and ventral views, each with its dorsal and api- cal edges armed with short, stout setae; in- terior bases of inferior appendages fused with apical edge of phallobase, forming cup embracing phallus ventrally. Phallus very short, tubelike, phallicata fused with phal- lobase, beanlike spinous bump present on each side of phallicata; paramere spines ab- sent. Female genitalia (Fig. 69-70): Sperma- thecal sclerite simple, trough-like in ventral view; in lateral view, its middle portion with slight bump dorsally. Length of forewing: Male 5.8—6.0 mm, female 6.4—6.7 mm. Material examined.—Holotype, ‘‘Sich- uan, Tu-pa-keo, 2255 m elevation, 7 Sept. 1929, Coll. Stevens’” (CFM). Other mate- rial examined: 1 6, 4 2, from Sichuan Province, Wen-chuan xian (N31.4, E103.6), Wo-long-zheng, Jiao-mu-shan-cun, Pi-tiao- he, 1850 m elev. 21 June 1990, Coll. Yang Lianfang and Li Youwen (NAU); | d from same river and date, 3 km from Jiao-mu- shan-cun, Coll. Chen Xiao-en (Clemson University Arthropod Collection). Diagnosis.—This species resembles Glossosoma (Protoglossa) kchinam in the similar shapes of the inferior appendages and the phallus, both species with the phal- licata bearing a setose patch, but the shapes of the side lobes of tergum X are obviously different in the two species. Distribution.—Known only from Sich- uan Province, southwestern China. ACKNOWLEDGMENTS We are especially thankful to Wolfram Mey, Museum fiir Naturkunde, Humboldt- Universitat, Berlin (MNH); O.S. Flint, Jr., National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM); Philip Parrillo, Field Museum, Chicago (CFM); C. Favret, Hlinois Natural History Survey, Champaign (INHS); and J. Huber, Canadian National Collection, Bio- logical Resources Program, Eastern Cereal and Oilseed Research Centre, Ottawa (CNC), for arranging loans of type speci- mens. We also thank Takao Nozaki from Kanagawa Environmental Research Center, Japan, for providing specimens and infor- mation. We thank Tatyana Vshivkova and Sun-Jin Park for help with preparation of the manuscript. We are grateful to the fol- lowing people for their cooperation in col- lecting these materials during our expedi- tions: Sun Chang-hai, Li You-wen, Wang Bei-xin, Du Yu-zhou, Wu Hong, Ma Yun, Wang Xin-hua, Zhou Chang-fa, Gui Fu- rong, and Chen Xiao-en. Financial support for the major expeditions was provided by the Special University Doctoral Foundation of China (No. 970204), the National Sci- ence Foundation of China (No. 39770113), and the Department of Entomology, Clem- son University. This is Technical Contribution No. 4813 of the South Carolina Agriculture and For- estry Research System, Clemson Universi- ty. LITERATURE CITED Banks, N. 1940. Report on certain groups of neurop- teroid insects from Szechwan, China. Proceedings of the United States National Museum 88(3079): 173-220. Botosaneanu, L. 1955. Note trichopterologice (1). Bu- letin Stiintific Sectia de Biologie si Stiinte Agri- cole si Sectia de Geologie si Geografie 7(3): 791— 802. . 1970. Trichopteres de la République Démo- cratique-Populaire de la Corée. Annales Zoologi- ci, Polska Akademia Nauk, Instytut Zoologiczny 27(15): 275-359. Curtis, J. 1834. Descriptions of some [hitherto] non- descript British species of may-flies of anglers. London and Edinburgh Philosophical Magazine and Journal of Science (Philosophical Transac- tions of the Royal Society), third series 4(21): 120-1125; 212-2118" Kimmins, D. 1953. Entomological results from the VOLUME 106, NUMBER 1 Swedish expedition 1934 to Burma and British In- dia: Trichoptera (Rhyacophilidae, subfamilies Hy- drobiosinae, Glossosomatinae and Agapetinae). Arkiv f6r Zoologi, Series 2, 6(8): 167-183. Klapalek, E 1892. Trichopterologicky vyzkum Cech vy. R. 1891. Rozpravy Ceské Akademie Cisare Fran- tiska Josefa pro Vedy, Slovesnost a Ument v Praze 1, I1(23): 443—462. Kloet, G. S. and W. D. Hincks. 1944. Nomenclatorial notes on two generic names in the Trichoptera. The Entomologist 77: 97. Kobayashi, M. 1982. A classification for Japanese spe- cies of Glossosomatidae (Trichoptera, Insecta). Bulletin of the Kanagawa Prefecture Museum 13: 1-18, 11 pls. Levanidova, I. M. 1982. Aquatic Insects of the Moun- tain Region of Far Eastern USSR. NAUKA (Pub- lishing House), Leningrad (St. Petersburg), 216 PPp- Martynov, A. V. 1914a (“1913”). Trichoptera of the Kamtschatka Expedition. d’Entomologie 13: 476—481. . 1914b. Notes on the Trichoptera collected by the Prof. P. Suskin’s expedition to the Altai during 1912. Revue Russe d’Entomologie 14: 72-84. . 1927. Contribution to the aquatic entomofau- na of Turkestan. Annuaire du Musee Zoologique de l’Académie des Sciences de ’URSS 28(2): 162-193, 11 plates. . 1934a. Tableaux analytique de la faune de VURSS, 13: caddisflies, Trichoptera, Annulipal- pia, I. Institut Zoologique de 1’ Académie de Sci- ences. Pp. 1—322. . 1934b. Trichoptera Annulipalpia of the USSR with descriptions of new or litthe known species and genera. Institut Zoologique de 1’ Académie de Sciences. Pp. 323-343. . 1935. On a collection of Trichoptera from the Indian Museum. Records of the Indian Museum 37(2): 93-209. McLachlan, R. 1879. A Monographic Revision and Synopsis of the Trichoptera of the European Fau- na part 8: 429-500, pls. xlv—li. Morse, J. C., editor. 2001. Trichoptera World Check- list. http://entweb.clemson.edu/database/trichopt/ index.htm, effective 22 May 1999, updated 28 July 2000, 8 January 2001. Morse, J. C. and L. Yang. 1993. Higher classification of the Chinese Glossosomatidae (Trichoptera), pp. 139-148. In Otto, C., ed. Proceedings of the Sev- enth International Symposium on_ Trichoptera, Umea, Sweden, 3—8 August 1992. Backhuys Pub- lishers, Leiden, The Netherlands, 312 pp. Ross, H. H. 1956. Evolution and Classification of the Mountain Caddis-flies. University of Illinois Press, Urbana, 213 pp. Ross, H. H. and Q. Hwang. 1953. Some interesting Revue Russe 73 Chinese Species of Glossosoma (Trichoptera: Rhyacophilidae). Proceedings of the Entomologi- cal Society of Washington 55(1): 6-9. Schmid, EF 1958. A propos de deux récent ouvrages sur la phylogénie et la zoogéographie des Tri- chopteres. Miscelanea Zoologica, Barcelona 1(5): 1-27. . 1971. Quelque nouveaux Glossosoma orien- taux (Trichoptera: Glossosomatidae). Le Natural- iste Canadien 98: 607-631. . 1980. Les Insectes et Arachnides du Canada, Partie 7: Genera des Trichopteres du Canada et des Etats Adjacents. Agriculture Canada Publica- tion 1692, 296 pp. . 1998. The Insects and Arachnids of Canada, Part 7: Genera of the Trichoptera of Canada and Adjoining or Adjacent United States. NRC Re- search Press, National Research Council of Can- ada, Ottawa, Ontario, Canada, 319 pp. Tsuda, M. 1940. Zur kenntnis der Japanischen Glos- sosomatinen (Rhyacophilidae, Trichoptera). An- notationes Zoologicae Japonenses 19(3): 191— 194. Vshivkova, T. 1986. Caddisflies of the family Glos- sosomatidae Wall. (Trichoptera) in Far Eastern U.S.S.R.: 1. Subfamily Glossosomatinae, pp. 58— 75. In Makarchenko, E.A., ed. Contributions to Freshwater Organisms of the Far East. Academy of Science USSR, Vladivostok, 140 pp. Yang, L. and J. C. Morse. 2002. Glossosoma subgenus Lipoglossa (Trichoptera: Glossosomatidae) of China, pp. 253-276. In Mey, W., ed. Proceedings of the 10th International Symposium on Trichop- tera, Potsdam, Germany July 30—August 5, 2000, Goecke & Evers, Keltern, Germany, Deutsches Entomologisches Institut, Nova Supplementa En- tomologica, 15, 664 pp. Addition in proof:—Since submission of our manuscript, we note that Mr. David Ruiter (in press, Western North American Naturalist) has recently reviewed the species of sister genus An- agapetus, transferring ““Glossosoma (Anagape- tus) shmidi’’ Levanidova, 1979 (Trudy Vseso- yuznogo Entomologicheskogo Obshchestva 61: 92-95), to Glossosoma as a fourth unplaced spe- cies in the genus (bringing the genus Glossoso- ma to 116 species). This species apparently has Synapomorphies #1 and #16, placing it close to the two clusters of genera Muroglossa-Glosso- soma-Synafophora and Protoglossa-Lippoglos- sa. However, it has none of the synapomorphies (#2, 3, 5, 9, 18) of either of these clusters of genera nor does it have any evident new synap- omorphies to relate it to one or the other of them. Therefore, we infer that it may represent a third linage, sister to them in a trichotomy. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 74-84 A NEW SPECIES OF CYAMOPS MELANDER 1913 (DIPTERA: PERISCELIDIDAE) FROM JAPAN AND A REVIEW OF JAPANESE PERISCELIDIDAE MASAHIRO SUEYOSHI AND WAYNE N. MATHIS (MS) Insect Ecology Laboratory, Forestry and Forest Products Research Institute, 1 Matsunosato, Tsukuba, Ibaraki, 305-8687, Japan (Japan Society of the Promotion of Sci- ence, Domestic Research Fellow) (e-mail: msuey @ffpri.affrc.go.jp); (WNM) Department of Systematic Biology, Entomology Section, MRC-169, P.O. Box 37012, Smithsonian Institution, Washington, DC 20013-7012, U.S.A. (e-mail: mathis.wayne@nmnh.si.edu) Abstract.—Cyamops hotei Sueyoshi, a new species of Periscelididae, is described from Japan, and the Japanese species of Periscelididae are reviewed. In addition to C. hotei, four periscelidid species are reported, of which Periscelis (Myodris) annulata (Fallén) and Stenomicra (Podocera) angustiforceps Sabrosky are recorded from Japan for the first time. A key to all Japanese species and a distribution map are provided. Key Words: The Periscelididae are a small family of approximately 84 species in nine genera from all zoogeographic regions. The family is divided into two subfamilies: Perisceli- dinae and Stenomicrinae (Mathis and Papp 1998). Adults of both subfamilies are very small (2—2.5 mm), and those of Periscelis Loew and Stenomicra Coquillett have been collected at sap runs and grassy areas, re- spectively. Adults of Cyamops Melander have been collected near wet habitats (cf. streams, waterfalls, marshes, etc.) (Khoo 1985, Baptista and Mathis 1994). The lar- val biology of only a few periscelidid gen- era is known. The larvae of Periscelis breed in sap runs on trees, and a puparium of Cy- amops and larvae of Stenomicra have been found in a bog and on banana, Gramineae, pineapple, and Pandanaceae, respectively (Ferrar 1987). The genus Cyamops, which occurs worldwide, is assigned to the subfamily Stenomicrinae and currently includes 26 species (Baptista and Mathis 2000). Only Diptera, Periscelididae, Cyamops, new species, Japanese species SIX species are known from the Old World, however, and except for an unidentified species from Japan, no species are known from the Palearctic Region (Cyamops sp.: Tamaki 1997, as a member of Aulacigastri- dae, and Kubo 2000; Cyamops sp. 4: Bap- tista and Mathis 2000). Recently, we found specimens in good condition that represent the unidentified species in collections of the Forestry and Forest Products Research In- stitute. The specimens were collected in a Malaise trap and are part of the Diptera sampled in an insect survey of the Ogawa Forest Reserve (Ibaraki), a temperate forest in Honshu, Japan. Research and analysis of these collections are progressing (submit- ted), and description of this new species of Cyamops is needed for these studies. More- over, we have also had a chance to examine all specimens previously listed in the above reports from Japan. In this paper, we describe a new species of Cyamops trom Japan as part of a taxo- nomic review of Japanese Periscelididae VOLUME 106, NUMBER 1 that includes a key and a distribution map for all Japanese species of the family. We also provide additional records of Stenomi- cra albibasis Sabrosky and S. fascipennis Malloch from Japan (Sabrosky 1965) and record S. angustiforceps Sabrosky and Per- iscelis annulata (Fallén) from Japan for the first time. MATERIALS AND METHODS A total of 301 specimens were examined. These are preserved dry except for some specimens in 80% ethanol. External struc- tures were observed under a stereoscopic microscope; some male and female abdo- mens were removed and treated in 10% KOH solution at 40° C for 10 hours, stained with Chlorazol Black E, neutralized with acetic acid, washed in distilled water, and then observed in pure glycerol. Dissected abdominal structures are preserved in pure glycerol in plastic microvials pinned under the dried mounted specimen or enclosed to- gether with other body parts in 80% etha- nol. Right wings were slide-mounted in Eu- paral. All specimens examined have a serial number label attached and are deposited in the Biosystematics Laboratory, Graduate Institute of Social and Cultural Studies, Kyushu University, Fukuoka (BLKU) (Pr.1001—1004, 1008-1246, 1248-1272, 1281-1285, 2005), the Forestry and Forest Products Research Institute, Tsukuba (FFPRI) (Pr.1005—1007, 1271), Tamaki’s personal collection, Moroyama (Pr. 2003-— 2004, 2006—2012), the Smithsonian Insti- tution, National Museum of Natural Histo- ry, Washington, D.C. (Pr.3001—3014), and the Bernice P. Bishop Museum, Honolulu (Pr. 4001). Abbreviations of localities are as follows: C: City, P: Prefecture, T: Town, V: Village. Species with an asterisk are newly recorded from Japan. The terminology and abbreviations mostly follow McAlpine (1981), excluding terms exceptionally men- tioned. WD Cyamops hotei Sueyoshi, new species (Figs. la—h, 2a—b, d, 3a—b, 4) Cyamops sp.: Tamaki 1997: 146; Kubo 2000: 378. Cyamops sp. 4: Baptista and Mathis 2000: 504. Male.—Head (Fig. 1a) higher than long; frons black in ground color, with velvet-like pruinescence; orbital plate, ocellar triangle, and vertex black in ground color, polished; face and gena yellow in ground color, with grayish pruinescence except narrow facial carina polished; face wide as distance be- tween posterior ocelli (Fig. 2a); parafacial dark brown in ground color, covered with silvery pruinescence; occiput, and postgena covered with grayish pruinescence; clypeus yellow. Head chaetotaxy: 2 orbital, 1 ver- tical (inner vertical seta absent), 2 paraverti- cal, 10 postocular, 9-10 peristomal, | genal, 2 postgenal setae, all black; ocellar, posto- cellar. Antenna: scape, pedicel, and post- pedicel (Stuckenberg 1999; Ist flagello- mere: McAlpine 1981) yellow in ground color; arista black, with 12 dorsal and 3 ventral branches; pedicel with several setae on anterior margin. Mouthparts yellow. Thorax entirely dark brown in ground color, covered with sparse grayish pruinesc- ence except for polished propleural lobe. Thoracic chaetotaxy: 2 dorsocentral, 2 no- topleural, | katepisternal, 1 supra-alar, | postalar and 2 scutellar setae all black; post- pronotal, anepisternal, and anepimeral setae absent; minute dorsocentral setulae anterior to dorsocentral setae present; 2 rows of mi- nute acrostichal setulae on presutural region present. Wing (Fig. 2d) generally hyaline, 3.5 times long as wide, broadly infuscate dark brown on apices of cell 15,3, 4,5, and m, and along veins R,,; and CuA,; veins dark brown; vein R,,, slightly sinuate. Halter yellow in ground color, with dark brown knob. Legs entirely yellow except apex of hind femur, apex of 4th tarsomere, and Sth tar- 76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON lavas ie postabdomen in ventral view. c, Epandrial complex (epandrium, surstylus, and cerci) in caudal view. d, Epandrial complex in left lateral view. e, Epandrial complex in anterior view. f, Hypandrial complex (hypandrium, gonites, and phallapodeme) in ventral view. g, Phallus in left lateral view. h, Phallus in caudal view. Abbreviations: bp: basiphallus; dp: distiphallus; ep: epandrium; hyp: hypandrium; or: orbital seta; ov: outer vertical seta; pa: phal- lapodeme; pp: posterior process of S6—7; prg: pregonite; s: surstylus; 4—6S: 4—6th abdominal sternites; 4—7T: Cyamops hotei, male (holotype). a, Head in left lateral view. b, 4—Sth abdominal segments and 4—7th abdominal tergites. Scale bars: a = 0.50 mm; b-f = 0.10 mm. somere dark brown; no distinct setae and spines. Abdomen entirely dark brown; S6—7 (Baptista and Mathis 1994; 6th sternite: Khoo 1985) with distinct posterior process (Fig. 1b: pp) and largely expanded as semi- circular plate anteriorly. Genitalia (Fig. 1c— h): epandrium (Fig. 1c—e: ep) spherical; both surstyli (Cumming et al. 1995; posterior surstylus: McAlpine 1981) (Fig. Ic—e: s) asymmetrical, right surstylus short, bowl- like, left surstylus long, hoe-like, with abrupt apex; hypandrium (Fig. If) asym- metrical, right pregonite (Fig. If: prg) bowl-like, left pregonite (Fig. 1f: prg) hill- like with 3 short longitudinal ridges; phal- lus (Cumming et al. 1995; aedeagus: McAlpine 1981) (Fig. 1g—h) consisting of some sclerites; basiphallus (Fig. 1g—h: bp) cylindrical, with longitudinal projection an- teriorly; distiphallus (Fig. I1g—h: dp) con- sisting of 2 transverse strap-like sclerites, large sclerite expanded apically, terminated with cylindrical apex and sclerite with hook. Body length 2.7 mm (anterior apex of head to posterior margin of abdomen). Wing length (base of costa to apex of vein Rifts) 2-1 mm: width*O.7 mm: Female.—Similar to male but with sex- ual dimorphism in head, fore femur, and wing markings: head (Fig. 3a) entirely black in ground color without yellow re- gions; frons, parafacial, gena, occiput, and postgena covered with grayish pruinesc- ence; facial carina broad as distance be- VOLUME 106, NUMBER 1 Fig. 2. view. a, Cyamops hotei, male (holotype). b, C. hotei, female (Pr. 1004). c, Periscelis annulata (Pr. 1001). d, C. hotei (holotype). e, Stenomicra albibasis (Pr. 1014). f, S. fascipennis (Pr. 1008). g, S. angustiforceps (Pr. 1029). tween outside of antennae (Fig. 2b); lateral corner of face and gena covered with sil- very pruinescence; clypeus (Fig. 3a: cl) black; scape, pedicel, and basal part of post- pedicel black, distal part of postpedicel or- ange; wing broadly infuscate, dark brown along apex of veins R,,;, R,,;, M, and CuA,, apical infuscation of wing reaching cell r,;; anteroventral side of femur with row of minute spine-like setae. Postabdominal structures (Fig. 3b): T6 fused with S6, forming ring-shaped 6th syntergosternite (Fig. 3b: 6ST) (Baptista and Mathis 1994); Wi dhe aL a —4 age he, Fe a jy hres g Heads and wings of Japanese Periscelididae. a—b, Head in anterior view. c—g, Right wing in dorsal T7 separate from S7; T8 long as T5, S8 membranous, attached to S7; T10 (10th ab- dominal tergite: Griffiths 1981; epiproct: McAlpine 1981) and S10 (10th abdominal sternite: Griffiths 1981; hypoproct: Mc- Alpine 1981) triangular; cercus short (Fig. 3b: c). Two spherical spermathecae (Fig. 3b: sp). Specimens examined.—Holotype <¢. (Ta- maki’s personal collection): ““South hill of Hiki/Ogose Town/Iruma, Saitama Pref./ 11.v.1993/N. Tamaki leg.”’, lime green cir- cle, ‘‘Pr. 2005”, **Holotype/Cyamops/hotei/ 78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Cyamops hotei, female (Pr. 1004). a, Head in left lateral view. b, Postabdomen in left lateral view. Abbreviations: c: cerci; cl: clypeus; or: orbital seta; ov: outer vertical seta; sp: spermatheca; 6ST: 6th synter- gosternite; 7—10S: 7—10th abdominal sternites; 7—1OT: 7—10th abdominal tergites. Sueyoshi”’ (red label). Paratypes: 3 ¢ 6 @, HONSHU. Ibaraki P: | 6, Ogawa, Kitai- baraki C, 27.vi—9.vii.1996 (Pr. 1005); 1 &, same locality, 25.vii—6.viii. 1996 (Pr. 1007); 2 2, same locality as holotype, 9— 25.vul.1996 (Pr. 1006, 1271). Kanagawa P: 1 ¢, Hitorizawa, Yokohama C, 29.v.1999 (Pe2006) 21) 2, -Koajiro, Miura G) 127 vi 1995 (Pr. 1272). Saitama P: 1 2, Mt. Ari- ma, Chichibu C, 14.vii.2001 (Pr. 1004); 1 ?, Minowada, Moroyama T, 3.vi.1994 (Pr. 2003); 1 3, same locality, 17.vi.1994 (Pr. 2004). KYUSHU. Oita P: 1 ¢, Beppo (= Beppu), 21.vi.1952 (Pr. 3001). Distribution.—Japan (Honshu and Kyu- shu: Fig. 4). Etymology.—tThe specific epithet is de- rived from the Japanese noun Hotei, mean- ing a Buddhist priest from China who has a fan and pouch in his hands, alluding to the flat and pouchlike surstyli of this spe- GIES: Remarks.—This species is distinguished from congeners by the following combina- tion of characters: anteroventral portion of male eye with enlarged facets (Fig. 2a); wing cell R,,, broadly hyaline and with dark brown marking at apex (Fig. 2d); fore femur with ventral seta at base; anterior scutellar seta well developed; hind femur with apical dark marking; and female ab- dominal tergite and sternite 7 separate (Fig. 3b). This species is very similar to C. ban- vaneue Baptista and Mathis in wing mark- ings but differs from the latter species by having dark markings on the male hind fe- mur restricted to the apical ;, and the left surstylus is not bifurcate and terminated at abrupt apex (Fig. Ic, e). Cyamops hotei is also similar to C. Jaos Baptista and Mathis in male genital structures but differs from it by the wing cell R,,, without a distinct white region at middle (Fig. 2d), and the male left surstylus is abrupt at its apex (Fig. hewes)r Tamaki (1997) recorded an undetermined species of Cyamops from Japan for the first time and Kubo (2000) also reported a male belonging to this genus. We examined all four specimens they reported and found that they belong to this new species. The male specimen of Cyamops sp. 4 (recorded in Baptista and Mathis 2000) was also exam- ined and found to be conspecific with C. hoteti. Baptista and Mathis (1994) documented the monophyly of Cyamops with eight au- tapomorphies: absence of inner vertical VOLUME 106, NUMBER 1 seta, middle of frons concave, side of frons below orbital plate silvery shiny, male clyp- eus retracted and tending to fill oral open- ing, both of surstyli asymmetrical, phallus with complex sclerites, female abdominal tergite and sternite 6 fused and forming a complete ring. Cyamops hotei has all of these characters and is thus a member of this monophyletic group. Baptista and Mathis (1994) also proposed three synapo- morphies that support some subgroups within Cyamops, and C. hotei has two of them: anteroventral facets of male eye en- larged and presence of spine-like short setae on female fore femur. Cyamops hotei has two spermathecae (Fig. 3b: sp), a condition that Baptista and Mathis (1994) consider plesiomorphic. These authors suggested an- other five characters as potential synapo- morphies of subgroups, although their po- larities were not determined. Among these five synapomorphies, we observed the fol- lowing in C. hotei: female abdominal ter- gite and sternite 7 separate (Fig. 3b); 2 pairs of dorsocentral setae posterior to transverse suture present; 2 pairs of scutellar setae pre- sent; anepisternal setae absent; wing cross- vein bM-Cu present (Fig. 2d). RECORDS OF JAPANESE SPECIES OF PERISCELIDIDAE Periscelis (Myodris) annulata (Fallén) (Fig. 2c, 4) Notiphila annulata Fallén 1813: 250. Periscelis annulata: Loew 1858: 118 [ge- neric combination]. Specimens examined.—HONSHU. Ibar- aki P: 1 6, Ogawa, Kitaibaraki C, 25.vii— 9.viii.1996 (Pr. 1003); 1 3d 1 2, same lo- cality, 21.v.2002 (Pr. 1001—1002). Tokyo: 1 ?, Tama Forest Science Garden, Hachioji Ca —29'y-2002 (Pr 1281)22 2, same lo- cality, 10—22.vii.2002 (Pr. 1282-1283); 1 2, same locality, 22.vii—26.viii.2002 (Pr. 1284); 1 2, Sugawara Shrine, Machida C, 15—29.vi.2002 (Pr. 1285). Distribution.—Europe (Duda 1934) and Japan (Honshu: Fig. 4). 1) Remarks.—This species is distinguished from congeners of the subgenus Myodris Lioy by the following combination of char- acters: scutum covered with grayish prui- nescence and with 3 longitudinal dark brown stripes; face with distinct medial bulging and facial carina; wing without dis- tinct dark markings; thorn of male cerci subapical; left surstylus as long as right sur- stylus; postgonite gradually tapered to apex. This species is very similar to P. (M.) ka- buli Papp but differs from the latter species by the above genital characters (Papp 1988). Stenomicra (Podocera) albibasis Sabrosky (Fig. 2e, 4) Stenomicra albibasis Sabrosky 1965: 214. Specimens examined.—HONSHU. Ibar- aka sBapalt “Os. Osawa Kataibarakas ©: 19.v1.2002 (Pr. 1010); 1 2, Okami Marshy, Satomi V, 18.v.2001 (Pr. 1008); 1 3, same locality, 14.v.2002 (Pr. 1009). Kanagawa P: 26, 9 2. Mt: Enkai;. Yokohama 7G: 21.vii.2002 (Pr. 1011-1021). Distribution.—Japan (Honshu: Sabrosky 1965) (Fig. 4). Remarks.—This species is distinguished from congeners by the following combina- tion of characters: wing slightly infuscate, yellow except for basal ¥;, with small black markings at apex and base of vein R,, base of vein R,,3, and base of alula (Fig. 2e); wing vein R,,, terminated near wing apex (Fig. 2e); anterior orbital seta as long as posterior orbital seta; head entirely dark brown in ground color, covered with gray- ish pruinescence. This species resembles S. bicolor Sabrosky from Reunion and S. ni- gricolor Sabrosky from Kenya in having a black head and thorax. However, it differs from S. bicolor by having entirely yellow legs, and from S. nigricolor by having an entirely yellow abdomen (Sabrosky 1975). Stenomicra (Podocera) fascipennis Malloch (Fig. 2f, 4) Stenomicra fascipennis Malloch 1927: 26. Stenomicra sp. 1: Tamaki 1997: 147. 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Stenomicra albibasis, & S. angustiforceps. Specimens examined.—HONSHU. Ibar- aki P: 1 36, Okami Marshy, Satomi V, 14.v.2002 (Pr. 1069). Kanagawa P: 1 @, Mt. Enkai, Yokohama C, 16.vii.2001 (Pr. 1269); Lod 2; same locality, 16.v.2002. (Pr 1067-1068); 3 3, same locality, 27.vi.2002 0 100 200 300 400 Sd0Km ———EE—EE— Distribution maps of Japanese periscelidid species. a, OQ Cyamops hotei; @ Periscelis annulata, @ (Pr. 1058-1060); 5 5 1 2, same locality, 28:vi.2002.(Pe" 1061=1066)2.15. cai 2 same locality, 21.vi1.2002 (Pr. 1070—1095). Saitama P: 6 d 8 2, Mt. Arima, Chichibu Cyl 4iviiw 2001 @r 1045=1057 41270) 2a . Kanao, Yori T, 28.v.1995 (Pr. 2010—201 1); VOLUME 106, NUMBER 1 1.2, Omaeda, Hanazono T, 28.v.1995 (Pr. 2007); 1 d 1 @, same locality, 15.vii.1995 (Pr. 2008-2009). Tochigi P: | 6 1 2, Yusen Gorge, Nasu T, 29.1x.2001 (Pr. 1043-1044). Tokyo Metropolis: 2 ¢ 6 2, Tokyo, Siveloss (Pr 3002=3009). Shizuoka P: 1 3d, Gotemba C, 9.x.1952 (Pr. 3010). Kyoto Paleo 3)¢., Kibune, KyotoxC, 10.v.1953 (Pr. 3011-3014). RYUKYUS. 20 6 25 @, Oku, Kunigami V, 27.vi.2001 (Pr. 1096— 1139, 1042); 2 d 2 9, Okuyona Pass, Ku- migami V, 27.vi.2001 (Pr. 1159-1162); 10 Peis. same, locality, 16:vir2002 (er 1140-1158, 1169-1172); 1 3, Yona, Ku- nigami V, 21.iii1.1997 (Pr. 1163); 2 2, same locality, 25.vi.2001 (Pr. 1164-1165); 1 5 2 2, same locality, 16.vii.2002 (Pr. 1166— mooi 2.6 | £, Gyoba, Ogimi V, Wevaie2002. (Pr 1173-1175); 1, 2, Oura, Nago,@- 17.iv-1996: (Pr 1188);-7 6 12-2, same locality, 28.vi.2001 (Pr. 1176-1187, 1189-1195); 3 gd 4 2, Gogayama, Nakijin Pele va 2002 (Pre 205-1211): 5 6 3-2 Okawa, Nago C, 18 vu 2002 (Pr. 1196-— 1203); 1 3, Sesoko, Motobu T, 17.vii.2002 (Pr. 1204). OGASAWARA. 1 & Senzan, Ogasawara V, 28 v 1958 (Pr. 4001); 15 3 18 2, Mt. Mikazuki, Ogasawara V, 26.vi.2002 (Pr. 1212-1244); 1 ¢, Higashi Kaigan, Ogasawara V, 30.vi.2002 (Pr. 1245); 6 6 3 2, Kitafukurozawa, Ogasa- wara V, (Pr. 1246, 1248-1255); 5 6 2 &, Kita Bay, Ogasawara V, 4.vii.2002 (Pr. 1256-1262); 2 ¢ 4 2, Mt. Sakai, Ogasa- wara V, 5.vii.2002 (Pr. 1263-1268). Distribution.—India, Ceylon, Nepal, Ma- laya, Taiwan, Philippines, Borneo, Guam, Solomon, Fiji, Hawaii (Sabrosky 1977, Khoo and Sabrosky 1989) and Japan (Oga- sawara, Honshu: Sabrosky 1965; and Ryu- kyus) (Fig. 5). Remarks.—This species is distinguished from congeners by the following combina- tion of characters: head entirely yellow in ground color, covered with grayish prui- nescence; wing infuscate, dark yellow, with 4 hyaline transverse bands (Fig. 2f); wing vein R,,, terminated near wing apex (Fig. 2f); anterior orbital seta as long as posterior 81 orbital seta. This species is very similar to S. biconspicua Sabrosky from South Africa in having four band-like white markings of the wing but differs from the latter by its slender and strongly curved surstylus (cf. Sabrosky 1965: fig. 5, Sabrosky 1975: fig. 1O): Sabrosky (1965) examined 13 specimens from Tokyo, Shizuoka, and Kyoto prefec- tures and recorded this species from Japan for the first time. We examined all of these specimens as part of this study. Tamaki (1997) reported six specimens of Stenomi- cra sp. 1. We examined these specimens and found that five are this species. The mesonotum and abdominal dorsum of this species are typically dark colored (i.e. black or brown) (Malloch 1927, Sa- brosky 1965), and the specimens from Hon- shu and the Ryukyus correspond well to this description, but not the specimens from Ogasawara. The mesonotum and abdominal dorsum of specimens from Ogasawara are predominately yellow brown and paler than those from Honshu and the Ryukyus. The male postabdomen is also distinctly yellow. Although we could assign the specimens from Ogasawara as a distinct species from S. fascipennis, we have identified them as conspecific because there are no distinct differences, including structures of the male genitalia, between the specimens examined except for the body color as noted. Stenomicra (Stenomicra) angustiforceps Sabrosky (Figs. 2g, 4) Stenomicra angustiforceps Sabrosky 1965: DANG* Stenomicra sp. 2: Tamaki 1997: 147. Specimens examined.—HONSHU. Kan- agawa P: 3 ¢ 7 2, Mt. Enkai, Yokohama C, 21.vii.2002 (Pr. 1022-1031). Saitama P: 1 2, Iwai, Moroyama T, 27.vi.1995 (Pr. 2012): RYUKYUS? S5udv2'eesOuray Nazo C, 28.vi.2001 (Pr. 1035-1041); 3 2, Oku, Kunigami V, 27.vi.2001 (Pr. 1032—1034). Distribution—Nepal (Sabrosky 1965), 82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON India (Sabrosky 1977), and Japan (Honshu and Ryukyus: Fig. 4). Remarks.—This species is distinguished from congeners by the following combina- tion of characters: wing entirely hyaline (Fig. 2g); wing vein R,,, terminated before wing apex (Fig. 2g); anterior orbital seta g 100 200 300 400 S00Km ae distinctly shorter than posterior orbital seta; postocellar seta present; scutum scarcely covered with whitish pruinescence, yellow in ground color, with pair of longitudinal brown vittae laterad of dorsocentral seta; postabdomen yellow. Tamaki (1997) reported Stenomicra sp. 2 VOLUME 106, NUMBER 1 based on a female specimen. We examined his specimen and found that it belongs to this species. KEY TO JAPANESE SPECIES OF PERISCELIDIDAE (Modified from Sabrosky 1965 and Mathis and Papp 1998) 1. Two pairs of orbital setae present (Fig. 1a); ocellar seta absent; wing costa extended to apex of vein M (Fig. 2d—g); postpronotal lobe WHUNOUE CISTINGR CARE 55 ooo coed abo b ob or 2 — One strong pair of orbital setae present (Mathis and Papp 1998: fig. 24.1); ocellar seta present; wing costa extended to apex of vein R,,; (Fig. 2d); postpronotal lobe with distinct setae 6 6 Ot 2, Ee cae Periscelis annulata . Orbital setae reclinate or lateroclinate (Grimal- di and Mathis 1993: figs. 15, 17); inner vertical seta present; supra-alar seta absent; wing cross- vein bM-Cu absent, wing cells bm and dm con- fluent; wing cell cup absent (Fig. 2e-g) .... 3 — Anterior orbital setae proclinate and posterior orbital setae reclinate (Fig. la); inner vertical seta absent; supra-alar seta present; crossvein bM-Cu present, wing cells bm and dm separate (Fig. 2d), wing cell cup present .. Cyamops hotei 3. Wing vein R,,; terminated at apex of wing (Fig. 2e—f); wing cell dm long; postocellar seta absent; anterior orbital seta as long as posterior orbital seta (Malloch 1927: fig. 11) — Wing vein R,,, terminated before apex of wing (Fig. 2g); wing cell dm short; postocellar setae present; anterior orbital seta distinctly shorter than posterior orbital seta (Malloch 1927: fig. 5 sao ae cen eS Stenomicra angustiforceps 4. Wing infuscate, light gray, with four transverse hyaline bands (Fig. 2f) .. Stenomicra fascipennis — Wing infuscate, light yellow, with hyaline at basal Y, (Fig. 2e) i) Stenomicra albibasis ACKNOWLEDGMENTS We thank Dr. K. Arakaki (Bernice P. Bishop Museum), Mr. H. Mitsui (Tama, To- kyo) and Mr. N. Tamaki (Moroyama, Sai- tama P) for the loan and donation of many specimens. We also thank members of the Department of Forest Entomology of the Forestry and Forest Products Research In- stitute for assistance with the first author’s field research. For reviewing a draft of this paper, we thank Dr. Stephen D. Gaimari (California State Collection of Arthropods). LITERATURE CITED Baptista, A. R. P. and W. N. Mathis. 1994. A revision of New World Cyamops Melander (Diptera: Per- iscelididae). Smithsonian Contributions to Zoolo- gy 563: 1-25. . 2000. Notes on the genus Cyamops Melander (Diptera: Periscelididae), including description of ten new species. Proceedings of the Entomologi- cal Society of Washington 102(3): 481-506. Cumming, J. M., B. J. Sinclair, and D. M. Wood. 1995. Homology and phylogenetic implications of male genitalia in Diptera~-Eremoneura. Entomologica Scandinavica 26(2): 121-151. Duda, O. 1934. Periscelididae, pp. 1-13. Jn Lindner, E., ed. Die Fliegen der palearktischen Region 6(1). E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart. Fallén, C. EK 1813. Beskrifning Oefver nagra 1 Sverige funna vattenflugor (Hydromyzides). Kongliga Ve- tenskaps-Academiens Handlingar 1813: 240—257. Ferrar, P. 1987. A guide to the breeding habits and immature stages of Diptera Cyclorrhapha. Ento- monograph 8: 77-78. Griffiths, G. C. D. 1981. Book Reviews, of Manual of Nearctic Diptera. Volume 1. J. E McAlpine et al., eds. Entomological Society of Canada, Bulletin 13: 49-55. Grimaldi, D. and W. N. Mathis. 1993. Fossil Periscel- ididae (Diptera). Proceedings of the Entomologi- cal Society of Washington 95(3): 383—409. Khoo, K. C. 1985(1984). The Australian species of Cyamops Melander (Diptera: Periscelididae). Aus- tralian Journal of Zoology 32(2): 527-536. Khoo, K. C. and C. W. Sabrosky. 1989. 75. Family Stenomicridae, p. 551. Jn Evenhuis, N. L., ed. Catalog of the Diptera of the Australasian and Oceanian Regions. E. J. Brill and B. P. Bishop Museum special publication, Honolulu, 86: 1— SS spp: Kubo, K., 2000. The insects of Enkai Mountain Re- gion, Diptera. Kanagawa-Chuho, Odawara, 130: 347-395. (In Japanese.) Loew, H. 1858. Ueber einige neue Fliegengattungen. Berliner Entomologische Zeitschrift 2: 101—122. Malloch, J. R. 1927. The species of the genus Sten- omicra, Coquillett (Diptera, Acalyptrata). Annals and Magazine of Natural History 20: 23-26. Mathis, W. N. and L. Papp. 1998. Family Periscelidi- dae, pp. 285—294. Jn Papp, L. and B. Davas., eds. Contribution to a Manual of Palaearctic Diptera, 3. Science Herald, Budapest, 880 pp. McAlpine, J. EF 1981. 2. Morphology and terminolo- gy—adults, pp. 9-63. In McAlpine, J. E et al., eds. Manual of Nearctic Diptera, Vol. 1. Mono- graph 27, Research Branch, Agriculture Canada, Ottawa, vi + 674 pp. Papp, L. 1988. Periscelis kabuli sp. n. and P. kazab sp. n. with notes on larvae and pupae of the fam- 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ilies Aulacigastridae and Periscelididae (Diptera). Acta Zoologica Hungarica 34(2/3): 273-284. Sabrosky, C. W. 1965. Asiatic species of the genus Stenomicra (Diptera: Anthomyzidae). Bulletin of the British Museum (Natural History), Entomol- ogy 17(5): 209-218. . 1975. The genus Stenomicra in the Ethiopian Region (Diptera, Aulacigastridae). Annals of the Natal Museum 22(2): 663-676. . 1977. Family Aulacigastridae, p. 230. In Har- dy, D. E. and M. D. Delfinado, eds. A Catalog of the Diptera of the Oriental Region, Volume III Suborder Cyclorrhapha (excluding Division As- chiza). The University Press of Hawaii, Honolulu, vi + 451 pp. Stuckenberg, B. R. 1999. Antennal evolution in the Brachycera (Diptera), with a reassessment of ter- minology relating to the flagellum. Studia Dipter- ologica 6(1): 33-48. Tamaki, N. 1997. Diptera of Saitama Prefecture, pp. 1—405. In Usui, T., ed. Insects of Saitama, Japan 2 (Diptera). Saitama Kontyu Danwakai, Omiya, 405 pp. (In Japanese.) PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 85-89 LARGIDEA DAVISI KNIGHT, A RARELY COLLECTED PLANT BUG (HEMIPTERA: MIRIDAE: DERAEOCORINAE) ASSOCIATED WITH PITCH PINE IN THE NORTHEASTERN UNITED STATES A. G. WHEELER, JR. Department of Entomology, Clemson University, Clemson, SC 29634, U.S.A. (e-mail: awhlr@clemson.edu) Abstract.—Described in 1917 and known previously only from Cape Cod, Massachu- setts, and Long Island and near New York City, New York (ca. 14 specimens), the der- aeocorine mirid Largidea davisi Knight is newly reported from Connecticut, Maine, New Hampshire, New Jersey, Pennsylvania, and Rhode Island. Additional records are given for Massachusetts and New York. This univoltine predator was found only on pitch pine (Pinus rigida Mill.), typically in extensive and remnant pine barrens. The gall-forming margarodid scale Matsucoccus gallicolus Morrison is suggested as prey of this late-season plant bug. Key Words: Insecta, Miridae, Deraeocorinae, Clivinematini, new records, Pinus rigida, pine barrens, Matsucoccus scales Largidea davisi Knight is the only east- ern member of a Nearctic genus of plant bugs. The other nine species of Largidea are essentially restricted to western North America (one [Illinois specimen, collected in 1895, is known of the otherwise western L. grossa Van Duzee [Knight 1941]). A mirid of the deraeocorine tribe Clivinematini, L. davisi has been termed a rare heteropteran of the northeastern fauna (Slater 1974) and has remained poorly represented in collec- tions since its original description (Knight 1917). Here I give new distribution records for this apparently predacious plant bug, provide notes on its seasonal occurrence and plant associations, and suggest a pred- ator-prey relationship. Largidea davisi Knight (Figs. I, 2) This dark reddish-brown, elongate-oval mirid, 6.0—6.5 mm long, can be recognized by the matted, pale yellow pubescence on its body; its short rostrum that is extended only to or slightly beyond the posterior margin of the forecoxae; the incrassate (@ ) or linearly thickened (d) second antennal segment, with the third and fourth segments short (0.37 mm) and thin; the coarsely and densely punctate pronotum with an im- pressed line, smooth and hook-shaped, from the anterolateral angle to the posterior mar- gin of the calli; and the broad, flattened bas- al tarsomere that is extended under the sec- ond tarsomere. For a diagnosis and descrip- tion of the tribe Clivinematini, see Ferrei- ra’s (2000) taxonomic review. DISTRIBUTION Historical.—Described from Promised Land, eastern Long Island, New York (Knight 1917), L. davisi later was recorded from Hyannis Port, Massachusetts, on Cape Cod (Knight 1923). Massachusetts was omitted as a state record in Carvalho’s (1957) catalog but was includéd by Henry 86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and Wheeler (1988) in the most recent cat- alog of North American Heteroptera. The only other published records are from New York: Yaphank (Long Island) and Bear Mountain, north of New York City (Leo- nard 1928). New records.—All specimens were col- lected from pitch pine by the author or (Connecticut) with T.J. Henry. Roman nu- merals denote nymphal instars. Voucher material has been deposited in the collec- tions of the Pennsylvania Department of Agriculture, Harrisburg, and the National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM). CONNECTICUT: Windham Co., Wind- ham Airport, 4 km NE of Willimantic, | Septe T99IsI"2! MAINE: York'Cor RE 1, Kennebunk, 6 sept. 2002.26 =2 2> Rt 4, 229) kin S08 jet. RE, 109) 6:9 kime SSE of Sanford, 7 Sept. 2002, 1 V. MASSACHUS- SETTS: Franklin Co., Montague Sand Plains, 3.5 km N of Montague, 14 Sept. 199i) Io 6 Sept, 20015 102 Re 202, 5:3°km SSE of Orange; 8 Sept. 2002, 2°9. NEW HAMPSHIRE: Hillsborough Co., Rt. 102, 1.4 km S of Rockingham Co. line, 5.7 km NNE of Hudson, 8 Sept. 2002, 7 °; Merrimack Co., Concord Barrens, Rt. 9, 14 Sept 199s di yQick& HID Ame 11993) lek: NEW JERSEY: Sussex Co., High Point State Park, Kuser Rd. W of Lake Marcia, 16 Sept. 2001, 2 2. NEW YORK: Ulster Co., Mohonk Preserve, Bonticou Crag, 5 km NNW of New Paltz, 21 Aug. 1992, 2 V; Sam’s Point Dwarf Pine Ridge Preserve, W of Lake Maratanza, NE of Cragsmoor, 5 Sept: 2001, 54d eS" 2a 4 Sept. 2002.40" 1 2. PENNSYLVANIA: Dauphin Co., Jef- ferson Twp., State Game Lands 210, 3 km SSE of Williamstown, 26—27 Sept. 1988, 2 2; Luzerne Co., Rt. 924, Humboldt, 2.3 km SW vofyjct4 8h... I Sept: 2001, Mad (Abs? 6e)3- Sept: 20022355, 23 PikecGonyRt: 739,125 km; SE,of LordssValleyal6! Sept: 2001, 3 2 & 3 Sept. 2002, 2 2; Schuylkill Co., jct. Rt. 61 & I-81 south, 1.2 km S of Frackville, 4 Sept. 2001, 1 ¢. RHODE IS- LAND: Providence Co., Sept. 19927 2-2. Slatersville, 19 SEASONALITY Except for a male collected on 10 August 1899 at Hyannis Port, Massachusetts, all other adults of L. davisi have been taken in September. The collection of equal numbers of males and females (5 of each) in Ulster County, New York, on 5 September 2001 and four males and one female on 4 Sep- tember 2002 suggests that adults had been present for only a week or 10 days. Mirids typically are protandrous, the males ap- pearing slightly earlier than the females and not living as long (Wheeler 2001). My lat- est collection of a male was 27 September in Dauphin County, Pennsylvania. The type series includes a pair taken in copula on 24 September 1910 (USNM collection). Despite extensive collecting of mirids on pitch pine from May to September, I found only four nymphs: a fourth instar on 12 Au- gust at Concord, New Hampshire, two fifth instars on 21 August in Ulster County, New York, and a fifth instar (see Fig. 1) on 7 September in York County, Maine. Collec- tions of L. davisi suggest that adults mostly appear from late August to early September. Assuming the date of 10 August given for the male taken on Cape Cod, Massachu- setts, in 1899 is correct (Knight 1923), this plant bug sometimes appears earlier in the season. The presence of late instars in mid- to late August suggests that overwintered eggs do not hatch until after mid-July. PLANT AND PREY ASSOCIATIONS The mostly Neotropical tribe Clivine- matini comprises 17 genera and 83 de- scribed species. Even though scant biolog- ical data are available, clivinematines are assumed to be predacious (Ferreira 1998). Two species of Clivinema and an Ambra- cius species feed on ensign scales (Orthe- ziidae); a Hemicerocoris species preys on soft scales (Coccidae) (Ferreira 1998, Wheeler 2001). The habits of Largidea spe- cies are unknown, although Kelton (1980) VOLUME 106, NUMBER 1 87 Figs. 1-2. speculated that L. shoshonea Knight feeds on aphids. Label data accompanying specimens of Largidea indicate that seven species are found on pines (Pinus spp.). Species of Largidea and its sister genus, the Neotrop- ical Adlargidea, likely are restricted to de- veloping on pines (Ferreira 1998). I have collected mirids from all species of Pinus occurring in the Northeast and have found L. davisi only on pitch pine (P. rigida Mill.). ‘“‘Pine,’’ recorded as the host at the type locality on Long Island, almost certainly refers to pitch pine. W.T. Davis and G.P. Engelhardt, who in 1910 collected the type specimens of this plant bug at Promised Land, Long Island, collected oth- er heteropterans that year from pitch pine at Promised Land (see Davis 1911). Spec- Largidea davisi. 1, Fifth instar (York Co., Maine, 7 Sept. 2002). 2, Adult female (Ulster Co., New York, 5 Sept. 2001). imens known historically from other local- ities—Cape Cod, Massachusetts; Bear Mountain, New York; and Yaphank, Long Island, New York—likely also were col- lected from P. rigida. The habits of L. davisi on pitch pine have not been determined. A potential prey spe- cies on pitch pine is the margarodid scale Matsucoccus gallicolus Morrison. This pine-bast scale is found mainly on pitch pine and is known from nearly all areas where L. davisi has been collected. The margarodid is particularly common on Cape Cod, Massachusetts, and Long Island, New York, as well as in parts of Connecticut, New Jersey, and Rhode Island (Morrison 1939, Parr 1939). The globular saclike adult females of this infrequently collected scale insect were taken with L. davisi at Monta- 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gue, Massachusetts, and in Dauphin, Luz- erne, and Pike counties in Pennsylvania. In Luzerne County, Pennsylvania, the mirid and the margarodid were collected from the same branches of pitch pine. In addition, the co-occurrence of the anthocorid Elato- philus inimicus Drake & Harris, a known predator of Matsucoccus species (e.g., Mendel et al. 1991), suggests the presence of M. gallicolus at other localities where L. davisi was found. Populations of Matsucoc- cus predators, such as Elatophilus species, are more likely to be detected than those of their inconspicuous prey (Morrison 1939, Mendel et al. 1991). DISCUSSION Previously published records of L. davisi evidently are based on fewer than 15 spec- imens: 10 from the type locality on Long Island, New York (Knight 1917); one from Hyannis Port, on Cape Cod, Massachusetts (Knight 1923); and likely only one from each of the two other localities in New York (Leonard 1928). Fifty-six adults were col- lected during the present survey, ranging from York County, Maine, in the north to northern Dauphin County, Pennsylvania, in the south. In elevation, L. davisi has been taken near sea level on Long Island, New York, and on Cape Cod, Massachusetts, to about 685 m near Lake Maratanza in Ulster County, New York. Largidea davisi appears to be an unusu- ally late-season, univoltine mirid. The dis- tinctive nymphs and adults are unlikely to have been overlooked during earlier-season collecting at sites where I found this plant bug in September, including the Montague Sand Plains in Massachusetts, the Concord Barrens in New Hampshire, the Mohonk Preserve and Sam’s Point Dwarf Pine Ridge Preserve in New York, and pitch pine com- munities in Luzerne and Schuylkill counties in Pennsylvania. Largidea davisi can be considered a characteristic, though uncommon and patchily distributed insect of northeastern pitch pine-scrub oak barrens and similar communities. It appears to be the least com- mon of the approximately 25 mirid species that develop on pitch pine (Wheeler, un- published data). It was found in several remnant pine barrens, such as Concord, New Hampshire, and on scattered pitch pine trees in ruderal sites along highways, in addition to being collected in the exten- Sive pine barrens in the Shawangunk Moun- tains of New York and the sand plains near Montague, Massachusetts. This mirid was not found in extensive pine barrens in southern Maine, such as Waterboro and Fryeburg; the Ossipee barrens in New Hampshire; the Albany (New York) Pine Bush; and the New Jersey Pine Barrens. Future research should focus on eluci- dating the feeding habits of L. davisi, ob- taining more information on its seasonal history, and determining whether pines oth- er than P. rigida serve as hosts. Clivine- matines have been assumed to be preda- cious, the few available records of their habits indicating that scale insects serve as prey. Cinara species and other pine-asso- ciated aphids also were found on pitch pines with L. davisi, and Kelton (1980) sug- gested that L. shoshonea Knight might feed on aphids inhabiting jack pine, Pinus bank- siana Lamb. For certain anthocorid species of the pine-associated genus Elatophilus, Kelton (1976) and others (cited by Mendel et al. 1991) speculated that these anthocor- ids are generalist predators on aphids or mites. But at least five species of Elatophi- lus now are known to be specialized pred- ators of Matsucoccus scales, and an inti- mate association among Elatophilus spe- cies, pines, and pine-bast scales has been hypothesized (Mendel et al. 1991, cf. Lattin and Stanton 1993). My hypothesis that members of the pine-associated genus Lar- gidea prey mainly on scale insects and that L. davisi feeds on a pine-bast scale, Mat- sucoccus gallicolus, awaits testing. ACKNOWLEDGMENTS I gratefully acknowledge J.E Stimmel (Bureau of Plant Industry, Pennsylvania VOLUME 106, NUMBER 1 Department of Agriculture, Harrisburg) for photographing the mirid and identifying the margarodid; PH. Adler (Department of En- tomology, Clemson University, Clemson, SC) for providing useful comments on an earlier draft of the manuscript; P.-C. Huth (Mohonk Preserve, New Paltz, NY) for ac- companying me in the field at Mohonk and facilitating my access to Sam’s Point Dwarf Pine Ridge Preserve; TJ. Henry (System- atic Entomology Laboratory, USDA, ARS, c/o National Museum of Natural History, Washington, DC) for checking the USNM collection for specimens of L. davisi and accompanying me in the field in Connecti- cut; M.D. Schwartz (Biological Resources Program, Agriculture and Agri-Food Can- ada, Ottawa) for providing label data from specimens borrowed from the USNM; and E.R. Hoebeke (Department of Entomology, Cornell University, Ithaca, NY) and R.T. Schuh (Department of Entomology, Amer- ican Museum of Natural History, New York, NY) for checking their respective collections for possible material of L. dav- ist. LITERATURE CITED Carvalho, J. C. M. 1957. A Catalogue of the Miridae of the World. Part 1. Cylapinae, Deraeocorinae, Bryocorinae. Arquivos do Museu Nacional (Rio de Janeiro) 44: 1-158. Davis, W. T. 1911. Noteworthy Hemiptera collected on Long Island, N.Y. Journal of the New York En- tomological Society 19: 112-113. Ferreira, P. S. E 1998. The tribe Clivinematini: Cla- distic analysis, geographic distribution and biolog- ical considerations (Heteroptera, Miridae). Revista Brasileira de Entomologia (Sao Paulo) 42: 53-57. . 2000. A taxonomic review of the tribe Cli- vinematini, with a key to world genera (Heterop- tera, Miridae). Studies on Neotropical Fauna and Environment 35: 38—43. Henry, T. J. and A. G. Wheeler, Jr. 1988. Family Mir- idae Hahn, 1833 (= Capsidae Burmeister, 1835): The plant bugs, pp. 251-507. /n Henry, T. J. and R. C. Froeschner, eds. Catalog of the Heteroptera, 89 or True Bugs, of Canada and the Continental Unit- ed States. E.J. Brill, Leiden. Kelton, L. A. 1976. The genus Elatophilus Reuter in North America with descriptions of two new spe- cies (Heteroptera: Anthocoridae). Canadian Ento- mologist 108: 631—634. . 1980. The insects and arachnids of Canada. Part 8. The plant bugs of the Prairie Provinces of Canada. Heteroptera: Miridae. Research Branch Agriculture Canada Publication 1703, 408 pp. Knight, H. H. 1917. New and noteworthy forms of North American Miridae (Hemip.). Entomological News 28: 3-8. . 1923. Family Miridae (Capsidae), pp. 422— 658. /n Britton, W. E., ed. Guide to the Insects of Connecticut. Part IV. The Hemiptera or Sucking Insects of Connecticut. Connecticut Geological and Natural History Survey Bulletin 34. . 1941. The plant bugs, or Miridae, of Illinois. Bulletin of the Illinois Natural History Survey 22(1): 1-234. Lattin, J. D. and N. L. Stanton. 1993. Taxonomic and biological notes on North American species of Elatophilus Reuter (Hemiptera: Heteroptera: An- thocoridae). Journal of the New York Entomolog- ical Society 101: 88—94. Leonard, M. D. 1928. A list of the insects of New York with a list of the spiders and certain other allied groups. Cornell University Agricultural Experi- ment Station Memoir 101, 1121 pp. Mendel, Z., E. Carmi, and H. Podoler. 1991. Relations between the genera Matsucoccus (Homoptera: Margarodidae) and Elatophilus (Hemiptera: An- thocoridae) and their significance. Annals of the Entomological Society of America 84: 502—507. Morrison, H. 1939. Descriptions of new species of Matsucoccus (Hemiptera: Coccidae). Proceedings of the Entomological Society of Washington 41: 1—20. Parr, T. J. 1939. Matsucoccus sp., a scale insect inju- rious to certain pines in the Northeast (Hemiptera- Homoptera). Journal of Economic Entomology 32: 624-630. Slater, J. A. 1974. A preliminary analysis of the deri- vation of the Heteroptera fauna of the northeastern United States with special reference to the fauna of Connecticut, pp. 145-213. /n Beard, R. L., ed. 25th Anniversary Memoirs, Connecticut Ento- mological Society, New Haven. Wheeler, A. G., Jr. 2001. Biology of the Plant Bugs (Hemiptera: Miridae): Pests, Predators, Opportun- ists. Cornell University Press, Ithaca, New York, 507 pp. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 90-96 PSEPHENOPALPUS BROWNI, A NEW GENUS AND SPECIES OF PSEPHENINAE (COLEOPTERA: BYRRHOIDEA: PSEPHENIDAE) FROM MEXICO ROBERTO ARCE-PEREZ Instituto de Ecologia, A.C. Km. 2.5 Antigua Carretera a Coatepec 351, Congregacién el Haya, Apartado Postal 63, 91070 Xalapa, Veracruz, México (e-mail: arcerob@ ecologia.edu.mx) Abstract.—Psephenopalpus browni, n. gen., n. sp. is described and illustrated from specimens collected in a stream running through a cloud forest in the Mexican town of Coatepec, Veracruz (1200 m). Psephenopalpus browni can be distinguished from other Psepheninae by the body shape, antennal length, and the size proportions of the phallobase and parameres (from Pheneps) and by the pronotum, maxillary palpi, and tarsi (from Psephenotarsis, Psephenops, Psephenus). Psephenopalpus appears to be more closely re- lated to Psephenops than to other Psepheninae. A key for American Psepheninae genera is provided. Resumen.—Se describe e ilustra a Psephenopalpus browni, n. gen. y n. sp. con base en especimenes colectados en un arroyo de agua dulce que atraviesa un bosque nebuloso en Coatepec (1200 m snm), estado de Veracruz, México. Psephenopalpus browni se puede diferenciar de otros Psepheninae por la forma del cuerpo, la longitud de las antenas, la proporcion entre la falobase y los parameros (Pheneps); también por la forma del pronoto, longitud y forma de los palpos maxilares y la regi6n tarsal (Psephenotarsis, Psephenops, Psephenus). Al parecer Psephenopalpus esta mas cercanamente relacionado a Psephenops que al resto de los Psepheninae. Se proporciona una clave para la identificacion de los géneros Americanos de Psepheninae. Key Words: Psephenidae, Psepheninae, new genus, description, key, State of Veracruz, Mexico The subfamily Psepheninae is represent- ed in the New World by the genera Psep- henus Haldeman 1853, Psephenops Grou- velle 1898, Pheneps Darlington 1936, and Psephenotarsis Arce-Pérez 2001 (Brown 1981, Spangler 1982, Arce-Pérez and No- velo-Gutiérrez 2001, Arce-Pérez and Shep- ard 2001). Psephenus contains 15 species with seven in North America, two of which are also found in Mexico; four in Mesoa- merica; three in Brazil; and one in Peru. Psephenops has eight species (Arce-Pérez and Novelo-Gutiérrez 2000, Bameul 2001, Arce-Pérez 2002) with a tropical distribu- tion, including two in Mexico, one in Gua- temala, one in Costa Rica, one in Haiti, one in the Antilles with one subspecies, one in Colombia that is also found in Costa Rica, and one in Argentina. Pheneps has eight species (Arce-Pérez and Novelo-Gutiérrez 2001, Souza Fernandez et al. 2001) also tropical, with one in Cuba, one in Haiti, one in Surinam, one in Venezuela also found in Brazil, and another four in Brazil. Psephe- VOLUME 106, NUMBER 1 notarsis includes one species in Mexico. In this paper, a new genus and new species are reported for Mexico. Twenty-six male specimens were collect- ed from a stream running through a cloud forest located in the Mexican town of Coa- tepec, Veracruz (alt. 1,200 m), municipality of Coatepec. Individuals were initially pre- served in 80% ethanol and some were pinned later. All collected material was ex- amined. Male genitalia were extracted in ethanol, and potassium hydroxide (KOH 5%) was used to clear genitalia to make il- lustrations. Microphotographs were taken with a JEOL scanning electron microscope, model JSM T20 with 200 magnification. Psephenopalpus Arce-Pérez, new genus (Figs. 1, 5, 7, 12-14) Description.—Head (Fig. 1) short, trans- verse; eyes spherical, very prominent. An- tenna long, filiform-subserrate, | 1-segment- ed, extending beyond elytral humerus; scape and pedicel yellowish, 9 flagellar seg- ments dark reddish brown. Maxillary pal- pus (Fig. 5) extremely long, extending be- yond sixth antennal segment, second pal- pomere much longer than the other palpo- meres. Pronotum (Fig. |) subtrapezoidal, its posterior margin quite smooth, never cren- ulate or rugose; lateral margins rounded, unexpanded at posterior middle, disc with feeble middorsal longitudinal carina; pro- sternum (Fig. 5) short, slightly elevated at middle, prosternal process long with sides parallel at middle, then gradually diverging to form a lanceolate apex; mesosternum clearly grooved and slightly bifurcate api- cally, so procoxa and mesocoxa not far apart. Tarsi 5-segmented (Fig. 7), first four tarsomeres short, subtriangular and covered with long setae, ventral vestiture spongy, fifth tarsomere much longer than first four together, cylindrical, slender, smooth; tarsal claws long, whole, curved, lacking denticle and accessory membranes. Male genitalia (Figs. 12, 13, 14): In dor- sal view (Fig. 12) subrectangular, trilobate; phallobase shorter than parameres; para- 9] meres shorter than median lobe, with exter- nal apical margin sinuate to 0.40, ending in a rounded apex, lateroapical portion mem- branous; (Fig. 13) middle lobe sclerotized, fingerlike in ventral view, (Fig. 14) dorsally convex, with sharp apex and curved in lat- eral view. Remarks.—Psephenopalpus can be dis- tinguished from other Psepheninae genera by the following characteristics: In Pheneps the body is more slender; the maxillary pal- pus is short; the antennae are long reaching 0.75 the length of elytra; the pronotum and elytra lack expanded margins (Fig. 2); tar- someres |—2 are clearly stouter than any of the three apical tarsomeres (Fig. 8); and the phallobase is notably larger than the para- meres. In Psephenops, the second maxillary palpomere is proportionally much shorter (Fig. 6); the pronotum is expanding later- ally on the posterior half (Fig. 3); and tar- someres | and 2, and occasionally 3, have wide ventral expansions that cover the api- cal tarsomeres (Fig. 9). In Psephenotarsis, the maxillary palpus is short; the antennae are short and moniliform; the pronotum is expanded laterally in the posterior half (Fig. 4); and the tarsomeres are notably triangu- lar, flattened and widely emarginate (Fig. 10). In Psephenus, the maxillary palpus is short; the basal half of the pronotum is oc- casionally expanded; the four basal tarso- meres are slender and subcylindrical and slightly emarginate in their apices, with del- icate, minute setae or papillae ventrally (Fig. 11); and the median lobe of the male genitalia is mostly membranous and shorter than or as long as the parameres. Psephenopalpus appears to be more closely related to Psephenops than other Psepheninae, mainly by the pronotal carina; filiform subserrate antennae; posterior mar- gins of sternites | and 2 slightly sinuate or emarginate; the basal tarsomeres with spongy ventral vestiture; and in general ap- pearance. The two may be distinguished by the following combination of characters (those of Psephenops in parentheses): Pro- notum with lateral margins rounded and 92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. 1, Psephenopalpus browni, adult male, dorsal aspect of head and pronotum. 2, Pheneps cursitatus Spangler, adult male, dorsal aspect. 3, Psephenops mexicanus Arce-Pérez and Novelo-Gutiérrez, adult male, dorsal aspect of head and pronotum. 4, Psephenotarsis triangularis Arce-Pérez and Novelo-Gutiérrez, adult male, dorsal aspect. 5, Psephenopalpus browni, adult male, ventral aspect of head. 6, Psephenops lupita Arce- Pérez, adult male, ventral aspect of head and prosternum. (Fig. 2 from Spangler 1987; Fig. 3 from Arce-Pérez and Novelo-Gutiérrez 2000; Fig. 4 from Arce-Pérez and Novelo-Gutiérrez 2001; Fig. 6 from Arce-Pérez 2002.) VOLUME 106, NUMBER 1 unexpanded (expanded laterally on poste- rior half); maxillary palpus extremely long, with the second palpomere much longer than the other palpomeres (maxillary palpus short, last palpomere larger); the basal tar- someres with short ventral expansions (tar- someres | and 2, and occasionally 3, with wide ventral expansions that cover the api- cal tarsomeres); and fifth tarsomere much longer than the first four together, cylindri- cal, slender and smooth (second tarsomere lobed ventrally, much longer than the other tarsomeres). Etymology.—From the Greek **Psephe- no” (hidden) and “‘palpus”’ (palpomeres), referring to the palpomere characteristic that distinguishes this genus from other Psepheninae. Psephenopalpus browni Arce-Pérez, new species Geraci eas. 125 13,14) Description.—Holotype male: Body oval tapered, depressed; integument reddish black, completely covered with dense ves- titure of short and long reddish yellow setae with golden highlights; venter with short yellowish gray setae with golden highlights. Head, pronotum, and scutellum black; an- tenna and elytron dark reddish brown, ely- tron without striae, with light yellow mark in mid basal region; legs with yellow coxae, trochanters, and femora; tibiae and tarsi dark reddish; total length 3.85 mm, maxi- mum humeral width 1.70 mm. Head (Fig. 1): Black, short, transverse, covered with short vestiture, yellowish se- tae and long dark reddish setae; eyes spher- ical, very prominent; clypeus subrectangu- lar, wider than its length, declivous at an angle of less than 90° from plane of frons, distal margin widely emarginate; labrum like clypeus but shorter; antenna long, fili- form subserrate, 11-segmented, extending past elytral humerus, scape and pedicel yel- lowish, 9 flagellar segments dark reddish; scape larger than all other segments, slight- ly curved, pedicel half as long as scape; fla- gellum with 9 slim, subserrate segments 93 covered with long reddish-black setae; maxillary palpus 4-segmented (Fig. 5), slightly longer than first six antennomeres combined, three basal palpomeres subcylin- drical, apical palpomere ovoid; first maxil- lary palpomere shortest, second 7.75 times longer than first and slightly sinuate, third palpomere slightly larger than second at middle; fourth slightly longer than third but ovoid and depressed with rounded apex and large sensorial region; relative proportion of segments: 0.07, 0.54, 0.31, 0.34; all pal- pomeres dark reddish and covered with red- dish setae; labial palpus very short (Fig. 5), 3-segmented, not extending beyond basal margin of first maxillary palpomere, two basal labial palpomeres rounded and wide, segment 3 subspherical and shorter; all pal- pomeres dark reddish and covered with red- dish setae. Thorax: Pronotum (Fig. 1) black and subtrapezoidal (posterior margin 1.55 mm, lateral margin 0.83 mm); anterior margin arcuate, posterior margin bisinuate and al- most as wide as elytral base; lateral margins rounded and unexpanded; posterolateral an- gles subacute; pronotal disc with mid-dorsal longitudinal carina weakly developed near base, slight depression at each side of ca- rina; in profile, pronotal disc convex; pros- ternum (Fig. 5) short and slightly elevated; long, concave, and lanceolate prosternal process reaching anterior half of mesoster- num. Mesonotum with scutellum short and black, with tip broadly rounded; elytron dark reddish brown with light-colored stain in mid-basal region, lacking striae, with short yellow setae and long reddish-yellow setae, sides parallel on basal 0.75, then gradually converging posteriorly and mar- gins expanding slightly; length from base to apex of suture 2.80 mm, width 1.70 mm across humerus. Mesosternum reddish black with wide groove for reception of prosternal process and slightly bifurcate at apex. Metasternum reddish black, large and voluminous, with longitudinal canal, deeper apically and ending in front of metacoxae. Legs (Fig. 7): Coxae, trochanters, and 94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON femora yellow; tibiae and tarsi dark reddish, almost black; procoxa globose, mesocoxa conical, metacoxa transverse; femora robust with dark reddish yellow setae, tibiae slen- der, protibia with distinct apical posterolat- eral denticle, metatibia longer than femur; tarsi 5-segmented, the first four tarsomeres short and subtriangular with slight ventral expansions and covered with long setae, ventrally with spongy vestiture; first tarso- mere covering half of second, vestiture of second tarsomere covering all or almost all of third, fourth tarsomere small and free or with sparse setae; fifth tarsomere much lon- ger than all others combined (0.32 mm compared to 0.44 mm), subcylindrical and smooth, tarsomere proportions 0.14, 0.20, 0.10, 0.10, 0.44 mm; claws long, curved, without denticles or accessory membranes. Abdomen: Ventrally convex, with seven sternites; first sternite slightly lifted in mid- basal region, second and third sternites flat in middle; posterior margin of sternites | and 2 slightly sinuate in middle; sixth ster- nite short, widely emarginate and hidden under fifth sternite; seventh sternite much smaller and oval in shape; pygidial plate oval, convex, with long, dark reddish setae. Genitalia (Figs. 12, 13, 14): Trilobate, total length 0.66 mm. Parameres in dorsal view (Fig. 12) voluminous, shorter than middle lobe, with external margin sinuated at apical 0.40, ending in a rounded apex with lateroapical membrane, length. 0.36 mm; penis in ventral view (Fig. 13) longer than parameres, fingerlike, widening at bas- al half, length 0.40 mm, with robust lon- gitudinal sclerite sublanceolate in shape, in lateral view (Fig. 14) dorsally convex, with curved sharp apex; dorsal area of phallo- base concave (Fig. 12); ventrally volumi- nous, subcylindrical; length 0.26 mm. Female: Unknown (females of this spe- cies may be larger and bulkier than males but less frequently collected). Variation in paratype series.—Some specimens have dark coxae, other speci- mens have a yellowish ventral region, mainly on the metasternum. Measurements (in mm): total body length 3.10—4; maxi- mum width 1.45—1.77; pronotal length 0.75—0.90, basal width 1.25—1.55; elytral length 2.67—2.95. Type material.—Holotype d labeled: “MEXICO, Veracruz, Municipality Coate- pec, Coatepec, Huehueyapan River, 1,200 m, mountainous cloud forest, 1-V-2000, Roberto Arce col.’ 7 ¢ paratypes, same data as holotype; 18 d paratypes, same as holotype except for, four 6-V-2000; five 19- TV-2000; one 29-III-2001, and eight 17-III- 2002. Holotype and paratypes deposited at Coleccion Entomoldgica, Instituto de Eco- logia, A. C., Xalapa, Veracruz, México (IEXA). Etymology.—It is with pleasure that I dedicate this species to Professor Harley P. Brown of the University of Oklahoma be- cause of his contributions to research and knowledge of American Psepheninae. Habitat.—Adult males were captured in a quick-flowing stream, on stones covered with moss and aquatic phanerogams, and near Psephenops lupita Arce-Pérez individ- uals. KEY TO NEw WORLD GENERA OF PSEPHENINAE BASED ON ADULTS [Modified from Brown (1976) and White and Brigham (1996)] 1. Antenna long, filiform, extending notably beyond elytral humeri (Fig. 2); pro- and me- sotarsomeres | and 2 markedly stouter than following three and slightly emarginate at apex (Big. 8) 9 hae caer ae eerie aa ee Pheneps - Antenna short, moniliform or subserrate, scarcely or not extending beyond elytral hu- meri; pro- and mesotarsomeres | and 2 of variable length but never stouter than follow- ONES (OVREE) TIBIANOVOMNEMES 3505000 0cc0g0006 2 2(1') Pronotum with a mid dorsal longitudinal ca- rina (Fage 3)! s:$, Sestacwes eae ees 3 Pronotum! without cannae eee ee 4 3(2) Pronotum expanded laterally on posterior half, with a well-developed carina (Fig. 3); maxillary palpus short, last palpomere lon- gest (Fig. 6); tarsomeres | and 2, and oc- casionally 3, with wide ventral expansions that cover apical tarsomeres (Fig. 9)..... MS Sa ee Ee ey ee ee Psephenops = Pronotum with lateral margins rounded, not VOLUME 106, NUMBER | 9 nN Figs. 7-14. 7, Psephenopalpus browni,adult male, tarsus showing extreme length of fifth tarsomere. 8, Pheneps antennalis Spangler and Steiner, adult male, protarsal segments. 9, Psephenops mexicanus, adult male, showing tarsomeres with wide expansions. 10, Psephenotarsis triangularis, adult male showing emarginate tarsomeres 1—4. 11, Psephenus sp., adult male, tarsus showing relatively slender tarsomeres 1—4. 12, Psephen- opalpus browni, male genitalia, dorsal aspect. 13, Psephenopalpus browni, male genitalia, ventral aspect. 14, Psephenopalpus browni, male genitalia, lateral aspect. (Fig. 8 from Spangler and Steiner 1983; Figs. 9-11 from Arce-Pérez and Novelo-Gutiérrez 2001.) expanded on posterior half, with a feeble deeply emarginate for reception of follow- carina (Fig. 1); maxillary palpus extremely ing tarsomere; tarsomeres I—2 with long long, second palpomere much longer than ventral setae or papillae, those on 2 reaching other palpomeres (Fig. 5); tarsomeres with- base of fifth tarsomere (Fig. 10)...... OlEmideaventrallexpan sions mittONtanSOUIELes | sy yy) uct tee eii Tei iewene arate circa Psephenotarsis much longer than first four together (Fig. 7) = Tarsomeres 1—4 subconical and _ slender, Oe aoa he gheat duty sh sis Psephenopalpus, gen. n. scarcely emarginate at apex; tarsomeres 1— 4(2') Tarsomeres 1—4 short, triangular, flattened, 2 usually with ventral setae or papillae that 96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON never cover contiguous tarsomere (Fig. 11) 3s, yas sl emhenoretoank seawater s. o3 Psephenus ACKNOWLEDGMENTS Thanks are due Dr. Miguel A. Moron (Xalapa, Mexico), Dr. Manfred Jach (Vi- enna, Austria), and Dr. Harley P. Brown (Oklahoma, USA) for their invaluable crit- icism on the final manuscript as well as for the English corrections. Finally, special thanks to Tiburcio Laez Aponte and Manuel Escamilla (Xalapa, Mexico) for the elabo- ration of microphotographs and drawings. LITERATURE CITED Arce-Pérez, R. 2002. A new species of Psephenops Grouvelle (Coleoptera: Psephenidae) from Mexi- co. Proceedings of the Entomological Society of Washington 104(4): 964—967. Arce-Pérez, R. and R. Novelo-Gutiérrez. 2000. First record of the genus Psephenops (Coleoptera: Psephenidae) from Mexico, with a description of a new species. Entomological News 111(3): 196— 200. . 2001. A new genus and species of Psepheni- nae (Coleoptera: Dryopoidea: Psephenidae) for Mexico. Proceedings of the Entomological Soci- ety of Washington 103(2): 389-395. Arce-Pérez, R. and W. D. Shepard. 2001. Sinopsis de la familia Psephenidae (Coleoptera: Dryopoidea) de Norte y Centroamérica. Folia Entomol6ogica Mexicana 40(3): 397—406. Bameul, EF 2001. Un nouveau Psephenops Grouvelle de la Guadeloupe, avec la description de larve (Coleoptera, Psephenidae). Nouvelle Revue d’ En- tomologie (N.S.) 18(2): 161-172. Brown, H. P. 1976. Aquatic dryopoid beetles (Cole- optera) of the United States. United States Envi- ronmental Protection Agency. Water Pollution Control Research Series 18050/72 (Second Print- ing). Aquatic Biology Section. Cincinnati, Ohio, 82 pp. . 1981. A distributional survey of the world genera of aquatic dryopoid beetles (Coleoptera: Dryopidae, Elmidae, and Psephenidae sens. lat.). The Pan-Pacific Entomologist 57(1): 133-148. Souza Fernandez, M. F, C. R. Vasconcelos da Fonseca, and P. J. Spangler. 2001. Estudo Taxionémico dos Psephenidae (Coleoptera: Byrrhoidea) da Ama- zonia Brasileira. Acta Amazonica 31(3): 469—500. Spangler, P. J. 1982. Coleoptera, pp. 328-397. Jn Hurl- bert, S. H. and A. Villalobos-Figueroa, eds. Aquatic Biota of Mexico, Central America and the West Indies. San Diego. University of San Diego, California, 529 pp. . 1987. A new species of water penny beetle, Pheneps cursitatus, from Cerro de la Neblina, Venezuela (Coleoptera: Dryopoidea: Psepheni- dae). Proceedings of the Entomological Society of Washington 89(2): 219-225. Spangler, P. J. and W. E. Steiner. 1983. New species of water beetles of the genera Elmoparnus and Pheneps from Suriname (Coleoptera: Dryopoidea: Psephenidae). Proceedings of the Entomological Society of Washington 85(4): 826-839. White, D. S. and W. V. Brigham. 1996. Aquatic Co- leoptera, pp. 399-473. In Merritt, R. W. and K. W. Cummins, eds. An Introduction to the Aquatic Insects of North America. Third Edition, Kendall/ Hunt Publishing Company, Dubuque, Iowa. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 97-101 A NEW SPECIES OF THE ANT GENUS ACANTHOSTICHUS MAYR (HYMENOPTERA: FORMICIDAE) FROM PARAGUAY, AND A DESCRIPTION OF THE GYNE OF A. BREVICORNIS EMERY WILLIAM P. MACKAY Centennial Museum, Department of Biological Sciences, The University of Texas, El Paso, TX 79968, U.S.A. (e-mail: wmackay @utep.edu) Abstract.—l| describe a new species of Acanthostichus from the Department of Chaco in Paraguay. Acanthostichus longinodis, described from the worker, is a member of the brevicornis species complex. It differs from the workers of all of the other South American species in having an elongate petiole, a relatively small subpetiolar process and a swollen posterior femur. I also describe the gyne of A. brevicornis Emery and provide new in- formation on the worker. Resumen.—Describo una nueva especie del género Acanthostichus encontrada en el departamento de Chaco, Paraguay. Acanthostichus longinodis es descrita basada en la obrera, y es miembro del complejo de especies brevicornis. Se diferencia en que el pecfolo es muy largo, el proceso subpetiolar es relativemente pequeno, y el fémur posterior es ancho. También describo la hembra de A. brevicornis Emery y presento nueva informacion de la obrera de esta especie. Key Words: The ant genus Acanthostichus Mayr is an interesting, rarely-collected group of ants. They are probably common, but rarely en- countered due to their subterranean habits. Most worker specimens are collected under stones or other objects, while males are at- tracted to lights. The genus is found throughout the Americas, from the southern part of the United States south to Paraguay, Uruguay, and northern Argentina, and re- cently has been revised (Mackay 1996). Acanthostichus longinodis Mackay, new species (Figs. 1—4, 10) Diagnosis.—This initially appears to be a typical South American Acanthostichus species, but can be distinguished from other South American species by the combination of the elongate petiole (Fig. 4), the small Neotropical, Cerapachyinae, Chaco region subpetiolar process (Fig. |) and the incras- sate posterior femur (Fig. 3). It can be sep- arated from two Mexican species with elon- gate petioles, A. quirozi Mackay and A. skwarrae Wheeler as the sides of the petiole are sharply separated from the dorsal sur- face by an abrupt bend which nearly forms a carina. The elongate petiole would pre- clude confusion with any other species in the genus. Description.—Worker (n = 3, all mea- surements in mm): HL (Head Length) 0.76—0.78, HW (Head Width) 0.58—0.60, SL (Scape Length) 0.28—0.30, SW (maxi- mum Scape Width) 0.10—0.11, EL (maxi- mum Eye Length) 0.04—0.05, WL (Weber’s Length, anterior edge of pronotal shoulder to posterior edge of metapleural lobe) 1.08— 1.14, PL (Petiole Length) 0.35—0.40, PW (Petiole Width) 0.23—0.26, SI (Scape index, 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SL/HL X 100) 35-40, CI (Cephalic Index, HW/HL xX 100) 76—77, PI (Petiolar Index, PL/PW X 100) 150-153, SL/SW 2.78- 2.88. Mandible without teeth; anterior border of clypeus weakly concave, nearly straight; frontal carinae closely spaced, slightly overhanging anterior margin of clypeus with head in full face view (Fig. 2); sides of head nearly straight, slightly widened an- teriorly (Fig. 2), posterior border of head slightly concave; scape short; eye tiny, ap- parently composed of three ommatidia; ma- lar groove well marked; mesosoma basical- ly rectangular in shape as seen from side and as seen from above, narrowed posteri- orly (seen from above), with narrowest part being posterior edge of propodeum; pro- podeal spiracle slightly below midline (Fig. 1); petiole greatly elongated (Fig. 4), ante- rior face nearly straight, slightly widened posteriorly; posterior femur incrassate (Fig. 3). Moderately hairy, with erect hairs on scape, mandible, dorsal surface of head, posterior border of head, ventral surface of head, dorsum of mesosoma, dorsum of pet- iole, and all surfaces of gaster. All surfaces smooth and glossy. Ferrugineous red, appendages and gaster slightly lighter. Distribution.—Known only from the type locality (Fig. 10). Type series.—Holotype worker (MNHP, Museo Nacional de Historia Natural del Paraguay), 2 paratype workers (CWEM, William and Emma Mackay collection; MCZC, Museum of Comparative Zoology, Harvard University), PARAGUAY: Dpto. Chaco: Parque Nacional Defensores del Chaco: Cerro Leon: zona pelada: Sitio #11; 9—15-viii-1983, Colr. J. A. Kochalka. Material examined.—Type series. Etymology.—From Latin, /ongus for long and nodus for swelling, describing the elongate petiole. Discussion.—This species would not be confused with any other South American species, based on the small subpetiolar pro- cess, the incrassate posterior femur, and the elongate petiole. The elongate petiole could cause confusion with two species from Mexico: A. guirozi and A. skwarrae. How- ever, it can be separated from the geograph- ically distant Mexican species as the sides of the petiole are sharply separated from the dorsal surface by an abrupt bend which nearly forms a carina. A modified key to include this new spe- cies will be placed at www.utep.edu/leb/ antgenera.htm. Biology.—The type series was collected under a stone. Acanthostichus brevicornis Emery (Figs. 5—9, 10) Acanthostichus brevicornis Emery 1894: 142, worker, French Guiana. This is one of the most common species in the genus, being previously found in Bra- zil, Argentina, and French Guiana. I report the first record from Paraguay, provide new information on the worker, and describe the gyne. Description.—Worker: Additional mea- surements (n = 3): HL 0O.71-1.07, HW 0.56—0.82, SL 0.30-—0.40, SW 0.11-0.18, EL 0.04—0.06, WL 0.92-1.46, PL 0.29- 0.47, PW 0.29-0.46, SI 37—42, CI 76-90, PI 100-106, SL/SW 2.20-2.73. These workers from Paraguay differ from the ‘“‘typical’’ A. brevicornis in being strongly dimorphic, increasing the range of worker size (above). Additionally the an- terior face of the petiole is nearly straight as seen from above. The shape of the an- terior face of the petiole in this species is variable, ranging from nearly straight to strongly concave. Gyne: HL 1.02, HW 1.01, SL 0.43, SW 0.19; EL 0.18; WE 167, PE O62 5b Wvelisor SI 42, Cl 99: P50) SE/SWV-2.-25) Subdichthadiiform, mandible without teeth (Fig. 6); anterior border of clypeus concave (Fig. 5); frontal carinae closely spaced, posterior border of head concave, sides of head rounded; malar groove poorly VOLUME 106, NUMBER 1 99 A. longinodis, lateral view. 2, Head of A. longinodis, full face view. 3, Posterior left femur of A. longinodis, front view. 4, Petiole of A. longinodis, dorsal view; arrow points anteriorly. 5, Head of A. brevicornis, full face view. 6, Mandible of A. brevicornis, front view. 7, Posterior left femur of A. brevicornis, front view. 8, Petiole of A. brevicornis, dorsal view. 9, Mesosoma and petiole of A. brevicornis, lateral view. developed, extending from base of mandi- ble slightly more than 0.1 mm, remainder of groove marked by very slight indenta- tion; scape thickened; eye large, but failing to reach side of head by about % minimum diameter; ocelli absent; mesosoma basically rectangular as seen from side (Fig. 9), and as seen from above; promesonotal suture poorly developed, metanotal suture marked on dorsum of mesosoma; propodeal spiracle placed above midline (Fig. 9), propodeum rounded between faces; subpetiolar process absent (Fig. 9), petiole wider than long as seen from above, slightly wider posteriorly (Fig. 8); posterior femur slightly widened (Fig. 7), much less so than that of worker; pygidium without spines. Erect hairs sparse; few hairs along ante- rior border of clypeus, and on mandible, along frontal carina, dorsum of head, pos- terior margin of head, dorsum of mesoso- ma, and legs; hairs on posterior face of pro- podeum fine, brushlike, with similar hairs on side and posterior margin of petiole: ventral surface of petiole with thick, closely spaced hairs; gaster with similar hairs on dorsal and ventral surfaces. All surfaces smooth and glossy. Yellow, mandibles and clypeus slightly darker. 100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 10. squares of A. brevicornis are from Mackay (1996). Material examined.—PARAGUAY, Can- indeyu: Col. “‘11 de Septiembre’’, 24°03’S 55°34'W, 19.vii.1997, A. Wild #AWO0S522 (7 workers, CWEM, MCZC; 1 2, MCZC). The full series consists of 43 workers [ma- jority not seen]. Specimens are also depos- ited in the LACM (Natural History Muse- um of Los Angeles County), MNHP, in the collection of Sean Brady, and in the collec- tion of Alex Wild. Discussion.—The gynes of this genus are @ A. longinodis @ A. brevicornis Distribution of A. /onginodis and the new locality of A. brevicornis in South America. The open known from only five species: A. emmae Mackay (fexanus species complex) and A. texanus Forel (texanus species complex), which are “‘normal’’, winged gynes, with ocelli, and A. laticornis Forel (serratulus species complex), A. guadratus Emery (ser- ratulus species complex) and now A. brev- icornis Emery (brevicornis species com- plex), which are subdichthadiiform (lacking ocelli, lacking flight sclerites, having a dis- tended gaster, and lacking the teeth on the VOLUME 106, NUMBER 1 pygidium). The gyne of A. brevicornis eas- ily can be separated from that of A. gua- dratus, as the malar groove is developed at least near the base of the mandible (appar- ently lacking in A. quadratus), the propo- deum is rounded between the faces (with an angular process in A. quadratus and a carina in A. laticornis) and the subpetiolar process is absent (developed into a tooth in A. quadratus and a broad flange in A. Jati- cornis). Separation from A. laticornis is more difficult. The gyne of A. brevicornis is smaller than that of A. laticornis. The to- tal length of the gyne of A. brevicornis is less than 9 mm, the gyne of A. laticornis is about 11 mm. The malar groove in the gyne of A. brevicornis is relatively short, extend- ing slightly more than 0.1 mm from the point of insertion of the mandible (extend- ing about half of the distance to the eye of A. laticornis or 0.4 mm). The lateral clypeal angles are absent in A. brevicornis (convex border is present), but are well developed in A. laticornis. The frontal carinae of A. brevicornis are very closely placed, with the distance between the external margins 0.23 mm (about 0.4 mm in A. quadratus, 0.47 in A. laticornis). This suggests an ad- ditional characteristic to separate the two species complexes. The frontal carinae of the gynes of the brevicornis complex may be closely spaced, those of the serratulus 101 species group are apparently more widely spaced. As gynes have not been collected without workers, the worker characteristics would allow separation of the gynes of the two species. Since the sample size of gynes is so limited, we have very little sense of intraspecific variation, which is really need- ed to establish how the species differ. Biology.—The series was nesting in the soil in grazed second growth vegetation. This species is not uncommon in that col- lection locality. Alex Wild (personal com- munication) saw it several times under logs and near the surface of the soil where he was digging. It was always found in dis- turbed habitats, e.g., grazed areas, lawns, nesting in red clay soil. ACKNOWLEDGMENTS I thank Alex Wild, for sending the spec- imens. He was supported by the United States Peace Corps in Paraguay. Alex Wild and Emma Mackay critically reviewed the manuscript. Publication was supported by the National Institutes of Health, Grant #5 G12RR08 124. LITERATURE CITED Emery, C. 1894. Studi sulle formiche della fauna Neo- tropica. Bullettino della Societa Entomologica It- alaliana 26: 137—242 + 4 plates. Mackay, W. P. 1996. A revision of the ant genus Acan- thostichus (Hymenoptera: Formicidae). Sociobi- ology 27: 129-179. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 102-109 A NEW FOSSIL NICOLETIIDAE (ZYGENTOMA, “APTERYGOTA’’) IN DOMINICAN AMBER Luis E MENDES AND GEORGE O. POINAR, JR. (LFM) Centro de Zoologia, Instuto de Investigagao Cientifica Tropical, R. da Junqueira, 14, 1349-007 Lisboa, Portugal (e-mail: czool@ckiict.pt); (GOP) Department of Zoology, Oregon State University, Corvallis, OR 97331-2907, U.S.A. (e-mail: poinarg@casco. net) Abstract.—Hemitrinemura exstincta, n. sp., is described from Dominican amber (ca. 15—45 Ma). The new species is considered to be the only Neotropical representative of a genus with (?) three living species known from Papua New Guinea, Melanesia, and Pol- ynesia. This species is the third Nicoletiidae fossil from New World amber; it is easily distinguishable from previously described taxa (two species of Trinemurodes Silvestri) by the complete praetarsus. Key Words: Zygentoma Several specimens of Microcoryphia (= Archaeognatha) and Zygentoma (= Thy- sanura s. Ss.) have been described from Do- minican amber. These include a single spe- cies of Meinertellidae (Microcoryphia), Neomachilellus (Praeneomachilellus) dom- inicanus Sturm and Poinar (1997) repre- sented by a large number (ca. 100) of spec- imens, 3 specimens of Ctenolepisma elec- trans Mendes (1998) in the Lepismatidae (scaled and with lateral compound eyes), 11 specimens of Archeatelura sturmi Mendes (1997a, b) in the Ateluridae (eyeless, usu- ally scaled, with short, droplike body and as arule short appendages), and two species of Nicoletiidae (eyeless, usually unscaled, with thin elongate body and with long ap- pendages), including one individual each of Trinemurodes antiquus Sturm and Mendes (1998) and 7. miocenicus Sturm and Men- des (1998). A myrmecophilic relationship was suggested with A. sturmi since ants were also present in one of the amber pieces of that series (Mendes 1997b). The other biogeography, Dominican amber, Hemitrinemura exstincta, Nicoletiidae, fossil taxa probably had edaphic (soil and/ or soil surface) habits similar to their extant relatives. The present fossil specimen is readily distinguishable from Trinemurodes Silves- tri, the only previous genus of the Nicole- tiidae described from Dominican amber, by its complete praetarsus with well-developed paired claws and empodium. The occur- rence of this new fossil species is quite in- teresting not only from the systematic point of view but also because of its geographical origin and the distribution of its closest liv- ing relatives. MATERIALS AND METHODS The specimen was obtained from mines in the Cordillera Septentrional of the Do- minican Republic. Dating Dominican am- ber is still controversial with the latest pro- posed age of 20—15 mya based on Fora- minifera (Iturralde-Vincent and MacPhee 1969) and the earliest as 45—30 mya based on coccoliths (Cépek in Schlee 1990). A VOLUME 106, NUMBER 1 range of ages for Dominican amber may be likely since the amber fossils are associated with turbiditic sandstones of the Upper Eo- cene to Lower Miocene Mamey Group (Draper et al. 1994). Dominican amber can be secondarily deposited in sedimentary rocks, which makes a definite age deter- mination difficult (Poinar and Mastalerz 2000). Dominican amber was produced by the leguminous tree, Hymenaea _ protera Poinar (1991). A re-construction of the Do- minican amber forest based on amber fos- sils indicated that the climate was similar to that of a present-day tropical moist forest (Poinar and Poinar 1999). The amber piece containing the fossil was reshaped and pol- ished in order to better view the specimen. Examination was made under magnifica- tions of up to 750, with a combination of direct illumination (optical tubes of Hund Weizler apparatus) and indirect lighting (with a Leitz SM-Lux optical microscope and a Wild MSA stereoscopic microscope). Also included in the same amber piece with the nicoletiid were various sized pieces of plant material. The holotype and only spec- imen is deposited in the Poinar Amber Col- lection maintained at Oregon State Univer- sity (accession # T-1-13). DESCRIPTION Hemitrinemura exstincta Mendes and Poinar, new species (Figs. 1-16) Diagnosis.—The new species possesses a long, thin, parallel-sided body with only se- tae and macrochaetae (scales lacking), a de- veloped prostheca, simple and complete praetarsi with long, delicate, smooth claws; thin tibiae with some long, stout, spiniform ventral setae; a short and wide urotergite X; urosternites I (?)—VII entire, II-IX stylets and II—VII vesicular structures; a subgenital plate wider than long; a short typical ovi- positor with thin setae only and a spinulated inner ventral area of gonapophyses LX. The new species is distinguished from other species of the genus by the degree of ap- 103 pendage elongation. It is intermediate be- tween the very elongated H. subarmata (Paclt), from Papua New Guinea (Paclt 1982), and the short, stout forms of H. pa- cifica (Carpenter) from Samoa (Polynesia) and Vanuatu (Melanesia) archipelagos and H. gracilis (Carpenter) from Western Sa- moa (Carpenter 1928). Description.—The basal ventral portion of the abdomen is damaged and the body is twisted in a U shape position. Some air bubbles and minute opaque particles occur on the dorsolateral surface at the end of the abdomen, at the middle dorsal area of the thorax and along the body. Some areas are either deformed or difficult to study in de- tail due to the position of the specimen. Body length: 4.1 mm; antennae length: maximum of 3.1 mm; cerci length: 2.3 mm; paracercus length (apically damaged): 2.4 mm. Body (Figs. 1—4): Thin and elongate, delicate, lacking scales; setae and macro- chaetae present, some near the border of the sclerites; legs long and stout, with acute or round points. Head (Figs. 4—9): Almost certainly wid- er than long (oblique) with several macro- chaetae mainly along lateral and posterior borders; antenna typical, scapus subcylindr- ical, ca. twice as long as wide, pedicellus not much longer than wide; flagellum with ovoid articles, each with a ring of thin setae, those of basal area round, apical ones clear- ly longer than wide; mandible large, apex not visible in detail; maxilla with prostheca, lacinial teeth acute, dark; galea ?; maxillary palp with subcylindrical articles, apical one with 6 papillae, ca. .25 longer than preced- ing and ca. 3.2 times longer than wide; la- bial posterolateral area not visible, labial palp (only one preserved) close to amber surface, impossible to see in detail, its api- cal article apparently ovoid or trianguloid and more or less as long as wide, with usual papillae. Thorax (Figs. 10-11): Longer than wide, with several marginal macrochaetae and some smaller discal setae and macrochae- 104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. Hemitrinemura exstincta, holotype. 1, Dorsal view, scale bar: 860 wm. 2, Ventral view, scale bar: 860 um. 3, Detail of abdomen, scale bar: 480 jm. VOLUME 106, NUMBER 1 Figs. 4-9. the air bubbles and granules in broken lines. 5, Head, dorsal. 6, Scapus and pedicellus of antenna. 7, Median flagellar chain. 8, Distal flagellar chain. 9, Maxillary palp. Scale bar: 0.1 mm. Abbreviations: a, antenna; g, galea; 1, lacinia; md, mandible; mp, maxillary palp; mx, maxilla. tae; leg I and leg II of only left side present and exclusively first one complete (right leg I and leg II and both leg III missing, only coxae preserved); femur of leg I and leg II elongate, thin (almost certainly distorted), tibiae very delicate, with short setae and 8 Hemitrinemura exstincta, holotype. 4, Ventral surface; damaged area of basal abdomen dotted, ventral macrochaetae; tibia I ca. 4% shorter than tibia II; tarsi with 4 divisions, with some strong ventral setae, praetarsus simple and complete, with long, thin lateral claws and delicate empodium (similar to claws though smaller, thinner and less arched): 106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tibia I, 0.16 mm long, clearly shorter than tarsus, tibia II, 0.26 mm, as long as tarsus. Abdomen (Figs. 12—16): Tergites similar to nota, with short setae and some posterior elongate macrochaetae (3+3 on posterolat- eral angle of urotergites I-VIII); urotergite X very short and wide, with almost straight posterior border (macrochaetae not pre- served); urosternite I polished away from amber piece, | 1—VII entire; H—VI with ves- icles, VII with pseudovesicles; H—IX with short, delicate, stylets; urostylet IX longer and stouter (ca. 0.20 mm) than urosternite VIII (ca. 0.14 mm), with some spiniform dark, long setae more conspicuous on IX; setae and macrochaetae of urosternites im- possible to observe; subgenital plate ellip- tical, shorter than wide at base; ovipositor short, with 8—10 divisions and with thin se- tae only, preserved in distorted position; in- ner ventral area of gonapophyses IX with spinulated integument; cercus and paracer- cus elongate, robust, clearly longer than half body length, with short setae and strong, elongate, macrochaetae. Etymology.—From the Latin exstinctus (a) (um), killed, disappeared, suppressed, extinct, in allusion to its fossil existence. Discussion.—The new taxon clearly shows a developed prostheca and subgenital plate and distally transformed gonapophys- es IX integument, which removes it from the Protrinemurinae. The presence of entire urosternites II—VII excludes it from the Ni- coletiinae as well as the Cubacubaninae. Unfortunately, the first urosternite is dam- aged, which makes it impossible to state ac- curately if this sclerite is entire (as occurs with the Subnicoletiinae) or if it is com- posed of one median sternite plus 1+1 cox- ites (as typical of the Coletiniinae) (Mendes 1988, 1994). The occurrence of 8 pairs of abdominal stylets (II-IX) and vesicular structures on urosternites II—VII is typical for the type genus, Coletinia Wygodzinsky 1980. How- ever, this predominantly northern Mediter- ranean genus possesses a much longer ovi- positor (exceeding the length of the IX sty- lets), a longer and more or less emarginated X urotergite and much more robust tibiae. Furthermore, the male paramera of Coleti- nia are entire and the ventral surface of X urotergite bear typical pegs, in contrast to the condition in Subnicoletiidae. In spite of the damaged urosternite I and absence of secondary sexual characteristics, we believe the Dominican amber fossil belongs to the Subnicoletiinae. The Subnicoletiinae (based on Mendes 1988, 1994 and Smith 1998) consists of 10 genera with a typical Gondwanian distri- bution known exclusively from the South- ern continents. The only genus in the family with living and fossil representatives (Sturm and Mendes 1998) is Trinemurodes Silvestri 1916, recognized by the absence of empodia. A lepismatoid-shaped body is also shared by other genera tentatively placed in this subfamily, including Hema- Escherich 1906, with modified scales, 4—6 pairs of vesicles and a Gond- wanian distribution, 7richatelura Silvestri 1932, from the Neotropics, with 3 pairs of stylets, Trichotriura Silvestri 1918 and Tri- chotriuroides Mendes et al. 1994 from Western Central Africa, with a reduced number of stylets (also on VII-IX only) and distinct tergal chaetotaxy. Subnicoletia Silvestri 1908a, a poorly de- scribed monotypical genus from the Guinea Gulf island of Sao Tomé (Sao Tomé a Prin- cipe Republic), has the stylets from uros- ternites IV—IX reduced to 6 pairs. The Aus- tralian genera Trinemura Silvestri 1908b and Subtrinemura Smith 1998, though much more similar to the fossil, show ab- dominal stylets on the urosternites I[I—-IX (lacking on II) as does Metrinura Mendes 1994, known from Australia and Celebes (= Sulawesi) with one isolated species in Colombia. Hemitrinemura Mendes 1994 is the only described genus that shares similar characters with the new Dominican fossil species, namely a long, thin unscaled body, entire urosternites (from I—VII), abdominal stylets on II-IX, vesicles on II-VI, pseu- telura VOLUME 106, NUMBER 1 107 Figs. 10-16. dovesicles on VII, and a complete praetar- sus, with paired claws plus empodium. According to Smith (1998), Hemitrine- mura includes not only the type species, H. subarmata (Paclt), from Papua New Guinea (Paclt 1982), but also two other species considered as possibly conspecific (Smith 1998), namely H. pacifica (Carpenter) from Samoa (Polynesia) and Vanuatu (Melane- sia) archipelagos and H. gracilis (Carpen- ter) from Western Samoa (Carpenter 1928). As previously reported (Smith 1998) the Pacific representatives show “‘antennae much shorter than body total length” and Hemitrinemura exstincta, holotype. 10, Left leg I, ventral. 11, Left leg Il, ventral. 12, Dorsal posterior abdomen. 13, Right cercus, dorsal, basal area. 14, Right cercus, distal area. 15, Terminal filament dorsal, basal area. 16, Terminal filament, distal area. Scale bar: 0.1 mm. Abbreviations: c, cercus; ov, Ovipositor; s. VIII, stylet VIII; s.LX, stylet LX; tf, terminal filament. “legs short and stout,’’ which seems to in- dicate a much closer similarity between H. subarmata (Paclt) and H. exstincta sp.n. In- deed, both species show quite thin and elongate legs and, despite the incomplete flagellar chain in the Dominican fossil, thin and elongate antennae. Hemitrinemura su- barmata, probably a troglobiont, is known only by males (Paclt 1982, Smith 1998). While the newly described species consists of one unique female; the New Guinea specimens are much larger (body + head ca. 10 mm), show longer and thinner max- illary palp articles and lack strong elongate 108 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON spines on the ventral surface of the legs. Despite possible shrinking resulting from amber preservation and even taking into ac- count that both males of H. subarmata are almost certainly somewhat distorted by de- hydration as reported by both Paclt (1982) and Smith (1998), the dissimilar characters show that H. exstincta is distinct from the New Guinea species. This is one of the few Dominican amber fossils with extant relatives having an Indo- West Pacific distribution. A Dominican am- ber marine water strider, Halovelia electro- dominica Andersen and Poinar (1998) also had a similar disjunct distribution with ex- tant members of the genus occurring in the Indo-West Pacific Region. Discovering a species of Hemitrinemura in Dominican amber suggests that the genus had a much more extensive distribution, including trans-Pacific, in the mid-Tertiary. The ex- tinction of H. exstincta from the New World could have been the result of climatic changes associated with cooling events that occurred in the Caribbean during the Plio- cene-Pleistocene periods, a condition that probably resulted in the extinction of many clades found in Dominican amber (Poinar and Poinar 1999). LITERATURE CITED Anderson, N. M. and G. O. Poinar, Jr. 1998. A marine water strider (Hemiptera: Veliidae) from Domini- can amber. Entomologica Scandanavica 29: 1—9. Carpenter, G. H. 1928. Apterygota. Insects of Samoa and other Terrestrial Arthropods. Part VII 3: 109— 116. Draper, G., P. Mann, and J. K Lewis. 1994. Hispaniola, pp. 129-150. In Donovan, S. and T. A. Jackson, eds. Caribbean Geology: An Introduction, The University of the West Indies Publisher’s Associ- ation, Kingston, Jamaica. 280 pp. Escherich, K. 1906. Beitrage zur Kenntnis der Thy- sanuren. 11 Reithe. Zoologische Anzeiger 30: 737-749. Iturralde-Vincent, M. A. and R. D. E. MacPhee. 1996. Age and palaeogeographical origin of Dominican amber. Science 273: 1850-1852. Mendes, L. EK 1988. Sur deux nouvelles Nicoletiidae (Zygentoma) cavenicoles de Grece et de Turquie et remarques sur la systématique de la famille. Re- vue Suisse de Zoologie 95: 751-772. . 1994. Evolutionary relationships among the Nicoletiidae (Insecta, Zygentoma). Acta Zoologi- ca Fennica 195: 98-103. . 1997a. First contribution to the study of the Dominican amber Zygentoma (Insecta). Family Ateluridae. Pedobiologia 41: 40—43. . 1997b. Some Zygentoma (Nicoletiidae, Ate- luridae) from the Neotropics with description of one new Metrinura species. Revue Suisse de Zoologie 104: 795—806. . 1998. Second contribution to the study of the Dominican Republic amber Zygentoma (Insecta): Family Lepismatidae. The Canadian Entomologist 130: 899-904. Mendes, L. FE, C. Bach de Roca, M. Gaju-Ricart, and R. Molero-Baltanas. 1994. Trichotriuroides boneti gen. n. sp. n. (Zygentoma, Nicoletiidae) and new data on Zygentoma in the collection of the Museo Nacional de Ciencias Naturales in Madrid (Spain). Eos 61: 21-29. Paclt, J. 1982. On some Solomon islands, Papua New Guinea and Sarawak Thysanura. Annotationes Zoologicae et Botanicae, Bratislava 151: 1—10. Poinar, Jr., G. O. 1991. Hymenaea protera sp. n. (Le- guminosae: Caesalpinoidea) from Dominican am- ber has African affinities. Experientia 47: 1075— 1082. Poinar, Jr., G. O. and R. Poinar. 1999. The Amber For- est. Princeton University Press. Princeton, New Jersey, 239 pp. Poinar, Jr., G. O. and M. Mastalerz. 2000. Taphonomy of fossilized resins: Determining the biostratino- my of amber. Acta Geologica Hispanica 35: 171— 182. Schlee, D. 1990. Das Bernstein-Kabinett. Stuttgarter Beitrage zur Naturkunde (C) No. 28, 100 pp. Silvestri, EF 1908a. Tisanuri raccolti da L. Fea alle isole del Cape Verde, alla Guinea Portoghese e alle is- ole S. Thomé, Principe e Fernando Poo. Annali del Museo civico di Storia Naturale di Genova 44: 133-187. . 1908b. Thysanura, pp. 46—68. /n Michaelsen, W. and R. Hartmeyer, eds. Die Fauna Siidwest- Australiens. Ergebnisse der Hamburger siidwest- australischen Forschungsreise. 1905, 2, Gustav Fi- scher Verlag, Jena. 1916. Descrizione di alcuni tisanuri indo- males. Bolletino del Laboratorio di Zoologia ge- nerale e agraria di Portici 11: 85-119. . 1918. Contribuzione alla conoscenza dei Ter- mitidi e termitofili dell’ Africa occidentale. II. Ter- mitofili. Parte prima. Bolleetino del Laboratorio di Zoologia generale e agraria di Portici 12: 287— 346. . 1932. Descrizioni di un nuovo genere ecitofilo di Lepismatidae (Insecta, Thysanura). Revista de Entomologia, SAo Paulo 2: 133-138. Smith, G. B. 1998. Review of the Australian Nicole- VOLUME 106, NUMBER | tiinae (Aygentoma: Nicoletiidae). Invertebrate Taxonomy 12: 135-189. Sturm, H. and L. E Mendes. 1998. Two new species of Nicoletiidae (Zygentoma, “‘Apterygota’’, Insec- ta) in Dominican amber. American Museum Novy- itaties No. 3226. 11 pp. Sturm, H. and G. O. Poinar, Jr. 1997. A new Neo- 109 machilellus species from Miocene amber of the Dominican Republic and its phylogenetic relation- ships (Archaeognatha: Meinertellidae). Entomo- logia Generalis 22: 157—170. Wygozinsky, P. 1980. A survey of the Nicoletiinae of Europe (Nicoletiidae, Thysanura, Insecta). Amer- ican Museum Novitates No. 2695. 24 pp. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 110-117 THREE NEW SPECIES OF CHILEAN CADDISFLIES (INSECTA: TRICHOPTERA) RALPH W. HOLZENTHAL Department of Entomology, 1980 Folwell Ave., Room 219, University of Minnesota, St. Paul, MN 55108, U.S.A. (e-mail: holzeOO1 @umn.edu) Abstract.—Three new species of caddisflies are described from Chile, Mastigoptila complicornuta, n. sp., Mastigoptila elae, n. sp. (Glossosomatidae), and Smicridea (Smi- cridea) figueroai, n. sp. (Hydropsychidae). The Mastigoptila species are characterized by asymmetrical male genitalia bearing long processes from the inferior appendages and phallocrypt and by complex phallic structures. The species of Smicridea is characterized by a pair of dorsomedial spines in the phallus. Key Words: new species, Chile, Neotropics, caddisflies, biodiversity, taxonomy, Glos- sosomatidae, Hydropsychidae Among all the South American countries, the Trichoptera fauna of Chile is perhaps the most well known. This is due mainly to the work of two researchers, the late FE Schmid, Canadian National Collection, and O. S. Flint, Jr., of the Smithsonian Institu- tion. Schmid, in a series of papers begin- ning in 1955 (e.g., Schmid 1958), described many new species and genera, almost en- tirely from material supplied to him by the Chilean naturalist and collector Luis E. Pena. Schmid’s other significant contribu- tion to our knowledge of the fauna was his monograph on Hydrobiosidae (Schmid 1989). Later, from about the mid-1960s to the present, Flint described much additional material, some supplied to him by Pena and other collectors, but most collected by him- self during several visits to the country (e.g., Flint 1967, 1974a, 1989, 1999, 2002). A few other workers, most notably L. Na- vas, described additional species. Together, Schmid, Flint, and Navas described 86% of the known Chilean caddisfly fauna (91, 72, and 18 species, respectively) (see Flint et al. 1999 for additional references to the Chilean fauna). Undoubtedly, Schmid and Flint’s interest in the fauna was influenced by its high de- gree of endemism; it is unlike anything in the rest of the Neotropical Region (Flint 1974b). At the species and generic level, Chile’s caddisfly fauna is almost 100% en- demic, but many of its families are shared with those of Australia and New Zealand. The fauna is clearly part of the well-docu- mented trans-Antarctic biota (Crisci et al. 1991). While the fauna is highly endemic, only about 210 species are known from the country (Flint 1974a, Flint et al. 1999); this is not very diverse by South American stan- dards. Even so, new species are still being discovered and described as Flint’s recent papers, as well as the present one, attest. In January of 2000, I had the pleasure of travelling to Chile with Dr. Fernando Mu- noz, University of Minnesota, at the invi- tation of Ms. Elizabeth Araya and Mr. Ri- cardo Figueroa, both then researchers in the aquatic biology section of ““-EULA-Chile” (Europe-Latin America, a research partner- ship between Chile and the European Union) at the Universidad de Concepcion. VOLUME 106, NUMBER 1 We collected at several localities in the foothills of the Andes from Concepcion south to Osorno, with most of the collec- tions coming from the Rio Bio Bio basin, the focus of study of the EULA researchers. The resulting material contained three new species, one each in the genera Mastigoptila (Glossosomatidae), Smicridea (Hydropsy- chidae), and Microthremma (Helicophidae). The specimens of Microthremma were sub- sequently sent to Dr. Flint for inclusion in his recent revision of the genus (Flint 2002) and were described as M. patagonica. In return, Dr. Flint kindly sent me specimens he collected in 1969 of a second unde- scribed species of Mastigoptila to include in this paper. Types of the species described herein are deposited in the University of Minnesota Insect Collection, St. Paul, Minnesota (UMSP), the National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C. (NMNH), and the Universidad de Concepcion, Concepcién, Chile (UDEC) as indicated below. Morphological terminolo- gy follows that of Flint (1989) for Smicri- dea, but a different interpretation from that presented by Flint (1967, 1974d) for the male genitalia of Mastigoptila is provided. Mastigoptila Flint Mastigoptila Flint 1967: 49 [type species: Mastigoptila curvicornuta Flint 1967, original designation].—Schmid 1958: 191 [diagnosis of Chilean species, as An- toptila Mosely]; Flint et al. 1999: 25 [sys- tematic catalog]. The description of the two new species below brings to ten the number of species now known in the genus (Flint et al. 1999), making it the largest of the three glosso- somatid genera occurring in Chile, all in the subfamily Protoptilinae and all endemic to the subregion. The other two Chilean gen- era, Scotiotrichia Schmid and Tolhuaca Schmid, are each monobasic. Valverde and Miserendino (1998) described the larval stages and biology of Mastigoptila longi- 111 cornuta from Argentina. Flint (1963) and Angrisano (1995) provided keys to the adults of the South American genera. Mastigoptila complicornuta Holzenthal, new species (Fig. 1) Like other species in the genus, this new species has highly complex, asymmetrical male genitalia. While these fit the general ground plan of the genus, they are unlike the male genitalia of any of the described species, except perhaps M. ecornuta which shares with the new species a hoodlike ter- gum X with an absence of processes. Male.—Length of forewing 5 mm. Body and appendages fuscous; forewings fus- cous, with small patch of cream colored se- tae at arculus and very faint tranverse line of lighter setae over cord. Genitalia as in Fig 1. Sternum VI with prominent poster- omesal process. Sternum VII with small, posteromesal point. Sternum VIII lacking process. Segment IX broadest laterally, an- terior margin broadly rounded, posterior margin angular, lightly sclerotized dorsolat- erally, excavated ventrolaterally, sternum produced posteriorly; in dorsal view, ter- gum IX with broad U-shaped excavation anteriorly, tergum forming very narrow, slightly produced sclerotized bridge; in ven- tral view, sternum IX with roughly V- shaped excavation anteriorly, produced pos- teriorly, sternum forming a heavily sclero- tized narrow bridge. Tergum X_ hoodlike, extending over phallic apparatus, setose apically, laterally, and ventrally, lightly sclerotized dorsally, posterior margin some- what irregular, ventral margin shelflike, produced mesally, heavily sclerotized me- sobasolaterally and forming apparent artic- ulation with segment IX; in dorsal view, tergum X irregular laterally, with broad, U- shaped excavation apically; internally with pair of hemispherical excavations and pair of terete, anteriorly directed apodemes. In- ferior appendages complex, asymmetrical, fused and produced medially into broad, quadrate, truncate, apically setose process; 112 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1 ' ' ' ‘ ‘ 1 1 1 ' 1 1 1 1 Fig. 1. Mastigoptila complicornuta, male genitalia. A, Segments IX and X, left lateral. B, Same, dorsal. C, Segment IX, ventral. D, Phallic apparatus, lateral. E, Phallocrypt, inferior appendages, and phallus, ventral. E Apex of phallic apparatus, enlarged, ventral. G, Inferior appendages and phallocrypt, right lateral. H, Segments VI-VIII, lateral. VOLUME 106, NUMBER 1 the inferior appendage complex fused ba- sally with base of lightly sclerotized phal- locrypt; left inferior appendage small, ear- shaped, bearing small, slender, recurved, posteriorly directed, spinelike basodorsal process; right inferior appendage larger, ear- shaped, with heavily sclerotized ridges ven- trally, bearing long, slender, highly re- curved, ventrally directed, spinelike, baso- dorsal process; with pair of very small se- tose processes at basolateral corners of fused inferior appendages. Phallocrypt a lightly screrotized cup, fused basally with base of inferior appendage complex, acting as receptacle for base of phallic apparatus; bearing long, slender, highly recurved, ven- trally directed, spinelike process originating from membranes of phallocrypt above base of phallic apparatus. Phallic apparatus asymmetrical, complex, tubular, with highly convoluted internal membranes basally; truncate subapically, apex flat, extended posteriorly as acutely triangular shelf; right side of phallic apparatus slightly inflated laterally, left side with small membranous setose protuberance; ventally with setose membranous mound on right side, large in- ternal toothlike spine on left side, medially with patch of fine microtrichia. Female.—Unknown. Type material.—Holotype, 6. CHILE: VIII Region del Bio-Bio: Bio-Bio: small imp to” Rio! Oueco, 5° km’ E) Ralco; nol Oess: 71 36,257 Ws els (500), 1m, 16.1.2000, Holzenthal and Munoz (UMSP). Paratypes. Same data as holotype, 4 ¢ (UMSP), 2%67(NMNH); 2 ¢ (UDEC). VI Region del Maule: Linares: Emb[alse] Bul- hileow36° 18'S. 71°25’ W, -11=12.4.1994,, C. and O. Flint, Jr., 2 ¢6 (NMNH). Etymology.—The name is derived from the Latin word for complex and refers to the complex, intricate structure of the male genitalia. The suffix “‘cornuta”’ is in keep- ing with that of other names in the genus, referring to the hornlike process of tergum X in those species (absent in M. complicor- nuta). Mastigoptila elae Holzenthal, new species (Fig. 2) This species is most similar to M. ven- tricornuta Flint. Both species share the pos- session of a prominent process on the ven- tral margin of tergum X and a similarly shaped inferior appendage complex. How- ever, in the new species the process on ter- gum X is much larger and the medial pro- cess of the inferior appendage is more quadrate than in M. ventricornuta. The two species differ significantly in the structure of the phallic apparatus, which in M. elae has an unusual receptacle for the tip of the recurved spinelike process of the inferior appendage. Male.—Length of forewing 3 mm. Body and appendages fuscous; forewings light fuscous, with smal! patch of cream colored setae at arculus and very faint tranverse line of lighter setae over cord. Genitalia as in Fig. 2. Sternum VI with prominent poster- omesal process. Sternum VII with small, posteromesal point. Sternum VIII lacking process. Segment IX broadest laterally, an- terior margin slightly rounded, posterior margin angular, lightly sclerotized dorsolat- erally, excavated ventrolaterally, sternum narrowly produced posteriorly; in dorsal view, tergum IX with broad U-shaped ex- cavation anteriorly, tergum forming very narrow, slightly produced sclerotized bridge; in ventral view, sternum [IX with roughly V-shaped excavation anteriorly, produced posteriorly, sternum forming a heavily sclerotized narrow bridge. Tergum X large, quadrate, hoodlike, extending over phallic apparatus, setose apically, laterally, and ventrally, lightly sclerotized dorsally, posterior margin truncate, produced baso- mesally into prominent process, not heavily sclerotized mesobasolaterally, without ap- parent articulation with segment IX; in dor- sal view, tergum X irregular dorsolaterally, truncate apically; internally with pair of narrow, sinuous excavations and pair of te- rete, anteriorly directed apodemes. Inferior 114 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON W/Z; ser a —— ey a g % “ft — Ye SSE . . =e. Fig. 2. Mastigoptila elae, male genitalia. A, Segment IX and X, left lateral. B, Segment X, dorsal. C, Phallocrypt, inferior appendages, and phallus, ventral. D, Phallic apparatus, lateral (showing insertion of apex of recurved basodorsal process of left inferior appendage). E, Right inferior appendage, lateral. appendages complex, asymmetrical, fused and produced medially into broad, rounded, setose process, in ventral view this process roughly triangular, apicolateral corners te- rete, excavated mesally; the inferior ap- pendage complex large, body heavily scler- otized with sclerotized ridges and folds, fused basally with base of lightly sclero- tized phallocrypt; left inferior appendage large, broadly S-shaped, bearing very long, slender, recurved, posteriorly directed, spinelike basodorsal process; right inferior VOLUME 106, NUMBER 1 appendage much smaller, roughly quadrate, with only very weakly developed basodor- sal process; with pair of larger setose pro- cesses at basolateral corners of fused infe- rior appendages. Phallocrypt a very lightly screrotized hemispherical cup, fused basally with base of inferior appendage complex, acting as receptacle for base of phallic ap- paratus; bearing shorter, thicker, curved, ventrally directed, spinelike process origi- nating from membranes of phallocrypt above base of phallic apparatus. Phallic ap- paratus asymmetrical, complex, tubular, roughly S-shaped, with highly convoluted internal and external membranes subapical- ly; apex flat, spatulate, extended posterior- ly; right side of phallic apparatus with unique, semisclerotized, rugose, cuplike re- ceptacle for apex of recurved basodorsal process of left inferior appendage, phallic apparatus without apparent internal spines, setae, or microtrichia. Female.—Unknown. Type material—Holotype, ¢. CHILE: VII Region del Araucania: Cautin: nr. Pucon, 4.1.1966, Flint and Cekalovic (NMNH). Etymology.—I am very happy to name this species in honor of Ms. Elizabeth Ar- aya in recognition of her hospitality and as- sistance during my visit to Chile and for her continued friendship. The name is derived from Elizabeth’s nickname, “Ely.” Smicridea McLachlan Smicridea McLachlan 1871: 134 [type spe- cies: Smicridea fasciatella McLachlan 1871, subsequent selection of Milne 1936]|.—Flint 1974c: | [revision of North and Central American species]; Flint 1989: 1 [revision of Chilean species]; Flint et al. 1999: 71 [systematic catalog]. This new species brings to 16 the number of Smicridea known from Chile. All belong to the subgenus Smicridea except one, S. murina McLachlan, in the subgenus Rhy- acophylax. Of the 16 species, 11 were de- scribed by Flint in his 1989 paper where he also described the immature stages and pro- 1S vided a key to the males and females. The genus is the largest among the Neotropical Hydropsychidae with more than 160 de- scribed species (Flint et al. 1999). Smicridea (Smicridea) figueroai Holzenthal, new species (Fig. 3) This new species seems to fall squarely within the smilodon group of Flint (1989), containing until now only the species S. re- dunca Flint and S. smilodon Flint; the new species 1s most similar to the latter. With those species, it shares the thin lobe on the ventral margin of tergum X and a short, compact phallus with the basal opening at a right angle to the stem and an enlarged apex bearing a pair of sclerotized spines. However, the details of the morphology of the phallic apex in S. figueroai are quite different, especially in its possession of dor- somedial spines rather than apical, ventrally directed spines in the other species. Male.—Length of forewing 6.5—7.0 mm. Color generally pale stramineous, append- ages paler, antennae annulate; dorsum of head and thorax with white setae; forewing whitish-yellow, apical two-thirds of wing faintly irrorate with light brown patches of setae, darker, larger patch at pterostigma, and series of more distinct patches of brown setae along basal half of costal margin. Eye of male in anterior aspect with diameter about two-thirds that of interocular dis- tance. Anterolateral process of sternum V shorter than sternum; abdominal segments VI and VII each with pair of internal, ellip- tical, reticulate sacs, each slightly shorter than the segment in which it lies. Genitalia as in Fig. 3. Segment [X with anterior mar- gin vertical. Tergum X deeply divided dor- somesally, tergite terete posteriorly in dor- sal view, slightly upturned in lateral view; ventral margin with large thin lobe basad. Inferior appendage with basal segment slen- der, slightly inflated apicad; apical segment short, blunt. Phallus with basal opening at right angle to stem, subapically enlarged, 116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Smicridea figueroai, male genitalia. A, Segments [X and X, lateral. B, Same, dorsal. C, Phallus, lateral. D, Apex of phallus, dorsal. E, Same, ventral. bulbous, especially dorsomedially, the en- larged area consisting of paired, broad, rounded lateral plates, flanking a membra- nous mesal region from which arises dor- somesolaterally a pair of hook-shaped, heavily sclerotized, posteriorly directed spines; phallus terminating in paired, lateral sclerites, cup- or scooplike mesally, longer than wide and projecting anteriorly well be- yond bulbous subapical portion. Female.—Unknown. Type material.—Holotype, ¢. CHILE: VIII Regién del Bto-Bio: Bio-Bio: small trib: ito Rio, Queco,. 5, km, Ey Ralco- 37°51.619'S, 71°36.257 W; el sO0R me 16.i1.2000, Holzenthal and Munoz (UMSP). Paratypes. Same data as holotype, 2 ¢d (UMSP), 1 ¢ (NMNH), 1 d (UDEC). Etymology.—I take great pleasure in naming this species in honor of my friend, Ricardo Figueroa, in recognition of his gen- erous assistance during my visit to Chile in VOLUME 106, NUMBER 1 2000 and his continued friendship and col- laboration. ACKNOWLEDGMENTS I am grateful for the support provided to me by EULA-Chile, Universidad de Con- cepcion, especially its director, Dr. Oscar Parra, during my research and collecting ef- forts in Chile. I am especially indebted to Elizabeth Araya and Ricardo Figueroa for their friendship and collaboration. Dra. Viv- ien Jerez graciously provided access to the insect collection in the Museo de Zoologia, UDEC. Fernando Munoz offered invaluable assistance in helping to arrange my visit to Chile and provided much help and com- radeship in the field. Dr. Oliver Flint gen- erously provided material, including one of the new species, for examination and inclu- sion in this study. Financial support to trav- el to Chile was provided by the Office of International Programs, University of Min- nesota. This material is based upon work supported by the National Science Foun- dation grant DEB 0117772. LITERATURE CITED Angrisano, E. B. 1995. Insecta Trichoptera, pp. 1199— 1237. In Lopretto, E. C., and G. Tell, eds. Ecos- istemas de Aguas Continentales: Metodologias para su Estudia, Tomo III, Identificaci6n de Or- ganismos. Ediciones Sur, La Plata, Argentina. Crisci, J. V., M. M. Cigliano, J. J. Morrone, and S. Roig-Junent. 1991. Historical biogeography of southern South America. Systematic Zoology 40: 152-171. Flint, O. S., Jr. 1963. Studies of Neotropical caddis flies, I: Rhyacophilidae and Glossosomatidae (Tri- choptera). Proceedings of the United States Na- tional Museum 114: 453-478. . 1967. Trichoptera collected by Prof. J. Illies on the Chilean subregion. Beitriage zur Neotro- pischen Fauna 5: 45—68. . 1974a. Checklist of the Trichoptera, or Cad- disflies, of Chile. Revista Chilena de Entomologia 8: 83-93. 117 1974b. A preliminary report of studies on Neotropical Trichoptera, pp. 47—48. Jn Malicky, H., ed. Proceedings of the Ist International Sym- posium on Trichoptera. Dr W. Junk Publishers, The Hague. 1974c. Studies of Neotropical caddisflies, XVII: The genus Smicridea from North and Cen- tral America (Trichoptera: Hydropsychidae). Smithsonian Contributions to Zoology 167: 1—65. 1974d. Studies of Neotropical caddisflies, XVIII: New species of Rhyacophilidae and Glos- sosomatidae (Trichoptera). Smithsonian Contri- butions to Zoology 169: 1—30. 1989. Studies of Neotropical caddisflies, XXXIX: The genus Smicridea in the Chilean sub- region (Trichoptera: Hydropsychidae). Smithson- ian Contributions to Zoology 472: 1—45. 1999. The Chilean genus Charadropsyche, with the description of its immature stages (Tri- choptera: Tasimiidae) (Studies of neotropical cad- disflies, LVII), pp. 99-105. In Malicky, H., and P. Chantaramongkol, eds. Proceedings of the 9th In- ternational Symposium on Trichoptera. Faculty of Science, Chiang Mai University, Chiang Mai, Thailand. . 2002. Studies of Neotropical caddisflies, LX: Three new species of the Chilean genus Micro- thremma, with a review of the genus (Trichoptera: Helicophidae). Entomological News 113: 225- 232. Flint, O. S., Jr., R. W. Holzenthal, and S. C. Harris. 1999. Catalog of the Neotropical Caddisflies (Tri- choptera). Ohio Biological Survey, Columbus, Ohio. McLachlan, R. 1871. On new forms, etc., of extra- European trichopterous insects. Journal of the Lin- nean Society of London, Zoology 11: 98-141. Milne, L. J. 1936. Studies in North American Trichop- tera. Part 3, pages 56—128. Cambridge, Massachu- setts. Schmid, E 1958. Contribution a Vetude des Trichop- teres Neotropicaux HI. Mitteilungen der Zoolo- gischen Museum Berlin 34: 183-217. . 1989. Les hydrobiosides (Trichoptera, Annu- lipalpia). Bulletin de l’Institute Royal des Scienc- es Naturelles de Belgique, Entomologie 59, Sup- plement: 1—154. Valverde, A. d. C. and M. L. Miserendino. 1998. Apor- tos al conocimiento do los estados preimaginales de Mastigoptila longicornuta (Trichoptera: Glos- sosomatidae). Revista de la Sociedad Entomol6- gica Argentina 57: 49—55. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 118-132 ANOPHELES (NYSSORHYNCHUS) KONDERI GALVAO AND DAMASCENO: NEOTYPE DESIGNATION AND RESURRECTION FROM SYNONYMY WITH ANOPHELES (NYSSORHYNCHUS) OSWALDOI (PERYASSU) (DIPTERA:CULICIDAE) CARMEN FLORES-MENDOZA, E. L. PEYTON, RICHARD C. WILKERSON, AND RICARDO LOURENCO DE OLIVEIRA (CFM) U.S. Naval Medical Research Center Detachment, Lima, Peru, APO AA 34031; (ELP) (deceased); (RCW) Department of Entomology, Walter Reed Army Institute of Research, Washington, D.C. 20307, U.S.A. (e-mail: wilkerson.richard@msc.si.edu); (RLO) Laboratorio de Transmissores de Hematozoarios, Instituto Oswaldo Cruz, Av. Brasil 4365, 21045-900 Rio de Janeiro, RJ, Brazil Abstract.—Anopheles (Nyssorhynchus) konderi Galvao and Damasceno 1942 is rede- scribed with illustrations of the male and female genitalia and the larval and pupal stages. A neotype for An. konderi is designated, and it is resurrected from synonymy with An. (Nys.) oswaldoi (Peryassu 1922). Key Words: Anopheles (Nyssorhynchus) konderi Gal- vao and Damasceno 1942 is similar to An. oswaldoi (Peryassu 1922) in larval, pupal and adult female characters, being distin- guished by only one character in the male genitalia. During the 1940’s, several authors considered An. konderi and An. oswaldoi as distinct species and made contributions to the knowledge of their geographical distri- bution and biology (Causey et al. 1946; Coutinho 1946; Deane et al. 1946, 1948). However, Lane (1953) considered An. kon- deri a synonym of An. oswaldoi and this concept was widely accepted. Indeed, the last revisions of Anopheles subgenus Nys- sorhynchus (Faran 1980, Faran and Linthi- cum 1981) agreed with Lane (1953). E.L. Peyton (apud Klein and Lima 1990) ob- served differences in the behavior and ma- laria transmission potential of material col- lected in Costa Marques, Brazil. He sug- gested the existence of two forms of An. oswaldoi: one was present in recently mod- Anopheles konderi, Anopheles, Culicidae, taxonomy, redescription, malaria ified open areas (An. konderi), and another restricted to forested areas (An. oswaldoi). Although the species has never been for- mally resurrected from synonymy with An. oswaldoi or morphologically well charac- terized in all stages, the name An. konderi has appeared in papers since Peyton’s state- ment (Lounibos et al. 1997, Marrelli et al. 1999). In the present paper, morphological and morphometric analyses of specimens of An. konderi and An. oswaldoi were conducted to distinguish these species and to rede- scribe the former. MATERIALS AND METHODS Progenies of females and immature stag- es of An. oswaldoi s.l. collected in six lo- calities in Brazil and one in Peru were in- cluded in the present study. To obtain prog- enies, females were blood fed and kept in individual oviposition vials. Some eggs from each female were fixed and stored in VOLUME 106, NUMBER 1 4% glutaraldehyde or alcoholic Bouin’s so- lution for morphological analyses. We follow the terminology of Harbach and Knight (1980) for morphological fea- tures and Wilkerson and Peyton (1990) for wing spot nomenclature. Abbreviations used are as follows: M, male; EK female; G, genitalia; Le, larval exuviae; Pe, pupal ex- uviae. The nomenclature adopted for the dorsal and ventral polarities of eggs is that of Clements (1992) and Valle et al. (1999), which is opposite of classical studies on the external morphology of the egg. The polar- ity is determined in the maternal organism during ovular development: the flat side of the egg or deck is considered the dorsal side and the submerged convex inferior side is ventral. Statistical analysis of morphometric characters was done using the Kruskal-Wal- lis test to verify the existence of significant differences among the samples of An. kon- deri from different localities. When mor- phometric characters were homogeneous between samples of An. konderi, they were subsequently compared with those in An. oswaldoi, using the Mann-Whitney test. Both the Kruskal-Wallis and the Mann- Whitney tests were done using the SPSS— Windows program version 8.0 (SPSS, Chi- cago), at a 5% significance level. The characters and ratios used for statis- tical analysis were: Female: Length of the wing, basal dark spots on hindtarsomere II, length of maxillary palpus/forefemur, total length of palpomere 3/size of basal white scaling on the same segment, proportion of basal dark-scaled band on fore- and mid- tarsomeres II and III, length of humeral pale spot/prehumeral dark, length of subcostal pale/sector dark, length of preapical pale/ preapical dark, length of apical dark/preap- ical pale and percentage of specimens with divided sector dark spot. Male: Length of the wing, basal dark spot on hindtarsomere II, ratio of length of parabasal seta/width of gonocoxite at base, length of the aedeagus/ length of claspette; length/width of sternum VIII (measured at base), length of gonocox- 119 ite/width of gonocoxite at base, width of gonocoxite taken at the widest point/width of gonocoxite at base. Pupa: Length of me- atus/length of trumpet, trumpet index (length trumpet/width trumpet), length of tracheoid/length of trumpet; length of seta 1-[V/length tergum V, and paddle index (length of paddle/width of paddle measured at the widest point). Larva: Clypeal index (distance between insertion of seta 3-C on one side/distance between insertions of se- tae 2-C), length of antenna/distance be- tween base of antenna and insertion of seta 1-A, length of anal papilla/length of seta 4- X, distance between apices of lateral arms of median plate of spiracular apparatus/dis- tance between spiracular opening (SO,), percentage of specimens with seta 1-X borne on saddle, type of pecten, and number of pecten spines. The type of pecten was clas- sified according to a formula, in which numbers were given to represent the size of spines: “0” for short spines, “1”’ for me- dium size spines (about twice as long as short spines “0”’) and ‘‘2” for large and long spines (about three times as long as Spine sine): TAXONOMIC TREATMENT Anopheles (Nyssorhynchus) konderi Galvao and Damasceno Anopheles (Nyssorhynchus) konderi Galvao and Damasceno 1942: 115-118, 132- 133. Type: Holotype male, right (south) margin of Rio Solimoes, Coari, State of Amazonas, Brazil (Departamento de Par- asitologia da Faculdade de Medicina de Universidade de Sao Paulo, S.P., Brasil), lost (Belkin et al. 1971). Galvao 1943: 156; Causey et al. 1946: 12; Coutinho 1946: 72; Deane et al. 1946: 27; Deane et al. 1948: 876; Lounibos et al. 1997: 148. Female (Figs. 1A, B, C).—Head: Integ- ument darkish brown. Interocular space with approximately 20 white long semi-de- cumbent fusiform scales. Vertex with nu- merous white erect spatulate scales. Occiput 120 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON = = ° £ = ° Fig. 1. and postgena with brown erect, spatulate scales. Postgena with elongate setae, few white spatulate scales at junction of eyes; ocular setae (8-12) long, brown. Clypeus brown, bare. Antenna: flagellum (1.14 mm) with 13 flagellomeres with darkish integu- ment and dirty white pollinosity, flagellar whorls pale; flagellomere | with decumbent white scales, and distal surface with. patch of white falcate scales dorsally. Pedicel in- tegument brown, with small patch of white decumbent scales dorsally. Proboscis 1.5— 2.7 mm (mean = 2.1 mm), integument dark brown; proximal third with brown erect scales, remainder covered by darkish brown decumbent scales. Proboscis nearly 1.0 (0.8—1.0) length of maxillary palpus. Labial basal setae 8, brown. Labellum brown, pal- er than proboscis, pollinose. Maxillary pal- pus |.7—2.7 mm (mean = 2.1 mm), integ- ument dark, covered with darkish narrow spatulate scales, 1.4 [(1.1—1.8) + 0.14, P = 0.03] [mean (range) + standard error of the mean, P] of forefemur; palpomere | length Anopheles konderi. A, Abdominal segment X of male. B, Female cibarium. C, Female cerci. Coari, Amazonas, Brazil. 0.14—0.21 mm (mean = 0.17 mm) covered with brown spatulate scales; palpomere 2 length 0.27—-0.4 mm (mean = 0.43 mm), covered with dark scales, dorsally sprinkled with few (7,8) white scales, and with apical narrow band of white scales; palpomere 3, length 0.64—0.94 mm (mean = 0.81 mm), almost completely dark, dark-scaled area 4.8 [(2.8-8.7) + 1.64, P = 1.64] times white-scaled area; palpomere 4 length 0.43—0.65 mm (mean = 0.5 mm), white- scaled dorsally, with narrow dark bands at ends; palpomere 5 length 0.22—0.31 mm (mean = 0.26 mm), completely white; ven- tral surface of palpomeres 1—4 dark-scaled. Cibarium (Fig. 1B) with 17 (14—21) cibar- ial teeth of variable form and size. Thorax: Scutum length approximately 1.0 mm; in- tegument darkish brown with pale scales and pollinosity; setae yellowish brown, nu- merous; integument mottled with 3 dark spots: 2 near scutal fossa and | on prescu- tellar area. Anterior promontory with white linear, erect scales. Antealar scales white, VOLUME 106, NUMBER | elongate and spatulate. Scutellum covered with small falcate grayish scales: scutellar setae brown, about 13 (11-15) long and 4 (4—9) short, distributed along caudal mar- gin. Antepronotum with spatulate, erect, brown scales, few pale scales basally; setae short, brown. Pleural integument brown to darkish brown. Usually 4 bronze upper me- sokatepisternal setae, about 7 (3-7) white upper mesokatepisternal scales, | brown lower mesokatepisternal seta, 2,3 white me- sokatepisternal scales, prealar scales and se- tae pale; about 3 yellowish upper mesepi- meral setae, lower mesepimeral scales ab- sent; upper proepisternal setae bronzy, thin and short; 2 brown lower proepisternal se- tae. Legs: Integument dark. Coxae: Anterior surface of forecoxa with patch of white scales, ventral 0.5 with brownish scales, 2— 4 brown setae; anterior surface of mid- and hindcoxae with pale scales transversally. Trochanters with patch of white scales, se- tae long and brown. Femora, tibiae and tarsi essentially dark-scaled, broadly pale gray- ish or light cream scales on ventral surfac- es. Femora and tibiae scattered with light cream scales dorsally, principally on fore- and midlegs. Foreleg: Femur length 1.28— 1.62 mm (mean = 1.45 mm); tibia, length 1.68—2.05 mm (mean = 1.86 mm); tarso- mere 1, length 1.0—1.34 mm (mean = 1.18 mm) with narrow band (mean = 0.04 of tarsomere) of white scales distally; tarso- mere 2, length 0.4—0.6 mm (mean = 0.5 mm) dark-scaled on basal 0.7 [(0.5—0.9) + 0.09, P = 0.128]; tarsomere 3, length 0.2— 0.5 mm (mean = 0.3 mm), dark-scaled on basal 0.3 [(0.1-0.6) + 0.14, P = OJ, re- mainder white; tarsomere 4, length O0.17— 0.26 mm (mean = 0.21 mm), completely dark-scaled, rarely with pale scales distally; tarsomere 5, length 0.14—0.2 mm (mean = 0.17 mm), dirty white, with ring of dark scales on basal 0.41. Midleg: Femur dark, length 1.54—2.08 mm (mean = 1.74 mm), with narrow band of white scales at base; tibia dark, length 1.8—2.37 mm (mean = 1.9 mm), with few pale spines at apex: tarso- mere | dark, length 1.2—1.7 mm (mean = 121 1.43 mm), with small band of white scales distally (0.04 of tarsomere); tarsomere 2, length 0.5—0.8 mm (mean = 0.66 mm), dark-scaled on basal 0.9 [(0.7—1) + 0.06, P = 0.69]; tarsomere 3, length 0.3-0.7 mm (mean = 0.5 mm), usually completely dark (0.9 [0.7—1] + 0.04, P = 0.45), dark-scaled, with sparse pale scales distally; tarsomere 4, length 0.29—0.34 mm (mean = 0.29 mm) usually all dark-scaled, with rare pale scales distally; tarsomere 5, length 0.17—0.21 mm (mean = 0.18 mm), dark-scaled on basal 0.4, remainder white. Hindleg: Femur, length 1.5—2.1 mm (mean = 1.7 mm), with 2 distal strong yellowish setae; tibia, length 1.7—2.5 mm (mean = 2.0 mm), with small ring of white scales distally; tarsomere 1, length 2.34—2.66 mm (mean = 2.63 mm), with narrow ring of white scales distally; tarsomere 2, length 0.63—0.89 mm (mean = 0.76 mm), dark-scaled on basal 0.14 [(O0.08—0.20) + 0.036, P = 0.23], remainder white; tarsomeres 3 and 4 white, tarsomere 3, length 0.4—0.69 mm (mean = 0.58 mm); tarsomere 4, length 0.31—0.48 mm (mean = 0.41 mm); tarsomere 5, length 0.26—0.34 mm (mean = 0.26 mm), dark-scaled on basal 0.5. Wings: Length 3.0—4.0 mm (mean = 3.4 mm = 0.207, P = 0.27), pale wing spots white, dark spots darkish brown to black. Basal pale spot plus prehumeral pale spot large, length 0.24 mm (0.20—0.26 mm), more than 2.5 of prehumeral dark; basal dark spot absent; humeral pale spot 3:6) (42-910) y= Al eRe — Oslo lgot spre humeral dark; subcostal pale spot 0.2 [(O— 0.4) + 0.09, P = 0.302] of sector dark; preapical pale spot (PP) 0.3 [(0—0.5) + 0.09, P = 0.38] of preapical dark (PD); api- cal dark spot 0.4 [(O.1-1) + 0.69, P = 0.684] of PP; accessory sector dark present in 82.4% (+ 0.921, P = 0.65) of specimens examined. Size of dark spots on Costa varying within progenies and between pop- ulations (n = 51). Veins CuA, M,,,, M,,>, M,, M, and IA covered with sparse thin fusiform scales; veins R,,3, R5, R, and R,,; densely covered with linear scales. Vein R, with 4,5 (5) dark spots. R,+R,,; with 1-3 122 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (2) dark spots; when only | dark spot pre- sent it occupies more than 0.7 of vein; when 2 dark spots present, basal white spot small- er than distal one. R, with 1-3 (2) dark spots; when 2 dark spots present, apical spot smaller; R, with 1—3 (2) dark spots; most frequently with 2 small spots, proxi- mal spot smaller than distal one; R,,; with 2 small dark spots, | basal and other distal. M,.,. with 1—3 dark spots, more frequently with 2 dark spots; when only | dark spot present it occupies more than 0.7 of vein, ventral surface of M,,, covered with dark scales. M, with 1,2 (2) dark spots, some- times sprinkled with white scales; M, with 1,2 (1) dark spots. M,,, with 3 small dark spots. CuA with | small dark spot distally; 1A with 2 dark spots; CuP with 1 small distal dark spot. R,,; separating from R,,, on level of distal 0.33 of sector dark spot. M,,, 0.33 longer than CuP. Cell R, 0.7 of cell M,.*Cell M,; 0.5 of M,,,5. Remigium with integument pale. Halter: Length 0.35 mm; integument of scabellum, pedicel and ventral surface of capitellum pale, remain- der of capitellum and distal 0.35 of pedicel dark. Abdomen: Integument dark brown, pollinosity grayish, scales falcate. Terga: tergum I with abundant brown setae; terga II-VI with posterolateral tufts of broad erect spatulate dark scales and distal median patches of grayish to cream scales in rough- ly triangular shape; tergum VIII covered with abundant cream scales; posterolateral setae numerous and long. Sterna with few brown setae; sternum I without white scales; sterna H—VII with white, spatulate scales laterally; posteromedian area with patch of spatulate brown scales. Sternum VIII length/width = 1.0 (0.5—1.5). Genita- lia (Fig. 1C): Sternum IX, length/width = 0.6 (0.2—1.0); cercus elongate, with spatu- late brown scales dorsally, and dark scales and setae ventrally; 2 postgenital setae, in- serted close together, strong, length 6.2 (4.6—10) distance between them; postgenital setae generally slightly smaller than cercus, length of postgenital seta/length of cercus = 0.7 (0.5—1.0); insula bare. Male (Figs. 2D, E, F).—Similar to fe- male except for sexual and few other dif- ferences as follow. Head: Interocular space with about 20—26 long white setae. Anten- na strongly verticilate, length 1.5—1.9 mm (mean = 1.7 mm), flagellomeres with in- tegument grayish, heavily plumose, setae long and pale. Proboscis length 1.9—2.4 mm (mean = 2.2 mm), maxillary palpus 1.0 (0.9-1.0) length of proboscis. Maxillary palpus: palpomere | as in female; palpo- meres 2 and 3 with one narrow band of white scales distally, palpomere 4 white- scaled on dorsal surface, with dark bands of dark scales at ends; setae of different lengths, some developed setae inserted ba- sally surpass palpomere 5; paipomere 5 covered dorsally with pale scales, setae pale and dark; palpomeres 4 and 5 expanded, with dark scales on ventral surface. Thorax: Lower mesokatespisternal scales (8—10) white. Legs: Hindtarsomere 2 dark-scaled on basal 0.15 [(0.10—0.20) + 0.020, P = 0.054]. Wing: Length 3.3 mm [(2.9-3.7 mm) + 0.21, P = 0.16]; subcostal dark spot 0.4 of sector dark; dark spots on internal veins smaller than in female. Halter: Length 0.16—0.27 mm (mean = 0.24 mm). Abdomen: Sternum VIII roughly trapezoi- dal in shape, moderately elongate, ratio length/width = 0.8 [(0.1-1.3) + 0.14, P = 0:11). Genitalia> (Fig: 2D) E, F) Lobestor sternum IX as wide as long, length 0.65 mm (0.4—0.8 mm). Parabasal seta 0.8 [(0.5— 1.6) = 0.14, P = 0.01] width of gonocoxite. Gonocoxite elongate, length 3.3 [(1.7—4.7) + 0.66, P = 0.009] width at base, expanded on basalO:S: [width h5°d:0—2.1)) 20262 = 0 width at base], outer surface strongly convex, inner surface concave, long setae on tergal surface extending to ventrolateral surface. Ventrolateral surface with numer- ous long spatulate scales, dorsomesal sur- face with narrow linear patch of small, short setae. Gonostylus moderately shorter than gonocoxite, length (0.85) of gonocoxite, strongly curved, with internal spicules be- yond middle; gonostylar claw spiniform and blunt. Ventral lobe of claspette with VOLUME 106, NUMBER 1 oimm Fig. 2. Anopheles konderi (drawn from specimens BR103(5)-1, BRO95(1)-2, and BRO97(2)-17, Costa Marques, Brazil). A-C, Pupa. A, Cephalotorax. B, Metathorax. C, Abdominal segments. D-E Male. D, Male genitalia. E, Ventral lobe of claspette. F Aedeagus. 124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON apical lobe moderately sclerotized, narrow, lobe length 3 times its width at base, with conspicuous median sulcus; refringent structure moderate in size; setae short and strong; basal lobule expanded laterally, with numerous long and strong setae distributed along basal margin. Dorsal lobe of claspette with pedicel moderately narrow, apex round, narrower than base, apex with 3,4 moderately broad leaflets. Phallosome: Ae- deagus length 1.3 [(1.0-1.6) + 0.21, P = 0.104] length of claspette; weakly rounded at apex, length of apex of aedeagus about 0.4 (0.3—0.6) of width, apical aedeagal sclerite narrow and curved in elbow-like lateral projections. Paraproct weakly scler- otized, narrow distally and expanded at base. Pupa (Fig. 2A, B, C).—Position and de- velopment of setae as figured; range and modal number of branches in Table 1. In- tegument brown, sclerotized. Cephalotho- rax: Integument more pigmented than ab- domen. Trumpet: length 0.63 mm (0.52—0.9 mm), laticorn, pigmented and_ spiculose, tragus elaborate, trumpet index 2.3 [(1.6— 5.7) = 0.06,,P < 0.0001], meatus length 0.2 [(0.1—0.4) + 1.36, P = 0.53] of trumpet length; pinna 0.12—0.28 mm (mean = 0.21 mm); tracheoid length 0.4 [(0.2—0.7) + 0.40, P = 0.4] of the trumpet length. Ab- domen: Seta 1-IV strongly sclerotized, length 0.35-0.55 mm (mean = 0.46 + 0.06); seta 1-IV moderately developed, 1.6 (1.1—2.2) length of tergum V; tergum V length 0.23—0.39 mm (mean = 0.29 mm); seta 9-II minute; 9-III—VIII thick, short, dark brown. Paddle: pale, slightly paler than abdomen, ovate, slightly longer than wide, index 1|.1—1.7 (1.4) + 0.24; external margin spiculose, paddle marginal spicules more developed on distal 0.5; midrib dis- tinct. Larva (Fig. 3).—Position and develop- ment of setae as in Fig. 3; range and modal number of branches in Table 2. Head: In- tegument pale, collar strongly pigmented. Antenna: length 1.0 mm (0.93—1.17 mm), antenna length 4.9 [(2.8-8.5) + 1.19, P = 0.22] distance from insertion of seta 1-A to base; ventral surface of antenna with short spicules, less numerous distally; seta 1-A inserted 0.2 mm (0.1—0.4 mm) from base of antenna, with 3—10 (5) branches; length of seta 1-A 0.7—1.5 (1.0) times width of an- tenna at point of insertion; seta 4-A_ bifur- cate; setae 2,3,5,6-A usually tapered at apex; 2,3-A almost same size; 5-A short, half size of 2,3-A, 6-A slightly shorter than 5-A. Setae 2-C widely separated, clypeal index 1.4 (0.8—1.9) + 0.27; setae 2,3-C al- most same size, branched, branches usually dendritic. Ventromentum pale, with 3 teeth on each side of 2 central more developed teeth. Dorsomentum dark, strongly sclero- tized, with 4 teeth on each side of one cen- tral more developed tooth. Thorax: Seta 1- P palmate; |—3-P arising from distinct tu- bercles; 1 1-P single or double, much shorter than 9,10,12-P but much more developed than 1-M,T; 3-T weakly developed, pal- mate. Abdomen: Integument pale. Seta 1I-I palmate, with 9—24 (14) moderately devel- oped, weakly pigmented branches; 1-II-VI palmate, leaflets usually broad, well devel- oped and strongly pigmented; I-X usually inserted outside saddle (86.3% of speci- mens examined). Saddle incomplete. Anal gills hyaline, length 0.43 mm (0.28—0.63 mm), 0.9 [(0.6—1.4) + 0.19] length of seta 4-X. Posterior margin of segment X with numerous short spicules. Spiracular appa- ratus: Lateral arms of median plate devel- oped, elongate, projecting toward spiracular process or spiracular opening; distance be- tween apices of lateral arms 1.3 [(0.8—1.8) + 1.17] of distance between SO,. Pecten with 16 [(12-20) + 4.76, P = 0.036] spines. Three types of pecten were found: type I = 2-0-2(n)-0-2(n)-0; type If = O(n)- 2(n)-1-2(n)-O and Type III = 1(n)-2-0-2(n)- Q-2(n)-0. The most common formula was type I [formula = 2-0-2(n)-0-2(n)-O], in which ‘“‘n” is the variable number of repe- titions of a kind of spine in the pecten. Egg (Fig. 4).—Boat-shaped in both dor- sal and lateral views; ventral view almost flat. Length 421 wm (379-520 wm); width VOLUME 106, NUMBER | onlupuap = q = I I = _ -- — #7 =F aa is ar = = aos ae =" 7 aos el = = az oy = = = =e - (1) eI ae a = ae (WD) =I ID) GE (CD) G7 Decal (ID) =| I = (py) S-C = I] ri =a (@) er — ID) | (ID) =| (€) PI — 7 CIDEGal = Ol = I I I I I I I I = ()) | 6 = = Me S=|I (@) €=| (©) v1 (@) | (2) | a ae aS I 8 a _ Gl (CID) fel K@) 1 (OO) CG) | (C) 8-1 0 | ~~ (Ce) II L —— ae (1) TI I) =I (QD) Gul (ID) =I (WD) =I (CD) el (1) 71 = (1) HI 9 = Fe CL) GY (WD) =| (1) tI (€) 8-1 (8) OI-€ (G2) =| U@)} (=| — (@) {eI S > (t) S-C (@) | (O) Wl (€) $-T (CO Se! (O) | (py) LI (bp) L-€ a= GO) rl 174 — —- (vy) S-I (CQ) II (¢) 9-] (p) O-T (ID) S| I (1) TI = (S) ital € (@) G1] = (¢) tI (€) 7-1 Ge) S| (@) GI OS) LAE (9) OI-1] MS) Ee Te (@) eI C I I I I I (1D) =" (8) CI-IT (Ol) vI-7 d = (ED) 1 I = Cina (vy) 9-€ (vy) 9-€ (vy) 9-€ (S) 9-€ (G2) HAS G2) LC Se = 0) a1PPed IIA IA IA A Al Il II I uinjouRay xproyjoyeyday ag sjusuIsag jePUILIOpGY BUdG “IBUIOINDH JOpeuas “LieuUef Op SelopuRD ‘OUlaA OO, ‘LROD) suoneindod uviizeig xIs Jo ‘(apoul) aduvI sMapuoy “uy JOJ Surpouriq [eras jedng ‘(SeNSeUILIN A) UPIANIOg duo pure (SaIeyUT] pur vitoinpeyy "T qeL 126 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON He EEF ZZ Aa Lhig¢=-———> ee WF Je Fig. 3. Anopheles konderi, larva (drawn from specimens BRO97(2)-10 and BRO97(2)-8, Costa Marques, Brazil). A, Head, al—dorsomentum, a2—ventromentum. B, Thorax and abdominal segments I-VI. C, Abdom- inal segments VII—X. D, Spiracular apparatus. Scales in mm. 127 VOLUME 106, NUMBER 1 asournjd = q Fr. imal ie <== ry am aa i = = <= = = (C)SI-T Sl a ra; I I I I I I I a ay, (6)cI-+ (QS) I-€ — (DP-I ia -F I = (P)8-€ (S)TI-S (S)8-+ (Q)TI-+ (6)TI-+ (L)9I-+ (L)€El+ ()F-I (()6-1 (E)S—I — (WL-C el Tra I — I I (Q)E-I (€)9-C (e)r-1 (DoT (O)r-1 (C)e-I (DTI I a (P)9-T cl =; I a (C)6-1 ()r-I ()t-I (CEI (Z)8-I (DTT (Q)P-1 (DTT I "1 oa (Of)9b-S1 Il =e I = (S)L-C (C)E-T I I I (EOS) I I I I za (C)t-I Ol — (faye = (Q)OI-€ (S)CI-€ (LCI + CXEN=E (Q)CI-€ (Q)L1-Z (E)OI-€ I I I = (9)6-C 6 = (Ore =; (p)6-C (€)¥-C (€)p-T (E)p-C (e)v—-C (Q)EC ~~ MAM ace WOW eI! (80)re+1 = (S)8-T 8 = Dye" = (WTI-€ (ET (€)h-C CE) 9SC (€)9-c (cE)Or-ec (6C)IP-It (CE)SP-ST (Q)¢-I (CE)CP-81 ae (60)0E-S L — (We ne (Q)6-€ | I I (d0r-81 (e)IP-L7 CME)Ib—-E? (OE-1 (O)r-I I — (SL)Fc-0l 9 (S)6-€ (Q)O0I-- (6-4 (S)8-T (E)8-€ (2) Taller (Q)OI-€ MAS ADGA (De-I (SC)LEF1 3 (LUCC-11 g d — (DET I I (€)h-C (Q)r-1 (¢)9-C (S)6-€ (y)9-€ (E)S-C (e)o-€ (CI)ST—-6 (ET (T)9-I v d = (QAR (€)S—Z I I (¢)9-] I I (DTT (C1)9I—-9 I I I (6)EI-T t al Woke (Mglae (S)TI-€ (S)8-€ (e)p-I I (€)8-C (Q)8-€ (p)L-C I LE SWiScD cae I (WANs C CS) San (WI GA scaUI VOSS CSG) Ceacln(SO9EOL (7o)ES=011 (Odct-6 = (FI )P7-6 I (9d)cE-8I (ED0cs6) (S) ONS I I (trl (F)9-C (H)9-€ (S)L~C (H)8-€ (S)8-€ (P)OI-€ a a i I <3 I 0 X sniereddy IITA IIA IA A Al il I I 1b W d uy PpPoH ‘ON qepnoesds pas s]uswidag [eulMOpqy XPIOUL, ‘(sensuvurlin x) uoneyndod urtAniag uo pure (soreyury pure rwoiInd Jopeuas ‘eur Op sBlapuRD ‘oUlaA OWOd ‘eOD) UeT[IzeIg XIS JO ‘(QpOW) BduRI “LapUOoY “Uy IOJ SurpouRsG [RIAs [RAIeT “TZ IQR], 128 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig 4. 130 pm (114-170 pm); length 3.4 (2.4— 3.8) of width. Floats moderately wide, about 0.87 (0.74—0.9) of total egg length, with approximately 36 (35—38) float ridges; float occupying nearly 0.33 of dorsal sur- face; deck narrow, hardly visible on poste- rior end in most of examined eggs but much wider on anterior end. Anterior deck en- closed by frill. Anterior end broadly round- ed; posterior end somewhat pointed. Type material.—Neotype male by pre- sent designation, with slide mounts of 1) associated larval and pupal exuviae, and 2) male genitalia, from the progeny brood of a female captured on animal bait at the type locality of An. konderi, data as follow: south margin of Solimoes River at Coari G°S7'S,, 63°12’ W);, “State of Amazonas, Brazil, specimen 1629, 15-VIII-1998, C. Flores-Mendoza coll., deposited at Instituto Oswaldo Cruz (IOC), Rio de Janeiro, Bra- ZAll. Material examined.—Anopheles konderi: a total of 763 specimens, consisting of 188 M, 2857 116° Pe, 16 Ee, 98 MG and) 25 FG, were studied, as follows: BRAZIL: State of Amazonas, Coari, Travessia, 17— 23-VII-1998, progeny brood of female captured on animal bait, C.Flores-Mendoza and ESouza colls., 11M, 12MG, 7MPeLe, Anopheles konderi, egg. Coari, Amazonas, Brazil, dorsal surface. Scanning electron micrograph. 11E 4FG, 4FPeLe. State of Rond6énia, Porto Velho, Sao Miguel at Madeira River (8°55'S, 64°12'W), 28-IV-1996, progeny brood of female captured on human bait, D.Lima coll-; 11M, 11MG, SMPeles (tle 6FG, 6FPeLe; Candeias de Jamari, Samuel Hydroelectric Dam (8°55'S, 64°08'W), 2- V-1997, progeny brood of female captured on human bait, C.Flores-Mendoza and M.Marrelli colls., 11M, 11MG, 7MPeLe, 11K 4FG, 4FPeLe; Costa Marques and vi- cinity (12°28’S 64°16’W), dates 1989-1992, progeny broods of females captured on human bait with the following specimen numbers (BR = Brazil, collection number, (progeny brood number), all de- posited in the Smithsonian Institution, Na- tional Museum of Natural History), J.B. Lima and T:A.Klein colls.: BRO95(1) 3M, IMG, 5E 1PeLe; BRO97(2) 1M, 1MG, 1PeLe; BR100(1) 1M, IMG, 5F; BR100(2) IM; “IMG, SF; BRI03@G); 2M; ING. SE: BR103(4) 2M, IMG, 5F; BR103(5) 2M, various 2MG, SE 1 Pele; sBRUI2ZC)eIMe IMG. SE: BRIN2G6).2M, IMG..4E; BRAS) 2Me IMG, -2F:" BRIT9@) IMe IMGs as: BR119(6) 1M, IMG, 5F; BR120(2) 1M, IMG, TF? BRI235C) SIM MG ak BR133(2) 2M, IMG, 5F; BR136(2) 2M, IMG, 5F; BR144(1) 2M, IMG, 3F; VOLUME 106, NUMBER I BR144(2) 2M, 1MG, 5F; BR161(3) pica 4; BR1I61G6) 1M, 1MG, BR170(1) 1M, IMG, 4F; BR170(5) IMG, 4F; BR174(5) 4M, IMG, BR175(4) 2M, 1MG, 6F; BR176(2) PMG. SF; BR1764) 4M, 1MG, 6F; BR176(13) 4M, IMG, 6F; BR178(4) 2M, IMG, 8F; BR277(14) 1M, 1MG, 12PeLe; Be 7715) 1M, IMG, 9PeLe; BR277(17) IM, IMG, 10PeLe; BR277(18) 1M, IMG, Pielke: BR277(19).1M, 1MG, 9PeLe; BR277(20) 1M, 1MG, 5PeLe; BR277(23) IM, IMG; BR289(1) 2M, 2MG; BR289(2) IM, IMG; BR289(3) IM, IMG; BR612(1) 13M, 1MG, 18F; BR612(2) 7M, IMG, 18F; BR61I2(3) 11M, IMG, 14F; BR613(1) 11M, IMG, 18F; BR613(2) 13M, 1MG, 14F; BR644(1) ISM, IMG, 10EF State of Acre, Senador Guiomar, Ramal Oco do Mundo (10°09’S, 67°44'W), 9—13-III-1998, progeny brood of female captured on ani- mal bait, R.Santos coll. 3M, 3MG, 2M hele, 36 2FG, tFPeLe; Sena Madureira; Sermeal. Sao Pedro de <«Ic6 (9°05’S, 68°45'W), 22-VIII-1998, progeny brood of female captured on animal bait, R.Santos Bone ooMe 2MG...1MPebe, 2E) 2FG, 2FPeLe. State of Espirito Santo, Linhares, Sooretama forest reservation (19°41’S, 39°59'W), 15—25-IV-1996, progeny brood of female captured on animal bait, C.Flores-Mendoza and C.Santos, colls., Brie IMG 2E PG; thPeLe: PERU: Lor eto Departament, Yurimaguas, Munichis (05°53'S, 76°12'W), 5—10-I-1999, progeny brood of female captured on animal bait, C.Flores-Mendoza, R.Fernandez and T.Santa Cruz colls. 11M, 11MG, 6MPeLe, RE tiRG i SePeLe. Distribution.—According to our data and the literature records, An konderi occurs in Brazil (states of Amazonas, Acre, Rond6n- ia, Espirito Santo, Para, S40 Paulo, Mato Grosso and Rio de Janeiro), Peru (Loreto Department) and Bolivia (Cochabamba). Bionomics.—Anopheles konderi has been collected most often close to or inside forest rather than in peridomestic environ- ments. It bites primarily outdoors, at sunset. 2M, 5) Be 2M, 6F; 3M, 129 In Coari, although An. konderi females were captured from sunset until 21:00h and around sunrise, the peak of biting activity was between 17:30 and 18:30h. Although it bites humans, An. konderi is mostly zoo- philic. In Coari collections performed in a corral, 26 out of 55 anophelines caught were An. konderi. In Porto Velho, using hu- man bait, only 17 An. konderi were cap- tured (among 270 anophelines), whereas in Samuel it accounted for 40 specimens out of 152 anophelines caught. In Senador Guiomar, no An. konderi was found among 485 anophelines collected on human bait, Whereas three females belonging to this species were obtained from a horse (among 110 anophelines). In Munichis, 10% of 1,207 anophelines collected using a horse- baited Shannon trap were An. konderi; no specimens were captured on human bait at this locality. Larvae of An. konderi have most often been found in shaded or partially shaded pools, small streams, and temporary lakes formed during the flooding of rivers. These sites usually have emergent vegetation and sometimes contain muddy water rich in de- composed plant debris. In Coari, 48 and 44 out of 93 anopheline larvae collected in a small stream that received the flow of a waste drainage pipe were An. konderi and An. (Ano.) mattogrossensis Lutz & Neiva, respectively. Larvae of An. konderi have been found together with An. (Nys.) nunez- tovari Gabaldon, An. (Ano.) mediopuncta- tus s.l. (Theobald), An. (Nys.) rangeli Ga- baldon, Cova-Garcia and Lopes, An. (Ano.) punctimacula Dyar and Knab and An. (Ano.) mattogrossensis (Galvao and Da- masceno 1942, Deane et al. 1948). Medical importance.—The role of An. konderi in malaria transmission is unknown, primarily because females belonging to this species could not be distinguished from those of An. oswaldoi. Experimental infec- tions with Plasmodium vivax suggested that An. konderi is less susceptible than An. os- waldoi (Marrelli et al. 1999). 130 DISCUSSION Throughout it range from Southeastern Brazil to the Amazon Valley it is possible that many literature records referring to An. oswaldoi are actually An. konderi. Precise identifications can only be verified by ex- amination of the male genitalia. The two species are sympatric at most collection sites. However, along the Solimoes and Amazon Rivers in the state of Amazonas it is our impression that either only An. kon- deri is present, or it is at least much more abundant than An. oswaldoi. At present only An. oswaldoi is reported from Vene- zuela, northern Colombia, Panama and Costa Rica, while only An. konderi has been found in Peru. Morphological and morphometric analy- ses of An. konderi from seven localities in Brazil and Peru showed that it is a highly variable species since variation was detect- ed between specimens from the same lo- cality and from the same progeny brood. In females, 11 out of 13 morphometric mea- surements analyzed did not show statisti- cally significant differences (P > 0.05). The ratios length of palpus/hindfemur (P = 0.003) and length of dark-scaled band on foretarsomere III/total length of tarsomere III were significantly variable (P = 0.001). Three out of seven morphometric measure- ments or ratios taken from the male geni- talia were significantly different: length of parabasal seta/width of gonocoxite at base (P = 0.001), length of gonocoxite/width of gonocoxite at base (P = 0.009), width of gonocoxite taken at the widest point/width of gonocoxite at base (P = 0.001). The five ratios or indices evaluated in the pupa were homogeneous between populations, where- as three out of seven morphological char- acters and ratios analyzed in the larva were heterogeneous: length of anal papilla/length of seta 4-X (P = 0.001), percentage of specimens with seta 1-X born on saddle (P = 0.035) and type of pecten (P = 0.037). The eggs oviposited by An. konderi females from five different localities in Brazil and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Peru were morphologically similar. How- ever, the mean length and width of eggs from these localities (length = 421 wm, width = 130 pm) are smaller than those found by Lounibos et al. (1997) for speci- mens from Alto Linares, Bolivia (length = 520 wm; width = 197 pm). Anopheles konderi is closely related to An. oswaldoi and therefore will key out with An. oswaldoi in the keys to females and larvae in the most recent revisions of the Albimanus Section of Anopheles sub- genus Nyssorhynchus (Faran 1980, Faran and Linthicum 1981). No diagnostic mor- phological or morphometric differences were found between the pupae or eggs of these species. However, they are readily distinguished by the shape of the aedeagus. In An. oswaldoi, the aedeagus is ovate and sclerotized at the apex, the length of the apex of the aedeagus is about 1.2 (0.8—2.1) of the width, while in An. konderi the ae- deagus is weakly rounded at apex, the length of apex of aedeagus is about 0.4 (O.3—0.6) of width, and the apical aedeagal sclerite is narrow and curved into elbow- like lateral projections. Males of An. kon- deri key out to An. evansae Brethes in the key for male genitalia in Faran and Linthi- cum (1981), but An. konderi can be distin- guished from An. evansae, as well as from the other Nyssorhynchus of the Albimanus Section, by the shape of the aedeagus. The morphological and morphometric charac- ters of the females and larvae of these spe- cies are also very similar. Statistical analy- sis showed some morphometric differences between females of these species: in An. os- waldoi foretarsomere 2 is dark-scaled on 0.6 (0.4—0.8) + 0.1 (while it is 0.7 [0.5— 0.9] + 0.09 in An. konderi, P < 0.0001), hindtarsomere 2 dark-scaled on 0.11 (0.08— 0.16) + 0.026 (while it is 0.14 [0.08—0.20] + 0.036 in An. konderi, P < 0.0001), sub- costal pale spot 0.3 (0 003-0.5) + 0.1 of sector dark (0.2 [0—0.4] + 0.09 in An. kon- deri, P < 0.0001), preapical pale 0.4 (0.2— 1.0) + 0.2 of preapical dark (0.3 [(O—0.5] + 0.09 in An. konderi, P = 0.009), acces- VOLUME 106, NUMBER 1 sory sector dark present in 82.4% (+ 0.921, P = 0.65) of specimens examined (around 60% in An. konderi). ACKNOWLEDGMENTS We are grateful to: Douglas Watts and the Fundagao Nacional de Satide for logistic support in Peru and Brazil, respectively; Pe- dro Cabello for helping in the statistic anal- yses; Taina Litwak, Lisa Roberts, and Glo- ria Gongalves for the illustrations; Jacenir Mallet for helping in processing the eggs for SEM; Bruce Harrison, Ralph Harbach and Dan Strickman for very helpful reviews of the manuscript; Janaina Neves for help- ing with the morphological and morpho- metric analyses; Dinair Couto Lima, Fatima dos Santos (FUNASA), Flaviano de Souza, Renan Souza dos Santos, Claudiney Santos, Roberto Fernandez, and Maria Santa Cruz for assistance in making collections; Judith Stoffer for digital setup of the illustrations and, Jose Bento Lima and Terry Klein for collecting and rearing specimens from Cos- ta Marques, Rondonia, Brazil. This research was partially performed under a Memoran- dum of Understanding between the Walter Reed Army Institute of Research and the Smithsonian Institution, with institutional support provided by both organizations. The opinions and assertions contained here- in are those of the authors and are not to be construed as official or reflecting the views of the Navy Department, the naval service at large, the Department of the Army or the Department of Defense. LITERATURE CITED Belkin, J. N., R. X. Schik, and S. J. Heineman. 1971. Mosquito studies (Diptera, Culicidae) XX V. Mos- quitoes originally described from Brazil. Contri- butions of the American Entomological Institute (Ann Arbor) 7: 1—67. Causey, O. R., L. M. Deane, and M. P. Deane. 1946. II. An illustrated key by male genitalic character- istics for the identification of thirty-four species of Anophelini from the northeast and Amazon re- gions of Brazil, with a note on dissection tech- nique, pp. 21-31. Jn Studies on Brazilian Anoph- elines from the Northeast and Amazon Regions. 131 American Journal of Hygiene, Monographic Se- ries 18. Clements, N. A. 1992. The biology of mosquitoes. The egg shell 3: 63-73. Ed. Chapman and Hall, Lon- don. Coutinho, J. O. 1946. Anofelinos do Rio de Janeiro (Distrito Federal) com referéncia aos transmis- sores de malaria. O Hospital 30: 651—662. Deane, L. M., O. R. Causey, and M. P. Deane. 1946. I. An illustrated key by adult female characteris- tics for the identification of thirty-five species of Anophelini, with notes on the malaria vectors (Diptera, Culicidae), pp. 1-18. Jn Studies on Bra- zilian Anophelines from the Northeast and Ama- zon Regions. American Journal of Hygiene, Monographic Series 18. . 1948. Notas sobre a distribuigao e a biologia dos anofelinos das Regides Nordestina e Amaz6n- ica do Brasil. Revista do Servico Especial de Sau- de Publica 1: 827-963. Faran, M. E. 1980. Mosquito Studies (Diptera, Culic- idae). XXXIV. A revision of the Albimanus Sec- tion of the subgenus Nyssorhynchus of Anopheles. Contributions of the American Entomological In- stitute (Ann Arbor) 15: 1-215. Faran, M. E. and K. J. Linthicum. 1981. A handbook of the Amazonian species of Anopheles (Nyssor- hynchus) (Diptera: Culicidae). Mosquito System- atics 13: 1-81. Galvao, A. L. A. 1943. Chaves para a determina¢ao das espécies do Subgenero Nyssorhynchus do Brasil. Arquivos de Zoologia, SAo Paulo 8: 141— 162. Galvao, A. L. A. and R. G. Damasceno. 1942. Anoph- eles (Nyssohynchus) konderi nova espécie de Anopheles do Vale do Amazonas e considerag6es sobre as espécies do complexo tarsimaculatus (Diptera: Culicidae). Folia Clinica et Biologica 14: 115-135. Harbach, R. E. and K. L. Knight. 1980. Taxonomists’ Glossary of Mosquito Anatomy. Plexus Publish- ing Inc., Marlton, NJ. xi + 415 pp. Klein, T. A. and J. B. P. Lima. 1990. Seasonal distri- bution and biting patterns of Anopheles mosqui- toes in Costa Marques, Rondonia, Brazil. Journal of the American Mosquito Control Association 6: 700-707. Lane, J. 1953. Neotropical Culicidae. Universidade de Sao Paulo, Vol 1, 548 pp. Lounibos, L. P., D. Duzzak, and J. Linley. 1997. Com- parative egg morphology of six species of the Al- bimanus Section of Anopheles (Nyssorhynchus) (Diptera: Culicidae). Journal of Medical Entomol- ogy 34: 136-155. Marrelli, M. T., N. A. Honorio, C. Flores-Mendoza, R. Lourengo-de-Oliveira, O. Marinotti, and J. K. Kloetzel. 1999. Comparative susceptibility of two members of the Anopheles oswaldoi complex, An. 132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON oswaldoi and An. konderi, to infection by Plas- (Diptera: Culicidae). Journal of Medical Entomol- modium vivax. Transactions of the Royal Society ogy 27: 207-224. of Tropical Medicine and Hygiene 93: 1—4. Valle, D., A. T. Monnerat, M. J. Soares, M. G. Rosa- Statistical Package for the Social Sciences, Version 8. Freitas, M. Pelajo-Machado, S. B. Vale, H. L. SPSS, Chicago. Lenzi, R. Galler, and J. P. B. Lima. 1999. Mos- Wilkerson, R. C. and E. L. Peyton. 1990. Standardized quito embryos and eggs: Polarity and terminology nomenclature for the costal wing spots of the ge- of chorionic layers. Journal of Insect Physiology nus Anopheles and other spotted-wing mosquitoes 45: 701-708. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 133-139 TWO NEW GALL-INDUCING SAPHENISTA WALSINGHAM (LEPIDOPTERA: TORTRICIDAE: COCHYLINI FROM COSTA RICA KENJI NISHIDA AND DAvID ADAMSKI (KN) Sistema de Estudios de Posgrado en Biologia, Escuela de Biologia, Universidad de Costa Rica, 2060 San, José, Costa Rica (e-mail: knishida@cariari.ucr.ac.cr); (DA) De- partment of Systematic Biology, Entomology Section, National Museum of Natural His- tory, Smithsonian Institution, Washington, DC 20560-0168, U.S.A. (e-mail: dadamski@ sel.barc.usda.gov) Abstract.—Saphenista muerta, n. sp., and S. bimaculata, n. sp., were reared from galls they induced on stems of Monnina crepinii Chodat (Polygalaceae) and Ageratina ixiocladon (Benth.) R.M. King & H. Rob (Asteraceae) respectively. Galls were discovered in a high elevation (3,000 m) oak forest on Cerro de la Muerte, Costa Rica. We present photographs of the imagos, habitat, and host; illustrations of the male and female genitalia, and descriptions of the galls. Resumen.—Saphenista muerta y S. bimaculata nov. spp., fueron criados de agallas que se encontraban en tallos de Monnina crepinii Chodat (Polygalaceae) y Ageratina ixiocladon (Benth.) R.M. King & H. Rob (Asteraceae), respectivamente. Estas especies fueron recolectados en un bosque de roble en Cerro de la Muerte, Costa Rica; alrededor de 3,000 m.s.n.m. Se presenta fotografias de adultos, habitat, y planta hospedera; ilustra- ciones de genitalia de macho y de hembra; y descripciones de las agallas. Key Words: Ageratina, Asteraceae, Central America, gall-inducer, high elevations, Mon- nina, Polygalaceae, Tortricinae Cochylini are an undisputably monophy- letic group currently recognized as a tribe within the Tortricinae (Tortricidae) (Horak and Brown 1991, Horak 1998). Historical- ly, the group has been treated by various authors as a family (e.g., Phaloniidae by Clarke (1968) and others, and Cochylidae by Powell (1983)), subfamily (Cochyinae), or subtribe (e.g., Cochylidii by Kuznetsov and Stekolnikov (1973), and Rawowski and Becker (1983)). The most convincing apo- morphy supporting its monophyly is the loss of the gnathos in the male genitalia. Although an uncus is present in nearly all species, it is well developed in the most primitive members of the tribe (e.g., Trachysmia), and hence its shared loss can- not be considered an autapomorphy for the entire tribe. Cochylini includes about 930 described species (Brown 2002) assigned to about 55 genera, and occurs in all faunal regions worldwide. The larvae feed inter- nally in flower heads, seed capsules, stalks, and roots, and most species associated with Asteraceae (Powell 1980). Cochilini larvae (e.g., Pogue 1988, Pogue and Friedlander 1987) are characterized by the presence of an anal fork (plesitomorphic condition), the occurence of SD1 and D1 on the same pi- naculum on abdominal segment 9 (shared with most Olethreutinae), conspicuously enlarged pinacula (a convergent character 134 with some internal-feeding Grapholitini), reduced number of crochets on the prolegs (convergent with many tortricids), and a bi- setose L-group on abdominal segment 9 (putative autapomorphy for the tribe). The genus Saphenista (Cochylini) was described by Walsingham (1914) on the ba- sis of several plesiomorphic characters. Based on the North American members, Pogue (1986) identified two autapomorp- hies for the genus: (1) valva widened ba- sally, gradually narrowed to a rounded apex; and (2) a pocket at the base of the valva with an associated costal groove that opens to the inside. He (Pogue 1986) also identified the presence of the hindwing cos- tal fold and 3-segmented maxillary palpus as holoplastic characters. In addition, all Saphenista have a distinctive transtilla with an elongate mesal process broadening dis- tally into a Y-shaped projection. The genus includes about 75 species and occures an all faunal regions except the Ethiopan (Po- gue 1986). Only a few larval host plants have been reported for Saphenista, and all but one are in the Asteraceae. The discovery of two undescribed spe- cies Of Saphenista in stem galls during a survey of gall-inducing Lepidoptera in Cos- ta Rica is noteworthy, and we take this op- portunity to describe these two species and comment briefly on their life histories. The galls, which were induced by the moth species described herein were collect- ed from the stems of Monnina crepinii Cho- dat (Polygalaceae) and Ageratina ixiocla- don (Benth.) R.M. King & H. Rob (Aste- raceae), in a high elevation oak forest at Villa Mills (3,000 m), Cerro de la Muerte, in the provinces of San José and Cartago, Costa Rica. Species of Monnina are shrubs or small trees (Fig. 5) found in cloud for- ests. Flowers are papilionaceous and blue to bluish-purple in color; leaves are uni- formly alternate. In Costa Rica, M. crepinii has been colleced between 1,800—3,000 m on both the Atlantic and Pacific regions of the central cordillera (INBio database 1977). Ageratina ixiocladon is a tropical PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON shrub (Fig. 6) that grows 3—4 m tall, and like M. crepinii, is usually found in partially open areas with limited sunlight (Fig. 4). In Costa Rica, A. ixiocladon is known from high elevations, about 2,500 m on both the Atlantic and Pacific slopes of the central cordillera (INBio database 1977). The forests of these high regions as de- scribed by Hartshorn (1983) and Kappelle (1996) are tropical montane rain forests or tropical montane cloud forests that are dominated by a few plant species. The tran- sitional zone is dominated mainly by Quer- cus costaricencis Liebmann and Q. cope- yensis, C.H. Muller (Fagaceae). Other plant species in moderate abundance are Miconia biperulifera Cogniaux (Melastomataceae), Vaccinium consanguineum Klotzsch (Eri- caceae), Weinmannia pinnata Linnaeus (Cunoniaceae), Schefflera rodriguesiana D.G. Frodin (Araliaceae), and Chusquea spp. (Poaceae). The climate is characterized by a wet and a dry season. The dry season lasts from De- cember or January to April. During this time of the year, rain is infrequent, although the humidity remains high and dense fog is common. The wet season lasts from April to November or December. Heavy rains are common during these months, and the area receives an average annual rainfall of 2,812 mm. The average temperature for this area is 10.9°C and sometimes can reach —3°C during the dry season. Galls were collected in the field, placed in plastic bags, and taken to an air-condi- tioned room where the temperature was kept between 16—18°C. Kornerup and Wanscher (1978) is used as a color standard for description of the adult vestiture. Gen- italia were dissected as described by Clarke (1941), except mercurochrome and chlora- zol black were used as stains. Adults were examined with dissecting and compound microscopes. Measurements were made us- ing a calibrated ocular micrometer. All specimens of this study are deposited in Instituto Nacional de Biodiversidad (IN- Bio), Santo Domingo de Heredia, Costa VOLUME 106, NUMBER 1 Figs. 1-6. Rica. Vouchers of Monnica crepinii and Ageratina ixiocladon are deposited in Her- bario de la Universidad de Costa Rica, Es- cuela de Biologia, San, José, Costa Rica. Saphenista muerta Nishida and Adamski, new species (Biss; °3:,7) Diagnosis.—Saphenista muerta appears most similar to S$. eneiema Razowski, 1990, Holotypes of Saphenista spp., gall, habitat, and colonies of Monnina crepinii and Ageratina ixiocladon. 1, Holotype of S. muerta. 2, Holotype of S. bimaculata. 3, Gall of S. muerta. 4, High elevation habitat of M. crepinii and A. ixiocladon. 5, Colony of M. crepinii. 6, Colony of A. ixiocladon. from Costa Rica. It differs by having a fore- wing pattern that is darker, a socius that is broader, a valva that is narrower, an aedea- gus with a longer apical spinelike process, and a vesica with a stout cornutus. Description.—Head: Frontoclypeus and vertex brownish yellow; outer surface of la- bial palpus pale yellowish brown, with third segment pale gray intermixed with white scales; inner surface white intermixed with 136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON few pale-gray scales; scape and basal 8—10 flagellomeres of antenna brownish yellow intermixed with gray scales, distal flagello- meres gray. Proboscis present. Thorax: Tegula and mesonotum brown- ish yellow, intermixed with gray scales on caudal part. Leg with coxa and femur pale brownish yellow intermixed with pale-gray and gray scales, tibia and tarsomeres dark gray intermixed with pale gray near distal end. Forewing (Fig. 1) length 7.0 mm (n = 1) all fasciae golden yellow intermixed with brownish-yellow scales; costal margin brown; costal strigulae silver, basal strigu- lae incomplete, distal strigulae complete; cell with a dark-brown streak about % length and a large dark-brown spot near dis- tal end; fringe pale yellowish brown; un- dersurface brown. Hindwing pale gray, slightly darker along anterior margin; costal fold present. Male genitalia (Fig. 7): Uncus and gnat- hos absent; socius elongate, setose, rounded distally; vinculum with two V-shaped parts, fused ventrally, posterior part dorsally fused with ventral articulation of valva, anterior part fused with dorsal articulation of valva; valva_ setose, slightly broadened basally, gradually narrowed distally forming a rounded apex; transtilla enlarged medially forming an elongate, dorsoposteriorly pro- jecting, Y-shaped process with two apically divergent tips; juxta platelike; aedeagus ba- sally broadened, gradually narrowed distal- ly, distal part with an elongate, protuberant spinelike process; vesica with an elongate cornutus. Female genitalia: Unknown. Type.—Holotype, ¢, “Costa Rica, San José [Province], Cerro de la Muerte, Villa Mills, Sendero, frente de Georgina, 3,000 m, 22-VI-1999, col/rear: Kenji Nishida, host plant Monnina sp. stem gall inducer, 22-VI-1999, pupa in gall 1-VI-1999, adult 22-VI-1999”’, ““COSTA RICA: INBio: CRIO02, 607856” [bar code label]; “‘d IN- Bio Genitalia Slide by D. Adamski No. 890” [yellow label]. The pupal exuvium is within a gelatin capsule and is attached to the pin of the holotype. Etymology.—The specific epithet is de- rived from the locality, Cerro de la Muerte, Mountain of Death, where the gall contain- ing the holotype was collected. Remarks.—Spindle-shaped galls (Fig. 3) were usually located near the midlength of the stem of Monnina crepinii near the nodes of the host plant. The galls are about 11 mm wide and about 20 mm in length. The stem width of the host is about 9 mm. An oval- shaped orifice (= exit hole) was covered with plant debris bounded with silk. The larva bores the central part of the pith of the stem. The larva of the holotype pupated in its gall chamber, spinning a loose co- coon, which was constructed with silk and frass. The pupa protruded prior to the emer- gence of the adult. Saphenista bimaculata Nishida and Adamski, new species (Figs. 2, 8-9) Diagnosis.—The genitalia of Saphenista bimaculata are most similar to those of S. gnathmocera Razowski, 1990, from Costa Rica; the species share a similarly shaped valva and aedeagus, but S. bimaculata dif- fers by having a narrower apical half of the valva and a stouter apical cornutus in the vesica of the aedeagus. Description.—Head: Frontoclypeus and vertex pale yellowish orange; outer surface of labial palpus brown intermixed with pale yellowish-brown scales or pale yellowish brown intermixed with few brown scales; inner surface pale yellowish brown inter- mixed with some white scales; scape of an- tenna pale yellowish orange, basal half of flagellum pale yellowish orange, distal half gray; proboscis present. Thorax: Tegula yellowish orange inter- mixed with pale yellowish-orange scales near distal margin; mesonotum missing most scales; legs with segments and tarso- meres brown with pale yellowish-brown apices. Forewing (Fig. 2) length 7.0—7.3 mm (n = 3) yellowish orange intermixed VOLUME 106, NUMBER | Siz Figs. 7-8. with pale yellowish-orange scales; a single dark-brown spot near midcell with a diffuse spot of dark-brown scales beyond cell; cos- ta with several small dark-brown spots and two large dark-brown spots, one near mid- length and one near half distance between the first spot and apex; several small, dark- brown, paired, submarginal spots present; posterior margin with irregular row of dark- brown scales; undersurface dark brown with dark-brown costal strigulae separated by pale yellowish-orange scales along cos- ta. Hindwing pale yellowish orange with several pale-brown irregular bands from midlength along costa to apex; costal fold present. Male genitalia (Fig. 8): Uncus and gnat- hos absent; socius elongate, setose, rounded Male genitalia, genital capsule and aedeagus of Saphenista spp. 7, S. muerta. 8, S. bimaculata. distally; vinculum with two V-shaped parts, fused ventrally, posterior part dorsally fused with ventral articulation of valva, anterior part fused with dorsal articulation of valva; valva setose, slightly broadened basally, gradually narrowed distally forming a nar- rowly rounded apex; transtilla enlarged me- dially with an elongate, dorsoposteriorly projecting, Y-shaped process with apically divergent tips; juxta platelike; apical part of aedeagus with a ventromedian lobe, distally narrowed and angled ventrally; vesica with an elongate basal cornutus, spinulate api- cally with a stout cornutus. Female genitalia (Fig. 9): Papillae ana- les elongate, setose, divergent anteriorly; ventral arms of sterigma slightly setose api- cally; apophyses posteriores dilated subba- 138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 9. Female genitalia of Saphenista bimaculata. sally, extending to near ostium; apophyses anteriores posteriorly bifurcate, one arm fused with dorsoanterior margin of sterig- ma, one curved medioventrally, forming part of a deeply notched seventh sternum; seventh sternum partically overlaid by pos- terior part of sixth sternum; ductus bursae, short, cylindrical; corpus burase ovoid, with elongate creases on side opposite ductus se- minalis, undulate and spinulate medially; signum absent. Types.—Holotype, 6, “COSTA RICA, San José [Province], Cerro de la Muerte, Villa Mills, Senderos Georgina, 3,000 m, 22-11-2000, Col[lected] and reared by Ken- ji Nishida, Host plant: Ageratina sp., Gall former on stem apex’’, “*d INBio Genitalia Slide by D. Adamski No. 891” [yellow la- bel]. Paratype: 1 2, Same label data as ho- lotype except, “3000 m 21-III-2000”’, “‘Pu- pation: 28-III-2000’’, ‘‘Gall former on apex of stem’’, “2 INBio Genitalia Slide by D. Adamski No. 892” [yellow label]. Etymology.—tThe specific epithet, bima- culata, is derived from the Latin maculata, meaning spot, and refers to the two large spots on the costa of the forewing. Remarks.—Galls were found on Agera- tina ixiocladon near the apex of the stem near the nodes (not figured). The globose or slightly elongate galls are about 6 mm wide and 7-18 mm long. The stem of the host is about 4 mm wide. Most of the larval fecal matter is ejected outside of the gall chamber through the exit hole, which later is covered with silk. Pupation occurrs with- in or outside the gall chamber. The pupa protruded prior to adult emergence. ACKNOWLEDGMENTS We thank Jozef Razowski, Institute of Systematics and Evolution of Animals, Pol- ish Academy of Sciences, Krakéw, Poland, for help with taxonomic placement of Sap- henista muerta and S. bimaculata; John S. Steiner and James F Di Loreto, Office of Imaging and Photographic Services, Smith- sonian Institution, Washington, D.C. for the photographs of the holotypes and digital or- ganization of all figures; Grace Wood and Alexander Rodriguez for identification of Ageratina ixiocladon and Ju-lin Weng for Monnica crepiniit; Harold E. Robinson and Deborah A. Bell, Department of Systematic Biology, Botany, Smithsonian Institution, for assistance with plant nomenclature and for biological information on Ageratina ixiocladon,; and Kuniko Arakawa for draw- ing Figs. 7-8. LITERATURE CITED Brown, J. W. 2002. How many species of Tortricidae are there? TORTS Newsletter 3(1): 1. VOLUME 106, NUMBER 1 Clarke, J. E G. 1941. The preparation of slides of the genitalia of Lepidoptera. Bulletin of the Brooklyn Entomological Society 36: 149-161. 1968. Neotropical Microlepidoptera, XVII. Notes and new species of Phaloniidae. Proceed- ings of the United States National Museum 125 (3600). 58 pp. Hartshorn, G. S. 1983. Plants: Introduction, pp. 118— 157. In Janzen, D. H., ed. Costa Rican Natural History. Chicago University Press, Chicago, IIli- nois, xi + 816 pp. Horak, M. 1998. Tortricoidea, pp. 199-215. Jn Kris- tensen, N. P., ed. Handbuch der Zoologie, Lepi- doptera, part 1, Volume 35, Berlin, New York. Walter de Gruyter & Co. 494 pp. Horak, M. and R. L. Brown. 1991. 1.2 Taxonomy and phylogeny, pp. 1—22. In van der Geest, L, P. S. and H. H. Evenhuis, eds. Tortricid pest, their bi- ology, enemies and control. Elsevier Science Pub- lishers B.V., Amsterdam. Kappelle, M. 1996. Los Bosques de Roble (Quercus) de la Cordillera de Talamanca, Costa Rica: Bio- diversidad, Ecologia, Conservacion y Desarrollo. Universidad de Amsterdam/Instituto Nactional de Biodiversidad. xvi + 319 pp. Koenerup, A. and J. H. Wanschner. 1978. Methuen Handbook of Colour. 2nd ed. Methuen and Co., Ltd., London, 243 pp. Kuznetsov, V. I. and A. A. Stekolnikov. 1973. Phylo- genetic relationships in the family Tortricidae (Lepidoptera) treated on the base of study of func- tional morphology of genital apparatus. Trudy 139 Vsesoyuznogo Entomologicheskogo Obschches- tua 56: 18—43. [In Russian. ] Pogue, M. G. 1986. A generic revision of the Cochy- lidae (Lepidoptera) of North America. Ph.D. Dis- sertation, Department of Entomology, University of Minnesota. xxiii + 280 pp. . 1988. Revision of the genus Lorita Busck (Lepidoptera: Tortricidae: Cochylini), with de- scription of a new species. Proceedings of the En- tomological Society of Washington 90: 440—457. Pogue, M. G. and T. P. Friedlander. 1987. Cochylis caulocatax Razowski (Lepidoptera: Tortricidae: Cochylini): A redescription of the male with de- scriptions of the female, larva, and pupa. Pro- ceedings of the Entomological Society of Wash- ington 95: 320-327. Powell, J. A. 1980. Evolution of larval food prederen- ces in microlepidoptera. Annual Review of Ento- mology 25: 133-159. . 1983. Tortricoidea, pp. 31—42. In Hodges, R. W., ed. Check list of the Lepidoptera of America north of Mexico. E.W. Classey, Ltd., and the Wedge Entomological Research Foundation, Lon- don. Razowski, J. 1990. Cochylini (Lepidoptera, Tortrici- dae) from Costa Rica. Miscellanea Zoologica 14: 85-103. Razowski, J. and V. O. Becker. 1983. Brazilian Cochy- lidii (Lepidoptera, Tortricidae). Acta Zoologica Cracoviensia 26(13): 421—464. Walsingham, Lord (de Grey). 1914. Lepidoptera: He- terocera. Biologia Centrali-Americana Lepidop- tera, Heterocera, Part 4, 482 pp. London. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 140-158 TWO NEW SPECIES OF MEALYBUGS (HEMIPTERA: COCCOIDEA: PSEUDOCOCCIDAE) FROM PATAGONIA, ARGENTINA MARIA CRISTINA GRANARA DE WILLINK AND DOUGLASS R. MILLER (MCGW) CONICET, Superior Institute of Entomologia “Dr. A. Willink’? (INSUE), National University of Tucuman, Foundation Miguel Lillo, Miguel Lillo 205, (4000) San Miguel de Tucuman, Tucuman, Argentina (e-mail: ewillink @arnet.com.ar); (DRM) Sys- tematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, Rm. 137, Bldg. 005, BARC-W, Beltsville, MD 20705, U.S.A. (e-mail: dmiller@sel.barc.usda.gov) Abstract.—Two new species of mealybugs, Eurycoccus chubutensis and Pseudanton- ina vernacula, have been discovered in the Patagonia Region of Argentina collected on Nothofagus dombeyi and on a grass, probably Stipa sp. Adult females and available im- mature stages of each species are described and illustrated and are compared with other mealybugs from the area. Key Words: The mealybug fauna of Argentina ap- pears to be inadequately described. A quick comparison of relatively well-known areas of the world gives important insight on the state of knowledge of Argentina’s fauna. The number of mealybug species reported in California (land area of about 158,000 square miles and a large diversity of habi- tats) is 201 (Ben-Dov 2002), the number of species in England (land area of about 50,000 square miles and a relatively uni- form diversity of habitats) is 46 (Ben-Dov 2002), and even in Maryland (about 10,000 square miles and relatively uniform habi- tats) there are 43 species known (Ben-Dov 2002). In comparison, the number of mealybug species in Argentina (land area of about 1,000,000 square miles and quite di- verse habitats) is only 46 (Ben-Dov 2002). The works of Leonardi (191 1a, b), Granara de Willink, (1979; 1981, 1983a, ‘b, 1986; 1991a, b, c), Williams (1973), Williams (1985b), and Williams and Granara de Wil- link (1992) have added significantly to Coccoidea, Nothofagus, Patagonia, grass, Neotropical knowledge about the mealybug fauna of the country, but much remains to be discovered and recorded. It is important to understand the mealybug fauna of Argentina, not only to fill major gaps in the knowledge base of the Pseudo- coccidae, but also to know the identity of species that could be pests either in Argen- tina or other parts of the world. With the vast temperate climate of major parts of the country it would seem likely that native mealybug species could easily adapt to the temperate climates of the heavily populated northern hemisphere areas of the world such as Europe, China, and the United States. Williams and Granara de Willink (1992) made the case that a comprehensive under- standing of the mealybug fauna of Central and South America was an effective weapon for combating mealybug species that could become invasive pests in other parts of the world. Their suggestion has been substanti- ated by the introduction of the invasive pa- paya mealybug (Paracoccus marginatus VOLUME 106, NUMBER 1 Williams and Granara de Willink) in the Ca- ribbean, Florida, and recently the Pacific (Meyerdirk, personal communication 2002). Unlike the years that it took to identify, de- scribe, locate the area of origin, and develop effective biological control strategies for the cassava mealybug (Phenacoccus manihoti Matile-Ferrero) (Miller and Rossman 1995), control for the papaya mealybug has been relatively rapid (Meyerdirk, personal com- munication 2002). The quick response was possible primarily because the mealybug was known, well characterized, and potential areas of origin for discovering biological control agents were understood before it be- came an invasive pest. Within Patagonia in Argentina most re- search on scale insects has focused on the mountainous, forested areas which are im- portant as sources of wood and other bio- logical resources. However, the steppe zone of Patagonia has numerous species of grass- es and other unique vegetation and harbors a wealth of unknown mealybug species that could be important in the future. These nat- ural pastures are especially resistant to the adversities of the harsh climate and are ben- eficial by maintaining soil moisture and preventing soil erosion. The grasses and low vegetation of the pampas are also im- portant as the primary food source of sheep and cattle and provide shelter for many small mammals and numerous birds. This work examines two unusual mealybug spe- cies from Patagonia and is part of a larger study on the scale-insect biodiversity of Ar- gentina. MATERIALS AND METHODS Specimens were prepared following the techniques described by McKenzie (1967) and were slide mounted in Canada balsam. Illustrations were made with a drawing tube and follow the international conventions generally used for illustrating scale insects (e.g., Williams 1985c). All specimens were examined in detail and were used to for- mulate the ranges and means for the nu- meric characters. Measurements of the ho- 141 lotype are given separately. Ranges are giv- en first, followed by the mean in parenthe- ses. Specimens are deposited in the Institute and Fundacion Miguel Lillo (MLA), of Tu- cuman, Argentina, and the Coccoidea Col- lection of the The National Museum of Nat- ural History, Beltsville, Maryland, USA (USNM). RESULTS Eurycoccus Ferris Eurycoccus Ferris, 1950; type species, by original designation, Pseudococcus jes- sica Hollinger 1916 Gunior synonym of E. blanchardii (King and Cockerell)). This genus includes 14 species from most continents of the world (Afrotropical 3, Australasian 2, Nearctic 4, Oriental 2, and Palearctic 3) but has never been re- corded from the Neotropical Region (Ben- Dov 2002). At present, Eurycoccus may contain several unrelated lineages, but we reluctantly decided to include the new Pa- tagonian species in it rather than add to the confusion by describing another monotypic genus. We have examined illustrations or spec- imens of all 14 Eurycoccus species (E. an- tiscius Williams, E. blanchardti, E. both- riochloae Williams, E. campbelli Kosztar- ab, E. coccineus (Newstead), E. copallinae Ferris, E. cuniculorum Williams, E. esakii (Kanda), E. glomerulus De Lotto, E. mon- odi Balachowsky and Ferrero, E. saudiensis Matile-Ferrero, FE. sternlichti Williams, E. tamariscus Williams, E. yuccae Ferris) and note similarities that occur in nearly all spe- cies (Table 1). The new species possesses all of these characteristics but differs by having: | more seta on each side of the basal segment of the labium; more numerous setae on the clypeolabral shield; an unusual type of dis- coidal pore that has a broad basal sclero- tized ring and a non-sclerotized area that protrudes from the ring; a rectangular ven- tral invaginated pocket on the intersegmen- tal line between segments VIII and IX; no 142 Table 1. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Distribution of characters considered important in diagnosing Eurycoccus. Large, Rotund Number of With Paired Cerarian Ventral Tubular Eurycoccus Species Body Coxal Pores Cerarii Setae Ducts antiscius yes no ] yes, elongate yes blanchardii yes no yes, conical yes bothriochloae yes yes 1 yes, conical yes campbelli no yes ] yes, conical yes coccineus yes rh i yes, elongate no copallinae yes ey 1 yes, conical yes cuniculorum yes yes 0) no yes esakit yes yes i yes, conical yes glomerulus no yes 1 yes, elongate yes monodi yes yes 1 yes, conical yes saudiensis no yes 0) no yes sternlichti yes yes i yes, conical yes tamariscus no yes I yes, elongate yes yuccae yes yes 1 yes, elongate yes tubular ducts; no elongate anal-lobe seta; panded, usually divided by intersegmental and multilocular and trilocular pores with Picea ah ee ercl yin bie oo an « nD Z - Circulus absent or small, round, usually obscure loculi. Although these characters Haat cee k y intersegmental line ..... 6 may be diagnostic of a group of mealybugs, 6(5) Translucent pores absent from hind tibia; with only a single representative of that cixculus present. ., 22... oe ae ee 7 clade we hesitate to describe it as a unique ~— Translucent pores present on hind tibia; genus. Other species of Eurycoccus have circulus absent PANS, DAMS CAEL AL ROS Eads 9 Dee 7(6) Ventral multilocular pores present on an- unusual small pores similar to those on the tapide Anarene Geren peer alee VI g gment New Specics.(E. cuntewlorien andi. tamar- 4 ee ae 8 iscus) and one lacks ventral tubular ducts = - Ventral multilocular pores absent from an- (E. coccineus). Therefore, we are placing terior margin of abdominal segment VII ihemew species im Murycoccus, 9 6) ee : bothriochloae Williams 8(7) Dorsal anal lobe with concentration of KEY TO SLIDE-MOUNTED ADULT FEMALES about 10 flagellate setae and 2 associated conical setae; antennae 8-segmented OF THE GENUS EURYCOCCUS Es sg aguas septs forbs OS ees aR OR esakii (Kanda) Tubular ducts present at least near vulva - Dorsal anal lobe with 2 or 3 slightly en- Agel St Led satel a OMT EMS Lee Tey «fem, ALeaaES 3 larged setae only; antennae 6- or 7-seg- - ihubularniductssabsentqar ae eee ee 2 Tented! Ars sy. esas cee glomerulus De Lotto 2(1) Translucent pores present on hind coxa 9(6) Dorsal anal lobe with pair of slightly en- and femur; multilocular pores present on larged setae and | or 2 flagellate setae . . POLAK so asaychoas tt “eo ces gs to bcete uso SOL | epenslamn Ri CEe ae tamariscus Williams chubutensis Granara de Willink and Miller, ~ Dorsal anal lobe without enlarged setae n. sp. but with cluster of about 10 flagellate se- = Translucent pores absent from hind legs; tae 3)3 rk aks ens ola % cuniculorum Williams multilocular pores absent from thorax 10(5) Dorsal oral-collar tubular ducts absent . . oF Ey ee Pe eee, eee coccineus (Newstead) ee eee esac oto oe eo ol 3(1) Translucent pores present on hind coxae — Dorsal oral-collar tubular ducts present SE Mepes eter SEene SNe ate eiaweecues oie Seek nae estcee 5 Se ee ee en COMP Dell MmNOsztdanal - Translucent pores absent from hind coxae 11(10) Translucent pores present on hind tibia 13 AERTS «epee Lee S Keates seetees Aha 4 - Translucent pores absent from hind tibia 4 (6) mediransluGentsporesspresen tao Mim G ete Triana Hence enone ae ee 12 and tibia blanchardii (King and Cockerell) 12(11) Ventral anal lobe with distinct anal bar; - Translucent pores absent from hind pair of most body setae elongate, longer than di- LESS Er ere eee tee antiscius Williams ameter of spiracular atrium ........ 56), Circulusslarcequadrate onslaterally vex- 9) ee are ee monodi Balachowsky and Ferrero VOLUME 106, NUMBER 1 Fig. 1. 13(11) 14(13) Ventral anal lobe without anal bar; most body setae short, shorter than diameter of spiracular atrium ..... sternlichti Williams Ventral oral-collar tubular ducts absent frOMMTMOTAx and wnead. yey ysis se 14 Ventral oral-collar tubular ducts present on thorax and head .. saudiensis Matile-Ferrero Anal-lobe cerarian setae conical; multiloc- ular pores restricted to posterior 2 or 3 ab- dominal segments ...... copallinae Ferris Anal-lobe cerarian setae flagellate, not conical; multilocular pores restricted to posterior 5 or 6 abdominal segments Abn eis Eien ne eee ee yuccae Ferris 143 AAAS a RR RES Sie 5 ac) Eurycoccus chubutensis, fourth-instar female (adult). Eurycoccus chubutensis Granara de Willink and Miller, new species (Figs. 1-3) Description.—Adult female: In life very convex, body yellowish, covered with white wax. Slide-mounted specimens (Fig. 1) circular or broadly oval, holotype 3.8 mm long (paratypes 2.5—4.3(3.6) mm), and 3.1 mm wide (paratypes 2.0—3.4(2.9) mm). Antenna well developed with 7- or 8-seg- ments, several specimens with third seg- 144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ment only partially divided, holotype anten- na 549 long (paratypes 429—585(519) wp). Legs well developed with numerous setae on all segments, with translucent pores on both surfaces of hind coxa and dorsal sur- face of hind femur; hind trochanter+femur of holotype 460 pw long (paratypes 331— 470(427) ww); femur of holotype 339p long (paratypes 315—352(338) w); tibia+tarsus of holotype 398 p long (paratypes 304— 415(370) p); tibia of holotype 281 p long (paratypes 195—284(249) jw); tarsus of ho- lotype 142 wp long (paratypes 133—140(137) wt); tibia considerably longer than tarsus, tibia/tarsus 2.0 in holotype (paratypes 1.4— 2.0(1.8)); and claw without a denticle on plantar surface; digitules on claw with clubbed apex; tarsal digitules with thin apex, not extending to tip of claw. With 2 pairs of ostioles, each ostiole lip with many trilocular pores (much more abundant than on surrounding derm) and setae, ostiole opening with sclerotized edge. Eyes on ven- ter near body margin, in 3 specimens a few trilocular or discoidal pores incorporated in sclerotization. Anal ring dorsal, near pos- terior end of body, oval, 132 wp long on ho- lotype (paratypes 74—138(119) ww) with 3 rows of pores, outer 2 rows weakly scler- otized, and 3 pairs of setae, longest seta on holotype 93 pw long (paratypes 91—122(106) w). Mouthparts well developed, clypeola- bral shield of holotype 363 pw long (para- types 319—385(352) yw), holotype with 14 setae on shield (paratypes 7—13(11) setae); labium 3-segmented, some specimens with narrow labium, others with structure broad, on holotype 382 w long (paratypes 380— 415(392) 2), basal segment with 4 setae on each side, middle segment with | seta on each side, and apical segment with 5 setae on each side and a group of 4 setae on each side along stylet canal at apex of labium; inner side of labium with | seta on each side in middle area of apical segment and 1 on each side near apex of apical segment. Circulus with well defined perimeter, divid- ed by intersegmental line between segments III and IV, rectangular, holotype 144 jw wide (paratypes 98—156(136) w). Spiracles con- spicuous, anterior pair on holotype 125 pw long (paratypes 185—230(201) w) and 82 wp wide (paratypes 185—230(201) w), anterior pair larger than posterior pair; trilocular pores and setae in membranous opening leading to spiracle, but not in sclerotized peritreme; often with few multilocular or trilocular pores incorporated into scleroti- zation on middle area of spiracle. Cerarii absent. Anal-lobe area without normal elongate anal-lobe seta, several se- tae at posterior apex slightly longer than re- maining dorsal setae, longest on holotype 86 pw long (paratypes 49—86(72) w). Dorsal surface: Trilocular pores round, with obliterated internal structure, about 4 w. in diameter, abundant over surface. Cone- shaped discoidal pores slightly larger than triloculars, abundant over surface. Dorsal setae flagellate, slightly enlarged basally, numerous, ranging from 15—42 wp long, about same length over surface except on segment VIII where longest setae range from 60 to 100 p long. Ventral surface: Trilocular pores abun- dant over surface. Cone-shaped discoidal pores most numerous along body margin, nearly absent medially. Small discoidal pores uncommon. Multilocular pores about 2 times larger than triloculars, about 7 p in diameter, present in medial and mediolat- eral areas from near vulva forward to area between mouthparts and front pair of legs, absent from marginal areas laterad of legs and spiracles. Setae flagellate, unusually short for mealybug, largest setae about 45 w long. Cisanal setae not always distin- guishable from other setae, about 75 p long, about same length as dorsal setae near anal ring. Invaginated pocket present in medial area on intersegmental line of segments VIII and IX, rectangular in shape, with def- inite opening. Material examined.—Holotype (IMLA) and 8 paratypes TMLA, USNM) from Ar- gentina, Chubut, Lago Puelo National Park, road to Los Hitos, on woody roots of Noth- VOLUME 106, NUMBER 1 ofagus dombeyii, Fagaceae, I-1999, Cuezzo y Granara de Willink. Description.—Second- (or third-) instar female (Fig. 2): Same as adult female ex- cept as follows: Body circular or broadly oval, 1.9 mm long, 1.3 mm wide. Antennae in only specimen available apparently ab- normal, 6 segments on | side and partially divided 7 segments on other, antenna 420 w long. Legs with hind trochanter+femur 275 w long; femur 207 pw long; tibia+tarsus 275 w; tibia 145 pw long; and claw 42 wp long; tibia slightly longer than tarsus, tibia/tarsus 1.1; digitules on tarsus and claw apparently broken. Each posterior lip of ostioles with many trilocular pores and | or 2 setae; an- terior lip usually without setae. Eyes with- out trilocular or discoidal pores incorporat- ed in sclerotization. Anal ring dorsal, near posterior end of body, oval, 83 long with 2 rows of pores, outer row weakly sclero- tized, and 3 pairs of setae (1 side of spec- imen abnormal, with 2 setae), longest seta 80 ww long. Mouthparts with clypeolabral shield with 8 setae, 225 pt long; labium 255 w long. Circulus 98 p wide. Anterior spi- racles 132 w long and 55 wp wide; trilocular pores in membranous opening leading to spiracle, but not in sclerotized peritreme. Cerarii absent. Anal-lobe area without normal, elongate anal-lobe seta, setae at posterior apex mostly broken. Dorsal surface: Trilocular pores about 3 2 in diameter, abundant over surface. Large discoidal pores slightly larger than trilocu- lars, without cone-shaped protrusion, abun- dant over surface. Small discoidal pores ab- sent. Dorsal setae ranging from 15—22 w long, about same length over surface except on segment VIII where longest setae about 34 p long. Ventral surface: Trilocular pores uncom- mon posteriorly and in medial areas. Large discoidal pores most numerous along body margin, nearly absent medially. Small dis- coidal pores absent. Multilocular pores about 2 times larger than triloculars, about 5 » in diameter, present in medial and me- diolateral areas of thorax. Setae about 34 wp 145 long. Cisanal setae about 33 p long. Invag- inated pocket may be represented by inter- nal, elongate oval structure visible near in- tersegmental line of segments VIII and IX, without definite opening. Notes.—The single specimen of this in- Star is in poor condition with many of the setae broken, the median area has a large cut and hole, and the abdomen is rolled. Thus, in some instances it was necessary to extrapolate patterns from both sides of the specimen. Material examined.—l specimen with same data as holotype (IMLA). Description.—First instar (sex not deter- mined) (Fig. 3): Same as adult female ex- cept as follows: Body oval, 0.9 mm long, 0.5—0.6(0.6) mm wide. Antenna 6-segment- ed, 290—300(297) ww long. Legs with hind trochanter+ femur 172—175(174) p» long; fe- mur 130—142(136) pw long; tibia+tarsus 182—210(198) yp; tibia 95—100(98) pw long; claw 38 p. Each posterior lip of ostioles with several trilocular pores; each anterior lip with or without large discoidal, several triloculars, and O—1(Q) seta. Eyes without trilocular or discoidal pores incorporated in sclerotization. Anal ring apical, oval, with 2 rows of pores, and 3 pairs of setae, lon- gest seta 80—112(97) wp long. Mouthparts with 6 setae on clypeolabral shield; 125— 142(133) pw long; labium 175—200(192) pw long. Circulus 65—78(72) wide. Anterior spiracle 52—75(64) w long and 32—35(34) ph wide; trilocular pores in membranous open- ing leading to spiracle, but not in sclero- tized peritreme. Cerarii absent. Anal-lobe area with nor- mal elongate anal-lobe seta, 125—150(138) p long. Dorsal surface: Trilocular pores about 3 w. in diameter, abundant over surface. Large discoidal pores without conical protrusion, slightly larger than triloculars, scattered over surface. Small discoidal pores absent. Dorsal setae ranging from 15—50 wp long, longest setae in posterior areas of abdomen. Ventral surface: Trilocular pores in small numbers in medial areas. Large dis- 146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Eurycoccus chubutensis, second- or third-instar female. VOLUME 106, NUMBER 1 147 Fig. 3. Eurycoccus chubutensis, first instar (sex undetermined). 148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON coidal pores most numerous along body margin, nearly absent medially. Small dis- coidal pores absent. Multilocular pores ab- sent. Setae ranging from 18—60 pw long. Cis- anal setae about 38—54(43) p long. Invag- inated pocket absent. Material examined.—This description is based on 3 embryos that are in poor con- dition with same data as holotype (IMLA). Biology.—The species was found on the woody roots of a large specimen of Noth- ofagus dombeyii (Mirb.) Oest. and was tended by the ant Camponotus chilensis Spinola. Comments.—Nothofagus is the only ge- nus in the family Fagaceae in the Southern Hemisphere and occurs in temperate areas of Australia, South America, New Caledon- ia, New Guinea, and New Zealand. Argen- tina has several native species of Nothofa- gus, but the scale insects that feed on them are virtually unknown. Williams (1985a) summarized available information on the Nothofagus scale fauna of South America. The scale-insect fauna of Nothofagus 1s probably best known in New Zealand, but even there the emphasis has been on four families, the coccids (Hodgson and Hen- derson 2000), eriococcids (Hoy 1962), mar- garodids (Morales 1991), and pseudococ- cids (Cox 1987) and other scale families are more poorly described. Eurycoccus chubutensis differs from oth- er species of Eurycoccus by having multi- ple setae on the clypeolabral shield, 4 setae on each side of the basal labial segment, large discoidal pores with a cone-shaped projection, an unusual invaginated pocket on the venter between segments VIII and IX, and multilocular and trilocular pores with obscured locular structure, and by lacking long anal-lobe setae and tubular ducts. Maskellococcus obtectus (Maskell) shows some resemblance to E. chubutensis by occurring on Nothofagus; having re- duced numbers of cerarii; and multilocular pores restricted to venter. Eurycoccus chu- butenis differs by lacking the narrow pear- shaped body characteristic of M. obtectus and in lacking tubular ducts. Neosimmondsia hirsuta Laing also re- sembles the new species because both lack cerarii, have numerous setae and trilocular pores, have translucent pores on the hind coxa, 2 pairs of ostioles, a circulus, and ventral multilocular pores on the venter only. Neosimmondsia hirsuta differs by having: tubular ducts; 6-segmented anten- na; dorsal setae that are as long as the anal- ring setae; no cone-shaped discoidals; no invaginated pocket; and elongate anal-lobe setae. Eurycoccus chubutensis has: no tu- bular ducts; 7- or 8-segmented antenna; dorsal setae that are shorter than the anal- ring setae; cone-shaped discoidal pores; an invaginated pocket; and no distinct anal- lobe setae that are longer than the other se- tae in the area. Pseudantonina Green Pseudantonina Green 1922; type species, Pseudantonina bambusae Green 1922, by monotypy. This genus includes 8 species (Ben-Dov 2002) that are recorded from several differ- ent zoogeographic regions (Neotropical 1, Nearctic 5, Oriental 1, and Palearctic 1) in- cluding Argentina (Pseudantonina aeria Williams and Granara de Willink). It is pos- sible that two or more unrelated groups are currently placed in the genus. Confusion about the status of several of the New World species is exemplified by Hendricks and Kosztarab’s (1999) treatment of P. aeria, P. arundinariae McConnell, P. gi- ganticoxa Lobdell, and P. texana Ferris as “Species of Uncertain Placement.” They also indicated that P. magnotubulata Bor- chsenius had been transferred to another ge- nus, but we can find no evidence that such action has taken place. Ben-Dov (2002), continued to include them in Pseudantoni- na, and Kosztarab (1996) mentioned P. arundinariae and P. giganticoxa as mem- bers of the genus along with two new spe- cies (P. nakaharai Kosztarab and P. wilkeyi VOLUME 106, NUMBER 1 149 Table 2. Distribution of characters considered important in diagnosing Pseudantonina. _—————— Hind Legs Distorted Many Transl. Pseudantonina Species Front Legs Pores Pores Near Hind Coxae Clustered Spirac. Pores Pairs of Cerarii Circulus Antennal Segs. rr aeria normal yes yes no no | absent 6 arundinariae small no no yes yes l absent 6 bambusae small yes no yes yes 0) absent 3-6 giganticoxa small yes yes no yes 0-1 absent 6 magnatubulata normal no yes no no 0) present 6 nakaharai normal yes yes no yes | absent 6 texana normal yes yes no yes ] absent 6 wilkeyi normal yes yes no yes 2or3 absent 6 Kosztarab). Although Hendricks and Kosz- 31) (Ginculustabsentaccss. os cece) eee ee 4 tarab (1999) was published after Kosztarab ~ Circulus present magnatubulata Borchsenius 4(3) Translucent pores absent from hind tibiae 5) (1996), it was essentially the same as the Hendricks dissertation completed three years earlier (Hendricks 1993). We have examined illustrations, descrip- tions, or specimens of all 8 Pseudantonina species (P.aeria, P. arundinariae, P. bam- busae, P. giganticoxa, P. magnatubulata, P. nakaharai, P. texana, P. wilkeyi) and note similarities that occur in nearly all spe- cies;(@iable-2). The new species possesses most of the characteristics commonly considered to be diagnostic of the genus and is therefore placed here until a detailed revision of Pseudantonina and relatives is undertaken. Other genera that share characters with Pseudantonina are: Antonina Signoret, Ac- rochordonus Cox, and Renicaula Cox, all of which occur on grasses. Antonina differs by lacking legs; Acrochordonus lacks pore clusters in or adjacent to the spiracular atria and has normally developed legs; and Ren- icaula lacks multilocular pores. KEY TO SLIDE-MOUNTED ADULT FEMALES OF THE GENUS PSEUDANTONINA —_— Multilocular pores not restricted to marginal ALCASTOMaVeNLCLies Mn 30-12) 6s ees s at Soc 3 _ Multilocular pores forming distinct band along ventral body margin 2(1) Pore cluster surrounding anterior spiracles without multilocular pores; middle legs with allRSEOmMentSi sere = arundinariae McConnell = Pore cluster surrounding anterior spiracles with multilocular pores; middle legs abor- bambusae Green i) tive, without all segments .. . — Translucent pores present on hind tibiae . vernacula Granara de Willink and Miller, n. sp. 5(4) Multilocular pores present on dorsal areas of thoraxqanG Wea ier ware eee eee a v7 = Multilocular pores absent from dorsal areas Ofsthoraxeandsheadaes estes eee 6 6(5) Anal ring small, with 2 or fewer rows of pores; trilocular pores sparsely scattered over body surfaces, absent from anterior lip of posterior ostiole cade” & aeria Williams and Granara de Willink - Anal ring large, with 3 or more rows of pores; trilocular pores abundant over body surfaces, present on anterior lip of posterior ostiole texana Ferris LE rash ovata hes a epee NS Se ORS ie Pia GREY eee 8 — Hind coxae longer than remainder of hind le Serie ety: ce ete ete Sta giganticoxa Lobdell 8(7) Anterior ostioles present; spiracles without pores in sclerotized atrium; anal-ring setae less than twice diameter of ring Ae ae or) ot fen ae nakaharai Kosztarab - Anterior ostioles absent; spiracles with pores in sclerotized atrium; anal-ring setae more than twice diameter of ring —-wilkeyi Kosztarab Pseudantonina vernacula Granara de Willink and Miller, new species (Figs. 4-7) Description.—Adult female: Found in grass sheath, body pink, lightly dusted with white wax. Slide-mounted specimens (Fig. 4) oval; length of holotype 2.7 mm long (paratype 2.9 mm), and 1.8 mm wide (paratype 1.9 mm), posterior abdominal segment narrow- 150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Pseudantonina vernacula, fourth-instar female (adult). VOLUME 106, NUMBER 1 est, partially sclerotized, with prominent lobes. Antenna 6-segmented, third segment partially divided on some specimens, ho- lotype antenna 167 w long (paratype 172 w.). Legs small compared to many mealy- bugs; with few setae; hind legs with tro- chanter and femur enlarged, with translu- cent pores on both surfaces of trochanter and femur, on dorsal surface of tibia; hind coxa noticeably smaller than trochan- ter+femur; trochanter of holotype 83 pw long (paratype with trochanter fused with femur); femur of holotype 54 wp long; tro- chanter+femur of holotype 127 wp long (paratype 82 w); tibia of holotype 37 wu (paratype 35 w); tarsus of holotype 32 pw (paratype 55 wy), tibia/tarsus on holotype 1.1 (paratype 0.6); tibia+tarsus of holotype 68 kt (paratype 90 yw); claw sometimes present; tarsal digitules thin, slightly thickened api- ces; claw digitules unequal or equal de- pending on specimen, thin, slightly thick- ened apices. Mouthparts well developed (but absent on paratype), clypeolabral shield of holotype 147 pw long, holotype with 2 setae on shield; labium 3-segmented, relatively short and broad, on holotype 59 w long, basal segment with 3 setae on each side, middle segment with | seta on each side, and apical segment with 2 setae on each side and a group of 3 or 4 setae on each side along stylet canal at apex of la- bium; inner side of labium without setae. With posterior pair of ostioles only, with lips sclerotized, usually without pores or se- tae, occasionally with anterior lip with | tri- locular pore. Anal ring invaginated in pock- et near posterior end of body; diameter on holotype 61 pw (paratype 72 w), with 2 rows of pores and 6 setae, longest seta on holo- type about 135 p long, broken on paratype. Circulus absent. Spiracles large, in sclero- tized invagination, peritreme large, with more than 20 multilocular pores in sclero- tization of each spiracle. Cerarii absent (depending on definition of cerarius) or reduced to 2 indistinct pairs on posterior abdominal segments. Dorsal anal-lobe conspicuous, protruding, sclero- 151 tized, with 14 slightly enlarged, curved se- tae, not clustered, 2 or 3 scattered trilocular pores near edge of sclerotization, and con- spicuous basal sclerotization, without basal clusters of pores or auxiliary setae, sclero- tization covering half of segment VIII; seg- ment VII with 3 or 4 slightly enlarged setae, with basal, triangular shaped sclerotized area. Marginal areas of segments IV, V, and VI each with | slightly enlarged seta, with- out associated trilocular pores or scleroti- zation. Ventral surface of protruding lobes sclerotized in lateral and medial area with 4 small setae and cluster of multilocular pores; without elongate anal-lobe seta. Dorsal surface: With numerous oral-col- lar tubular ducts on surface, length about 12 yw, diameter at dermal orifice about 2 p, diameter at inner end about 5 p, with glan- dular filament. Setae short, flagellate, thin varying from 10-18 wp long. Trilocular pores small, with thick walls, scattered over surface. Ventral surface: Setae flagellate, short; setae around vulva undifferentiated; cisanal setae, short about 25 w long, 2 pairs. Tri- locular pores primarily in lateral areas, un- common. Multilocular pores with 9 or 10 loculi forming a submarginal line from pos- terior spiracle to abdominal apex, also pre- sent on head and near anterior spiracles. Discoidal pores rare. Oral-collar tubular ducts similar to dorsal ducts, scattered over surface, less abundant than on dorsum. Material examined.—Holotype adult fe- male from: Argentina, Neuquén, on Ruta 40 between La Angostura and Confluencia, 4- I-99, on Gramineae, possibly Stipa, Granara de Willink (IMLA). Paratype female with same data in USNM. Description.—Third-instar female (Fig. 5): Same as adult female except as follows: Slide-mounted specimen elongate oval; length 1.7 mm long, 0.6 mm wide, posterior abdominal segment with sclerotization and slightly protruding anal lobes. Antenna 6- segmented, 165 and 167 pw long. Legs well developed, without translucent pores; hind trochanter+femur 88 and 92 pw long; tibia Figs: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Pseudantonina vernacula, third-instar female. VOLUME 106, NUMBER 1 48 and 50 pw long; tarsus 50 and 51 p long; tibia+tarsus 99 and 100 pw long; tibia/tarsus 0.9 and 1.0; tarsal digitules very different in size, 1 conspicuous and extending be- yond tip of claw with small apical club, oth- er setose without club not reaching tip of claw; claw digitules equal or unequal de- pending on specimen, with apical club. Mouth parts well developed, labium short and broad, with 3 segments, 50 p long; cly- peolabral shield 138 p long. With posterior pair of ostioles only, without pores or setae. Anal ring invaginated, near apex of abdo- men; diameter 45 jw; with 2 row of pores; anal ring setae with narrow apices, longest seta 62 w long. Spiracles with enlarged per- itreme, with at least 3 or 4 trilocular pores in peritreme sclerotization. Cerarii present from segments VI to VIII; on segment VI represented by | slightly en- larged seta, no basal sclerotization; on seg- ments VII with 2 or 3 associated enlarged setae and | trilocular pore, no basal scler- otization; on segment VIII with 4 associ- ated setae, no trilocular pore, and_ basal sclerotization dispersed beyond base of cer- arian setae. Anal lobes protruding, sclero- tized, ventral surface with | small setae, with elongate anal-lobe seta 70 and 78 pw long, 2 or 3 marginal enlarged setae, and 1 trilocular pore. Dorsal surface: With more than 3 lon- gitudinal lines of flagellate setae on each side of abdomen excluding cerariian setae; trilocular pores most abundant along pos- terior body margin, with | or 2 on head and thoracic body margin. Setae and tubular ducts scattered over surface. Tubular ducts decreasing in size anteriorly. Ventral surface: With 3 to 5 longitudinal lines of flagellate setae on each side of body; 2 pairs of cisanal setae, short, about 18 wp long. Quinquelocular pores absent. Trilocular pores along body margin, in spi- racular atrium, absent from head. Multiloc- ular pores present on head and thorax near legs and spiracles and along body margin of abdomen. Discoidal pores inconspicu- ous, | longitudinal line on submargin of 153 body. Oral-collar tubular ducts present near body margin. Material examined.—This description is based on | specimen with same data as ho- lotype (IMLA). Description.—Second-instar female ? (Fig. 6): Same as adult female except as follows: Slide-mounted specimens elongate oval; length 1.0—1.1(1.1) mm long, 0.3-— 0.4(0.4) mm wide, posterior abdominal seg- ment with sclerotization and slightly pro- truding anal lobes. Antenna 6-segmented, 150—162(157) » long. Legs well developed, without translucent pores; hind trochan- ter+femur 90—92(91) w long; tibia 58— 61(60) p long; tarsus 50—58(54) w long: tibia+tarsus 105—112(110) p long; tibia/tar- sus 1.0—1.2(1.1); tarsal digitules very dif- ferent in size, | conspicuous and extending beyond tip of claw with small apical club, other setose without club not reaching tip of claw; claw digitules unequal or equal de- pending on specimen, with apical club. Mouth parts well developed, labium short and broad, with 3 segments, 45—48(47) w long; clypeolabral shield 115—125(119) pw long. With posterior and anterior pairs of ostioles, without pores or setae on posterior ostioles, anterior ostioles with 1 seta on posterior lip. Anal ring slightly invaginated, apical; diameter 32—35(33) w; with 2 row of pores; anal ring setae with blunt apices, longest seta 35—41(38) pw long. Spiracles with slightly enlarged peritreme, with at least 1 trilocular pore in peritreme sclero- tization. Cerarii present from segments I or II to VIII; on segments I to VI each represented by | slightly enlarged seta, | widely spaced slender seta and | associated trilocular pore, no basal sclerotization; on segments VII with 2 closely associated setae and | triloc- ular pore, no basal sclerotization; on seg- ment VIII with 2 closely associated setae, | trilocular pore, and basal sclerotization dispersed beyond base of cerarian setae. Anal lobes slightly protruding, sclerotiza- tion more broadly dispersed, ventral surface with | small seta, with elongate anal-lobe 154 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Pseudantonina vernacula, second-instar female ?. VOLUME 106, NUMBER 1 seta 55—78(68) » long, and with | trilocular pore. Dorsal surface: With 3 longitudinal lines of flagellate setae on each side of abdomen excluding cerariian setae; trilocular pores primarily along body margin. Setae and tri- locular pores more abundant on thorax and head. Tubular ducts marginal. Ventral surface: With 3 to 5 longitudinal lines of flagellate setae on each side of body; 2 pairs of cisanal setae, short about 18 wp long. Quinquelocular pores absent. Trilocular pores along body margin, in spi- racular atrium. Multilocular pores present on head and thorax near legs and spiracles and along body margin of abdomen. Dis- coidal pore forming | longitudinal line on submargin of body. Oral-collar tubular ducts present near body margin. Material examined.—This description is based on 3 specimens with same data as holotype (IMLA). Description.—First instar (Fig. 7): Same as adult female except as follows: Slide- mounted specimens elongate oval; length 0.7-0.9(0.8) mm long, 0.2—0.5(0.4) mm wide, posterior abdominal segment with slight sclerotization; without protruding anal lobes. Antenna 6-segmented, 140— 168(160) w long. Eye present near base of antenna. Legs well developed, without translucent pores; hind trochanter+femur 92—102(96) pw long; tibia 58—65(63) pw long; tarsus 52—65(61) p long; tibia+tarsus 1 15— 125(121) p long; tibia/tarsus 1.0—1.2(1.0); tarsal digitules very different in size, | con- spicuous and extending beyond tip of claw with small apical club, other setose without club not reaching tip of claw; claw digitules unequal or equal depending on specimen, | thicker than other, with apical club. Mouth parts well developed, labium short and broad, with 3 segments, 32—45(40) p long; clypeolabral shield 90—110(104) w long. With posterior and anterior pairs of ostioles, with O—1(1) trilocular pore on posterior lip of each ostiole, without setae. Anal ring not invaginated, apical; diameter 28—31(30) p; with 2 rows of pores; anal ring setae with 155 blunt apices, longest seta 28—35(30) pw long. Spiracles without enlarged peritreme, with- out pores in peritreme sclerotization. Cerarii present from segments I or II to VIII; on segments I to V each represented by 1 slightly enlarged seta, | widely spaced slender seta and | associated trilocular pore, no basal sclerotization; on segments VI and VII with 2 closely associated setae and | trilocular pore, no basal sclerotization; on segment VIII with 2 closely associated se- tae, 1 trilocular pore, and basal sclerotiza- tion restricted to area near base of cerarian setae. Anal lobes not protruding, scleroti- zation confined to cerarius, ventral surface with | small seta, with elongate anal-lobe seta 80—115(103) p long, and without tri- locular pores. Dorsal surface: With 2 longitudinal lines of flagellate setae on each side of abdomen excluding cerarlian setae and | longitudinal line of trilocular pores on each side of ab- domen excluding pore loosely associated with cerarii. Setae and trilocular pores more abundant on thorax and head. Tubular ducts and discoidal pores absent. Ventral surface: With 3 longitudinal lines of flagellate setae on each side of body; 2 pairs of cisanal setae, short, 18— 24(21) ww long. Quinquelocular pores in submarginal longitudinal line on abdomen, also present medially on thorax and head. Trilocular pores rare, present near spiracles. Multilocular pores present on head and tho- rax near legs and spiracles. Discoidal pore forming | longitudinal line on each sub- margin of body. Oral-collar tubular ducts present near body margin. Material examined.—This description is based on 10 specimens that are in reason- able condition with same data as holotype (IMLA). DISCUSSION One of the more interesting findings while preparing this paper was the discov- ery of additional setae on the basal segment of the labium and on the clypeolabral shield. According to Koteja C1974) oall 156 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Pseudantonina vernacula, first instar (sex undetermined). VOLUME 106, NUMBER 1 mealybugs have 3 setae on each side of the basal segment of the labium. In EF. chubu- tensis there clearly are 4 in all instars ex- amined. Although we have looked at hun- dreds of illustrations, descriptions, and specimens of a diverse array of mealybugs, we have not located other species with 4 setae on the basal segment of the labium but feel certain that they exist. We have, however, discovered two species that are exceptional in that they have only 2 setae on each side of the basal labial segment, i.e., Plotococcus minutus Williams and Granara de Willink (Williams and Granara de Willink 1992) and Macrocepicoccus lor- anthi Morrison (Miller and Denno 1977). Unfortunately, the number and distribution of setae on the mouthparts were not consid- ered important until the work of Koteja (1974), and many descriptions and illustra- tions either do not show them at all, or de- pict them inaccurately. Further, most mealybug species that we have examined have a total of 2 setae on the clypeolabral shield. The few exceptions found are in the so called anomalous mealy- bugs such as Allomyrmococcus acariformis Takahashi (with more than 30 clypeolabral setae), Xenococcus annandalei Silvestri (with 4 such setae) (Williams 1978), and several species of Eumyrmococcus (with 4 setae) (Williams 1998). In FE. chubutensis there may be from 7 to 14 setae on the cly- peolabral shield in the adult female and at least 6 in the immatures. It will be interest- ing to see if more species of South Amer- ican mealybugs have this unusual charac- teristic. The invaginated pocket posterior of the vulva on Eurycoccus chubutensis is another structure not commonly reported in the mealybug literature. Similar structures were illustrated in Rhizoecus by Williams (1996). ACKNOWLEDGMENTS We are grateful to the National Council of Scientific researches and Technics (CONICET), Agency of Promotion and Technological Development (FONCyT), 157 Advice of Investigations of the National University of Tucuman (CIUNT) for sup- port of this project, and to Dra. Fabiana Cuezzo (Instituto Superior de Entomologia, UNT, ARG) for ant identifications. We also thank Douglas J. Williams (Department of Entomology, The Natural History Museum, London, UK), Norman E. Woodley (Sys- tematic Entomology Laboratory, USDA, Washington, DC, USA), and David A. Nickle (Systematic Entomology Laborato- ry, USDA, Beltsville, MD, USA) for their comments and reviews of the manuscript. LITERATURE CITED Ben-Dov, Y. 2002. Pseudococcidae Jn Scalenet.http:// www.sel.barc.usda.gov.scalenet/scalenet.htm Cox, J. M. 1987. Pseudococcidae (Insecta: Hemiptera). Fauna of New Zealand. No. 11. DSIR Science In- formation Publishing Centre, Wellington, New Zealand, 229 pp. Ferris, G. F 1950. Atlas of the Scale Insects of North America. (ser. 5) [v. 5]. The Pseudococcidae (Part I). Stanford University Press, Palo Alto, Califor- nia, 278 pp. Granara de Willink, M. C. 1979 (1978). Contribucion al estudio de las cochinillas harinosas (Homop- tera: Pseudococcidae) en citricos de la Provincia de Tucuman. Jornados Fitosanitarias Argentinas |: 125-160. . 1981. Nueva especies de Hypogeococcus Rau de Tucuman, Republica Argentina (Homoptera Pseudococcidae). Neotropica 27: 61—65. . 1983a. Spilococcus radicalis nueva especies de Pseudococcido de la Republica Argentina (Hom. Pseudococcidae). Acta Zoologica Lilloana 37: 55-58. . 1983b. Tres especies de Phenacoccus de la Provincia de Tucumaan, Argentina (Hom. Pseu- dococcidae). Acta Zoologica Lilloana 37: 59-64. . 1986. A new species of Cryptoripersia from Argentina (Homoptera: Pseudococcidae). Acta Zoologica Lilloana 38: 129-132. . 1991a. Cochinillas harinosas de importancia economica encontradas en la Argentina: actuali- zacion sistematica y nueva lista de hospederos. Boletin de la Academia Nacional de Ciencias (Cordoba, Argentina) 59(3/4): 259-271. . 1991b. Una nueva especie de Ferrisia Fulla- way en la Republica Argentina (Homoptera: Pseu- dococcidae). Insecta Mundi 5(3—4): 181-184. . 1991c. Cochinillas harinosas de importancia economica encontradas en la Argentina: Actuali- zaciOn sistematica y nueva lista de hospederos. 158 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Boletiin de la Academia Nacional de Ciencias. (Cordoba, Argentina) 59: 259-271. Green, E. E. 1922. The Coccidae of Ceylon, Part V. Dulau & Co., London, 472 pp. Hendricks, H. 1993. Revision of the Tribe Serrolecan- iini Shinji (Pseudococcidae) with discussion on other legless mealybugs. Ph.D. Dissertation, De- partment of Entomology, Virginia Polytechnic In- stitute and State University, Blacksburg, Virginia, 262 pp. Hendricks, H. and M. Kosztarab. 1999. Revision of the Tribe Serrolecaniini (Homoptera: Pseudococci- dae). de Gruyter, Berlin & New York. xiv, 213 pp. Hodgson, C. J. and R. C. Henderson. 2000. Coccidae (Insecta: Hemiptera: Coccoidea). Fauna of New Zealand; No. 41. Manaaki Whenua Press, Lincoln, Canterbury, NZ, 259 pp. Hollinger, A. H. 1916. The shell-bark hickory mealy- bug. Canadian Entomologist 48: 411—413. Hoy, J. M. 1962. Eriococcidae (Homoptera: Coccoi- dea) of New Zealand. New Zealand Department of Scientific and Industrial Research Bulletin 146: 1-219. Kosztarab, M. 1996. Scale insects of Northeastern North America. Identification, biology, and distri- bution. Virginia Museum of Natural History, Mar- tinsburg, Virginia, 650 pp. Koteja, J. 1974. Comparative studies on the labium in the Coccinea (Homoptera). Zeszyty Naukowe Akademii Rolniczej] w Warszawie, Rozprawy Naukowe 89: 1—162. Leonardi, G. 191 la. Contributo alla conoscenza delle coccinglie della Republica Argentina. Bollettino del R. Laboratorio di Entomologia Agraria di Por- tici 5: 237-284. . 1911b. Contributo alla conoscenza delle coc- cinglie della Republica Argentina. Bollettino della Real Scuola Superiore di Agricoltura. Portici (ser 2) 10: 3-50. McKenzie, H. L. 1967. Mealybugs of California with taxonomy, biology, and control of North Ameri- can species (Homoptera: Cooccoidea: Pseudococ- cidae). University of California Press, Berkeley, 526 pp. Miller, D. R. and R. EK Denno. 1977. A new genus and species of mealybug with a consideration of mor- phological convergence in three arboreal species (Homoptera: Pseudococcidae). Systematic Ento- mology 2: 111-157. Miller, D. R. and A. Y. Rossman. 1995. Systematics, biodiversity, and agriculture: Systematic analyses of small but important organisms provide crucial information for improvement of agriculture. Bio- science 45(10): 680—685. Morales, C. EF 1991. Margarodidae (Insecta: Hemip- tera). Fauna of New Zealand. No. 21. DSIR Plant Protection, Auckland, New Zealand, 123 pp. Williams, D. J. 1973. Two cactus-feeding mealybugs from Argentina (Homoptera, Coccoidea, Pseudo- coccidae). Bulletin of Entomological Research 62: 565-570. . 1978. The anomalous ant-attended mealybugs (Homoptera: Pseudococcidae) of South-East Asia. Bulletin of the British Museum (Natural History) Entomology 37: 72 pp. . 1985a. Some South American scale insects (Homoptera: Coccoidea) on Nothofagus. Journal of Natural History 19: 249-258. . 1985b. A new species of Ferrisia (Hemiptera: Pseudococcidae) on cassava (Manihot esculenta) from Paraguay and on ragweed (Ambrosia tanui- folia) from Argentina. Bulletin of Entomological Research 75: 255-257. . 1985c. Australian mealybugs. British Muse- um (Natural History). 431 pp. . 1996. Four related species of root mealybugs of the genus Rhizoecus from east and southeast Asia of importance at quarantine inspection (He- miptera: Coccoidea: Pseudococcidae). Journal of Natural History 30: 1391-1403. . 1998. Mealybugs of the genera Eumyrmococ- cus Silvestri and Xenococcus Silvestri associated with the ant genus Acropyga Roger and a review of the subfamily Rhizoecinae (Hemiptera, Coc- coidea, Pseudococcidae). Bulletin of the Natural History Museum. Entomology Series (London) 67: 1-64. Williams, D. J. and M. C. Granara de Willink. 1992. Mealybugs of Central and South America. CAB International, London, England, 635 pp. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 159-165 DOLERUS ANATOLH, N. SP., THE FIRST PALEARCTIC MEMBER OF THE SUBGENUS NEODOLERUS GOULET (HYMENOPTERA: TENTHREDINIDAE) MIKK HEIDEMAA AND ALEXEY ZINOVJEV (MH) Institute of Zoology and Hydrobiology, University of Tartu, Vanemuise 46, 51014 Tartu, Estonia, c/o Institute of Plant Protection, Estonian Agricultural University, 50412 Tartu, Estonia (e-mail: mikk@eau.ee); (AZ) Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, St. Petersburg 199034, Russia (e-mail: zag-entom @ yahoo.com) Abstract.—A new tenthredinid, Dolerus (Neodolerus) anatolii Heidemaa and Zino- vjev, n. sp., is described from the Russian Far East and South Korea. It is the first species of the subgenus Neodolerus Goulet from outside the Nearctic Region. The subgenus now includes eleven Nearctic and one Palearctic species. Shared characters between Neodol- erus species and the Palearctic Dolerus (Poodolerus) vulneratus Mocsary are discussed. Key Words: Dolerus vulneratus, Dolerus shanghaiensis, geographic distribution, Hol- arctic, Hymenoptera, Neodolerus, new species, Poodolerus, sawfly, taxon- omy, Tenthredinidae, zoogeography Neodolerus Goulet, 1986 (type species Dolerus sericeus Say, 1824), a subgenus of the Holarctic genus Dolerus Panzer, 1801, has been known from 11 Nearctic species. Together with the nitens group, which cor- responds to Poodolerus Zhelochovtsev, 1988, it is regarded as an early lineage of Dolerus by Goulet (1986). Both are char- acterized by a shared apomorphic character, the relatively long sternum 9 in males (short in Selandriinae). Neodolerus is defined by the following synapomorphies: a deeply outlined furrow on the outer surface of the metatibia, a large flat surface at the poste- rior angle of the median lobe (small in other Selandriinae), and the markedly raised pos- teriomedial surface of tergum 8 in the males. Here, we describe a new species of Neo- dolerus from the Russian Far East and South Korea. It does not fit any Dolerus species known from the Palearctic (Zhelo- chovtsev 1928, 1935; Malaise 1931; Muche 1965; Haris 1996; Wei 1997; Wei and Nie 1997; Haris 2000, 2001), and it is not one of the Nearctic species treated by Goulet (1986). The species is named after the out- standing Russian symphytologist, Anatolii Nikolaevich Zhelochovtsev (1903-1976), who was the first to recognize it as a new species but never named or published on it. MATERIAL AND METHODS The description of new species is based on 3 female specimens, 2 collected in the Russian Far East and | in South Korea. The morphological terminology follows Goulet (1986) and Viitasaari (2002). The right half of valvula | (lancet) of the holotype and the left half of one paratype were dissected, processed in 10% KOH, and mounted in Canada balsam (holotype) and euparal (paratype) between rectangular cover slides. They were then placed in paper frames and pinned with the corresponding specimen. Measurements were made on ail three spec- 160 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON [Sudzuch. Zap. = Lazovsky Reservation] [Ta-Chingou-za Bay = Proselochnaya Bay] [24.-27.06.1948 A. Sharov leg.] Dol. n. sp. pr. [prope = near] taeniatus 1 arom Il Ge D: Dolerus anatolii, holotype female. Labels imens. The upper and lower limit for each measurement is given in the description. The line drawings of valvula | are from digital micrographs taken with an Olympus BX50 System Microscope and Olympus DP11 Camera. Other illustrations are from micrographs taken with an Olympus stereo- microscope SZX9 and digital camera C- 4040Z00M. Dolerus (Neodolerus) anatolii Heidemaa and Zinovjev, new species — (Figs. 1-10) Type material.—Holotype: female in coll. A. N. Zhelochovtsev, Zoological Mu- seum of Moscow State University (Russia); with three labels (labels 1 & 2 in Fig. 1.): [Sudzuch. Zap. = Judzuhe Zapovednik, now the Lazovsky Reservation, Ta-Chin- gou-za (= Ta-Chingou-za Bay, now Prose- lochnaya Bay, approximately in the middle between Valentin 44.12N; 134.30E and Preobrazheniye 42.90N; 133.91E), 24- 27.06.1948, A. Sharov leg. (pale label, par- tially printed, partially handwritten, in cy- rillic)], “‘Dol. n. sp. [LB = line break] pr [= prope] taeniatus”’ [in pencil, authentic handwriting of A. N. Zhelochovtsev]; ‘“HOLOTYPUS 2 DOLERUS ANATOLI HEIDEMAA & ZINOVJEV 2003” [red handwritten label]. Flagellomeres 5—7 of the right antenna are missing, the remaining flagellomeres of both antennae are glued on a piece of paper placed on the same pin as the specimen. The preparation of the right half of valvula 1 bears the handwritten code: “Dol. sp. 6493-3,” and is pinned with the’ specimen) PARATYPES @ 2): 2 with two labels ““RUSSIA: Primorskiy kray [LB] Sikhote-Alin Biol. Station [LB] 30 km SE Chuguevka [LB] 44.05 N 134.12 E [LB], 6 3:V.2983' » 6508 anit Bless A.TAEGER” [white printed label]; ““PAR- ATYPUS 2 DOLERUS ANATOLI HEI- DEMAA & ZINOVJEV 2003” [red hand- written label]. The preparation of the left half of valvula | is pinned with the speci- men. Deposited in the Deutsches Entomo- logisches Institut, Eberswalde, Germany; | 2 with two labels: ‘“‘Mirugam (Pugdaesa) [128.57 E; 37.80 N] [LB] 1300m Mt. Odae- san [LB] Kangwondo. Korea [LB] 28.V.1991 [LB] A. Shinohara” [white print- ed label, the date partially handwritten]; “PARATYPUS 2% DOLERUS ANATOLI HEIDEMAA & ZINOVJEV 2003” [red handwritten label]. The left flagellum, mid- tibia, and midtarsus are missing. Deposited in the Department of Zoology, National Science Museum (Nat. Hist.), Tokyo, Japan. Female.—Body black. Wings hyaline, slightly infuscated towards apices. Abdo- men and legs of holotype somewhat brown- ish, probably due to storage conditions of specimen. Body length 9.6—10.2 mm. Dis- tance from distal margin of tegula to pter- ostigma 5.0—5.55 mm. Head: Width at level of compound eyes 2.35-2.52 mm, maximum width behind eyes 2.26-2.41 mm (measured in dorsal view). Head behind eyes subparallel in dor- sal view (Fig. 2). Length of head behind compound eye about 0.4 (25/60) of length of eye in dorsal view (head positioned so hind margins of lateral ocelli and com- VOLUME 106, NUMBER 1 pound eyes aligned). Setae from silvery to brownish silvery and from brownish silvery to brownish on labrum and mandible. Lon- gest setae on postocellar area shorter than diameter of middle ocellus, on frons in front of middle ocellus setae about as long as ocellus diameter. Setae on paraantennal field up to 0.11 mm long. Antennal setae blackish, on scape up to 0.11 mm long. Length of antennal segments measured in mm: scape (with radicula) 0.49—0.50; ped- icel 0.27—0.32; flagellomere 1—0.83—0.95, 2—0.69-—0.81, 3—0.60—0.70, 4—0.51- 0.60, 5—0.44—-0.52, 6—0.40-0.45, 7— 0.38—0.45. Upper area between ocelli and compound eye somewhat depressed. Punc- tures behind eyes mostly separated, only some in upper third (just behind eye) fused. Punctures between compound eyes with glabrous interspaces narrower than punc- ture diameter, forming meshlike sculpture. Punctures on genae and temples less dense, with glabrous interspaces; puncture diame- ter on temples 0.02—0.06 mm. Postocellar area slightly raised, convex, with longitu- dinal median groove indistinct (holotype, | paratype) or missing (1 paratype), bordered by distinct parallel or slightly convergent postocellar furrows. OCL: 0.45—0.49, OOL: 0.55—0.59, POL: 0.28—0.31 mm. Length of left compound eye 0.67—0.73, height 0.96— 1.05 mm. Posterior part of vertex with transverse depression bordered posteriorly with occipital carina. Occipital carina more distinct at vertex and gena level. Clypeus slightly asymmetric, with right lobe more prominent than left lobe (Fig. 3). Clypeus with indistinct transverse carina and irreg- ular punctures, its slightly triangular emar- gination about 0.8 as deep as its length. Distance from lower margins of toruli to lower margin of clypeus 0.54—0.60 mm. Thorax: Lateral half of median lobe with 10-14 large punctures. Middle part of me- dian lobe laterally from median mesoscutal groove longitudinally depressed. Punctures on mesoscutum fine, more or less regular, with glabrous interspaces mostly wider than puncture diameter. Punctures on anterior 161 and central parts of mesoscutellum smaller and sparser than on posterior and lateral parts. Flat surface at posterior angle of me- dian lobe glabrous or slightly alutaceous. Setae on upper part of thorax silvery to brownish, longer than those on upper head. Punctures on mesopleuron up to 0.13 mm, often polygonal, with glabrous ridges of un- even height between them (Fig. 4). Setae on mesopleuron up to 0.14 mm long. Lat- eral portions of pectus surface rather flat. Pectus with small and sparse punctures in middle and posterior parts, in its anterior part punctures larger (up to 0.06 mm) and denser; pectus microsculpture indistinct, scalelike. Anterior part of thorax at border of mesopleuron and pectus without punc- tures, but with some scalelike microsculp- ture. Distance between cenchri 0.50—0.59 mm. Lateral parts of mesoscutellar append- age transversely concave, meshed. Metepi- meron and metepisternum with irregular rough sculpture, their shape as in Fig. 5. Outer surface of metacoxa with numerous sparse punctures, denser and larger at base than at apex; microsculpture scalelike, rath- er obsolete. Basal upper corner of metacoxa glabrous. Abdomen: Terga shiny. Tergum | with at least some small punctures. Macrosculpture on next terga obsolete, indistinct punctures starting from tergum 5. Sculpticells on terga 1—5 scalelike, obsolete. Terga 1—4 dorsally almost without setae, more numerous and dense from tergum 5. Silvery-gray setae cover all sterna and lateral parts of all terga entirely. Brownish setae may be present on terga 8-9. Valvula 3 (apical sheath) in lat- eral view about as long (1.0—1.10 mm) as valvifer 2 (basal sheath), its shape as in Fig. 6. Longest setae of valvula 3 (sensory tuft) 0.29—0.33 mm, curved at apical third and forming ca. 30° angle in dorsal view (Fig. 7). Cercus 0.17—0.19 mm long. Setae on cercus and valvula 3 brownish. Valvula | (lancet) with 16 segments, apical half dor- sally concave (Fig. 8). Annuli of middle segments outlined by stiff setalike sensilla; winglike processes of annuli absent; serru- 162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-7. Dolerus anatolii, holotype/paratype female. 2, Head in dorsal view (holotype). 3, Outline of clypeus (holotype). 4, Mesopleuron surface (holotype). 5, Metepimeron and metepisternum (holotype). 6, Apical part of abdomen in lateral view (paratype). 7, Valvula 3 (apical sheath) in dorsal view (holotype). VOLUME 106, NUMBER 1 Figs. 8-10. Dolerus anatolii, holotype female. 8, Valvula | (lancet) with details of 2 serrulae. 9, Apical part of the valvula 1. 10, Basal part of the valvula 1. lae as in Fig. 9. Apical and basal parts of valvula 1 as in Figs. 9-10. Serrulae 1-3 without teeth, the rest with approximate number of teeth as follows: 4:4; 5:5; 6:7; 7: Seo asee oO: LO29° 1S: 12:7: 13:9: 14:6: 15:4; 16:1 (serrulae numbered from base of valvula | to apex). Male.—Unknown. Host plant.—Unknown. Distribution.—Eastern Asia: southern Russian Far East and South Korea. DISCUSSION The holotype female of the species bears a handwritten label by A. N. Zhelochovtsev (Fig. 1.) indicating that he was planning to describe it as a new species. However, there is no indication that he noticed its resem- blance to any Nearctic species. Superficial- ly, D. anatolii resembles D. pachycerus Hartig, 1837 (= D. taeniatus Zaddach, 1859), of the subgenus Juncilerus Zhelo- chovtsev (= Achaetoprion Goulet), but it differs by the ovipositor and presence of a furrow on the metatibia. Dolerus anatolii, possessing the following combination of characters, does not fit any of Zhelochov- tsev’s subgenera: comparatively long ovi- positor, valvula | without lateral teeth, and 164 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the enlarged irregular punctures on the me- sopleuron. Within the subgenus Neodole- rus, Dolerus anatolii is closest to D. par- asericeus MacGillivray, 1908; however, D. anatolii differs by the mostly separated punctures behind the compound eyes with only some in the upper third (just behind the eyes) fused, the mesoscutellar append- age transversely concave, outer surface of the metacoxa with numerous punctures, and the lamnium with 16 segments. The host plant is known for only one Neodolerus species, Dolerus sericeus Say, which feeds on wheat and probably on oth- er grasses (Goulet 1986, Leblanc and Gou- let 1992). Until now, Neodolerus was con- sidered endemic to North America. With the discovery of D. anatolii in the eastern Palearctic the possibility of a Palearctic or- igin for Neodolerus cannot completely be excluded. Presence of Neodolerus in east- ern Palearctic is a sequent example reflect- ing the well-known relationships between the temperate fauna of eastern Asia and eastern North America. Because many sci- entists studying eastern Asiatic species were unaware of the eastern North Ameri- can taxa, they often described new genera. Goulet (1986) proposed that Neodolerus might be a lineage within his nitens group (Poodolerus Zhelochovtsev). Poodolerus is distributed mostly in the Palearctic (over 55 species in western Palearctic), with seven Nearctic and one Holarctic species (D. as- per Zaddach, 1859), and one species intro- duced into North America (D. nitens Zad- dach, 1859). Discussion of the phylogenetic relationships within Poodolerus is not with- in the scope of this paper, but it is note- worthy that characteristic features of Neo- dolerus occur in some species of Poodole- rus, for example, in D. vulneratus Mocsary, 1878. This species has two characters in common with Neodolerus species: the dis- tinctly outlined furrow on the outer surface of the metatibia and the large flat triangular surface at the posterior angle of the median lobe (3 females were examined). Dolerus vulneratus is a rare Palearctic species known from Estonia (Viitasaari et al. 1998), Russian Karelia (Lindqvist 1969), Siberia, the Russian Far East (Zhelochovtsev and Zinovjev 1996), and South Korea (Haris 2001). On the other hand, the penis valve of D. vulneratus 1s very different (Zhelochovtsev 1988: 181, fig. 10) from all Neodolerus spe- cies and resembles species of the Dolerus alpinus and D. affinis groups. We did not study any material of Dolerus shanghaien- sis Wei and Nie (the name is preoccupied by Dolerus shanghaiensis Haris, 1996) that according to its description and illustrations of the ovipositor and penis valve first re- sembled a Neodolerus species. However, the specimens of its type series show no furrow on the outer surface of the metatibia (Meicai Wei, personal communication). This species most probably belongs to the subgenus Poodolerus. Further study incorporating molecular methods may shed more light on the phy- logenetic relationships between the Neodol- erus and allied Poodolerus species. ACKNOWLEDGMENTS David R. Smith (Systematic Entomology Laboratory, USDA, National Museum of Natural History, Washington, D.C.) and Henri Goulet (Canadian National Collec- tion of Insects, Ottawa) commented on and improved the manuscript. Irina Kadis (Ar- nold Arboretum, Harvard University, Cam- bridge, Massachusetts) improved the style. Alexander V. Antropov (Zoological Muse- um of Moscow State University, Moscow), Akihiko Shinohara (Dept. of Zoology, Na- tional Science Museum (Nat. Hist.), To- kyo), and Andreas Taeger (Deutsches En- tomologisches Institut, Eberswalde) loaned specimens. Matti Viitasaari (Dept. of Ap- plied Biology, University of Helsinki) ar- ranged the loan of D. sericeus and helped with some morphological terms. Arkady S. Lelej (Institute of Biology and Pedology, Vladivostok) provided us with up-to-date geographical information concerning the type locality. Wei Meicai (Central South VOLUME 106, NUMBER 1 Forestry University, Zhuzhou, Hunan) in- formed us about the requested character states of D. shanghaiensis Wei and Nie. Martin Karner and Taavi Virro (Institute of Zoology and Hydrobiology, University of Tartu) assisted in preparing the digital mi- crographs for the line drawings of the ovi- positor. Our cordial thanks is extended to them. LITERATURE CITED Goulet, H. 1986. The genera and species of the Ne- arctic Dolerini (Symphyta, Tenthredinidae, Selan- driinae): Classification and phylogeny. Memoirs of the Entomological Society of Canada No. 135, 208 pp. Haris, A. 1996. New east-Palearctic Dolerus species (Hymenoptera, Symphyta, Tenthredinidae). Acta Zoologica Academiae Scientiarum Hungaricae 42(3): 187-194. . 2000. Study on the Palearctic Dolerus Panzer, 1801 species (Hymenoptera: Tenthredinidae). Fo- lia Entomologica Hungarica/Rovartani K6zlemén- yek 61: 95-148. . 2001. Six new Dolerus Panzer, 1801 species from Japan, Turkey and the United States (Hy- menoptera: Tenthredinidae). Folia Entomologica Hungarica/Rovartani K6zlemények 62: 83—93. Leblanc, L. and H. Goulet. 1992. Descriptions of lar- vae of eight Nearctic species of Dolerus (Hyme- noptera: Tenthredinidae) with focus on six Equi- setum-feeding species from the Ottawa region. Canadian Entomologist 124(6): 999-1014. Lindqvist, E. 1969. Blattwespen-Studien (Hymenop- tera, Symphyta). Notulae Entomologicae 49: 38— 48. Malaise, R. 1931. Entomologische Ergebnisse der 165 schwedischen Kamtchatka-Expedition 1920— 1922. Arkiv f6r Zoologi 23A(8): 1-68. Muche, H. 1965. Mongolische Dolerini (Tenthredini- dae, Dolerinae). Reichenbachia 5(20): 181-189. Viitasaari, M. 2002. The suborder Symphyta of the Hymenoptera, pp. 11-174. Jn Viitasaari, M., ed. Sawflies I (Hymenoptera, Symphyta). A Review of the Suborder, the Western Palaearctic Taxa of Xyeloidea and Pamphilioidea. Tremex Press, Hel- sinki, 516 pp. Viitasaari, M., M. Heidemaa, M. Nuorteva, and A. Zi- novjev. 1998. An annotated checklist of the saw- flies (Hymenoptera, Symphyta) of Estonia. Pro- ceedings of the Estonian Academy of Sciences. Biology Ecology 47(2): 126-147. Wei, M. 1997. New species of sawflies (Hymenoptera: Tenthredinidae) in the collection of Entomological Museum of Northwestern Agricultural University. Entomotaxonomia 19(Suppl.): 17—24. Wei, M. and H. Nie. 1997. Seven new species of Do- lerus Panzer (Hymenoptera: Tenthredino-morpha: Tenthredinidae) from China. Entomotaxonomia 19(Suppl.): 60—68. Zhelochovtsev, A. N. 1928. Uber paliarktische Doler- inae. Zoologischer Anzeiger 79(3—4): 105-112. . 1935. Notes sur les Dolerinae (Hym.) Paléarc- tiques. Archives du Muzée Zoologique de l'Université de Moscou II: 79—84. (In French with Russian summary.) . 1988. Symphyta, pp. 7-234. In Medvedjevy, G. S., ed. Opredeliatel Nasekomykyh Evropeiskoi Chasti SSSR. III. Perepondhatokrylye 6. Oprede- liteli po faune SSSR 158. Nauka Leningrad. (En- glish translation: 1994. Keys to the Insects of the European Part of the USSR, Volume III, Hyme- noptera, Part VI, Symphyta. E. J. Brill, New York, 432 pp.) Zhelochovtsev, A. N. and A. G. Zinovjev. 1996. A list of sawflies and horntails (Hymenoptera, Symphy- ta) of the fauna of Russia and adjacent territories. II. Entomological Review 76: 451—470. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 166-175 ACALYPTRATE DIPTERA ASSOCIATED WITH STANDS OF CAREX LACUSTRIS AND C. STRICTA (CYPERACEAE) IN NORTHEASTERN OHIO B. A. FOOTE Department of Biological Sciences, Kent State University, Kent, OH 44242, U.S.A. Abstract.—Eighty-seven species of acalyptrate Diptera were swept from stands of the sedges Carex lacustris and C. stricta in a small freshwater marsh located near the city of Kent in Portage County in northeastern Ohio. Information is presented on stand prefer- ence, seasonal distribution, and larval feeding habits for the 45 species for which more than one specimen was collected. The most species-rich families were Chloropidae (28 spp.), Ephydridae (15), and Sciomyzidae (14). Key Words: There seems to be a general impression among wetland biologists that higher flies are relatively unimportant in marshes, prob- ably because many of the species of marsh- inhabiting cyclorraphous Diptera are not truly aquatic and thus are not thought to be involved in aquatic food chains leading to fish or waterfowl production (Keiper et al. 2002). However, species richness of Diptera can be high (Todd and Foote 1987a, Beau- lieu and Wheeler 2002), and populations of certain species can be huge in freshwater wetlands (Larson and Foote 1997, Keiper and Walton 2000). Because of their abun- dance, species richness, and diversity of feeding habits, acalyptrate flies undoubtedly play important roles in marshland ecology, and probably are particularly important as food resources for small birds and mam- mals (King and Brazner 1999). This is the eighth paper in a series of publications focusing on the acalyptrate Diptera of freshwater marshes in northeast- ern Ohio (Todd and Foote 1987a, b; Rogers et al. 1991; Wearsch and Foote 1994; Kei- per et al. 1998; Larson and Foote 1997; Foote, in press). The present study gives survey data obtained during one warm sea- freshwater marsh, Carex, acalyptrate Diptera, Ohio, larval feeding habits son of collecting acalyptrate Diptera in stands of two sedge species, Carex lacustris Wild. and C. stricta Lam., in a small marsh located in northeastern Ohio. Information is given on stand preference, seasonal occur- rence, and larval feeding habits of 45 of the 87 associated species. MATERIALS AND METHODS The freshwater marsh, ““Horning Road Marsh,” utilized in this study is located 0.8 km east of the main campus of Kent State University (Portage Co.). It encompasses some 0.5 h and supports a mosaic of eight vegetation types occurring in nearly mono- cultural stands (Todd and Foote 1987a). Carex lacustris is a cespitose, rhizomatous, broad-leaved (8—15 mm) species that forms a nearly closed canopy over the marsh muds. In contrast, C. stricta is a narrow- leaved (2-5 mm) clumped sedge, a growth form that allows considerable light to reach the marsh surface. Acalyptrate Diptera were obtained by sweep samples consisting of 15 back and forth movements of a standard aerial insect net through the vegetation. Sampling took VOLUME 106, NUMBER 1 place weekly for 22 weeks between May 6 and September 29, 1989. RESULTS AND DISCUSSION A total of 93 species of acalyptrate Dip- tera was obtained in the two stands of Car- ex (Table 1). Sixty-four species were found in C. lacustris, and 74 in C. stricta. In the list below, 48 species are covered in greater detail with respect to their occurrences in the two stands, relative abundances, and larval feeding habits. ANNOTATED LIST OF SELECTED SPECIES Family Agromyzidae Cerodontha (Cerodontha) dorsalis (Loew).—Adults of this species were more abundant in C. /acustris, being recorded from mid-May to mid-August. It appeared to be bivoltine, as no adults were recorded between late June and late August. The lar- vae are leaf miners of grasses (Spencer and Steyskal 1986). Grasses were intermixed with C. lacustris in this stand. Cerodontha (Butomomyza) subangulata (Malloch).—Larvae are known to mine leaves of Carex (Spencer and Steyskal 1986). Family Anthomyzidae Anthomyza variegata (Loew).—Six indi- viduals were swept from C. lacustris. Lar- vae of this stem-boring species overwinter in culms of the host plant. Mumetopia occipitalis Melander.—Seven specimens were taken in C. /acustris, and 11 in C. stricta. The larvae are reported to mine the stems of wetland monocots (Ferrar 1987): Family Chamaemyiidae Plunomia elegans Curran, P. tibialis Malloch, P. transversa Malloch.—Thirty- five adults of these three species were taken from C. stricta during late May and early June. Nothing is known of the larval feed- ing habits of any of these species, but other species of the family prey on aphids and scale insects (Sluss and Foote 1971, 1973). 167 Family Chloropidae Chlorops certima Adams.—This was an abundant species between late May and mid-June in C. lacustris. Larvae are stem borers of several Carex species (Rogers et al. 1991). They reported that there is a sin- gle annual generation, with overwintering occurring as third-instar larvae in dead culms. Conioscinella nuda (Adams).—This was a common species in both species of Carex, Where its larvae fed as secondary invaders of stems damaged by phytophagous larvae of the dipterous family Scathophagidae. Dasyopa sp.—One adult of this appar- ently new species was taken in each Carex stand. Nothing is known of the life history or larval feeding habits of any species of the genus. Elachiptera erythropleura Sabrosky.— Taken only in C. stricta, the larvae of this species are secondary invaders of monocot stems damaged by other insect larvae (Val- leyct al: 1969): Elachiptera nigriceps (Loew).—This was an abundant species in both species of Carex. Its larvae feed as secondary invad- ers of stems damaged by phytophagous lar- vae (Valley et al. 1969). Epichlorops exilis (Coquillett).—This was an abundant species between late May and early July in C. lacustris (Fig. 2), where its larvae were stem miners (Rogers et al. OS) Eribolus longulus (Loew).—A common species in wetlands, adults were abundant throughout the warm season in both sedge stands. Its larvae are secondary invaders of the stems of wetland monocots that have been damaged by phytophagous larvae (Valley and Foote 1996). Incertella bispina (Malloch).—Adults were taken only in C. /acustris. Its larval feeding habits are unknown. Incertella (Becker).—This was an abundant species in both species of Car- ex. Its larvae fed as secondary invaders of stems of monocots. incerta 168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Species, numbers, and trophic guilds of acalyptrate Diptera collected in stands of Carex lacustris and C. stricta in northeastern Ohio. eS Number Collected in Carex Carex Species lacustris stricta Trophic Guild AGROMYZIDAE Agromyza albipennis Meigen | 0) Leaf miner Amauromyza sp. 0 1 Leaf miner Cerodontha dorsalis (Loew) 5 1 Leaf miner Cerodontha magnicornis (Loew) i 0) Leaf miner Cerodontha subangulata (Malloch) ] 0) Leaf miner Metopomyza interfrontalis (Melander) 2 0) Leaf miner ANTHOMYZIDAE Anthomyza variegata (Loew) 0) 2, Stem borer Mumetopia occipitalis Melander 7 11 Stem borer Undetermined sp. 0) 4 Unknown AULACIGASTRIDAE Cyamops halteratus Sabrosky 0) | Unknown CHAMAEMYIIDAE Chamaemyia juncorum (Fallén) 0) l Scale predator Plunomia elegans Curran 0) 29 Unknown Plunomia tibialis Malloch 0) 4 Unknown Plunomia transversa Malloch 4 | Unknown CHLOROPIDAE Apallates neocoxendix (Sabrosky) 0) | Unknown Aphanotrigonum scabra (Aldrich) 0) 4 Unknown Chlorops certima Adams 2 a Stem borer Chlorops sulphurea Loew 0) Stem borer Conioscinella flavescens (Tucker) 0) | Secondary invader Conioscinella nuda (Adams) 14 >) Secondary invader Dasyopa sp. ] l Unknown Elachiptera costata (Loew) | | Secondary invader Elachiptera erythropleura Sabrosky 0) 3 Secondary invader Elachiptera nigriceps (Loew) 33} 48 Secondary invader Elachiptera penita (Adams) 3 l Unknown Epichlorops exilis (Coquillett) 163 4 Stem borer Eribolus longulus (Loew) 60 84 Secondary invader Eribolus nana (Zetterstedt) | 0) Secondary invader Incertella bispina (Malloch) 9 0) Unknown Incertella incerta (Becker) 148 61 Secondary invader Incertella infesta (Becker) 2) 2 Secondary invader Incertella sp. (new?) 2 0) Unknown Liohippelates pallipes (Loew) 8 14 Unknown Meromyza americana Fitch 4 6) Stem borer Olcella trigramma (Loew) 1 0) Unknown Oscinella frit (Linnaeus) 98 44 Stem borer Pseudopachychaeta approximatinervis (Zett.) 2 2 Seed predator Rhopalopterum carbonaria (Loew) 156 112 Secondary invader Rhopalopterum soror (Macquart) i 0) Scavenger Rhopalopterum umbrosa (Loew) 2 4 Unknown Stenoscinis atriceps (Loew) l 6 Secondary invader Thaumatomyia glabra (Meigen) 3 8 Aphid predator DIASTATIDAE Diastata repleta (Walker) 0) l Scavenger ee VOLUME 106, NUMBER 1 169 Table 1. Continued. NO Number Collected in Carex Carex Species lacustris stricta Trophic Guild DROSOPHILIDAE Drosophila palustris Spencer 0) 2 Scavenger Drosophila quinaria Loew 0) l Scavenger Scaptomyza adusta (Loew) 2 5 Scavenger Scaptomyza pallida (Zetterstedt) 81 455 Scavenger EPHYDRIDAE Coenia curvicauda (Meigen) 4 2 Scavenger, phycovore Discocerina obscurella (Fallén) | 2 Scavenger, phycovore Hyadina albovenosa Coquillett | 3 Phycovore Hydrellia griseola (Fallén) 2, 3 Leaf miner Lytogaster excavata (Sturtevant and Wheeler) D; 7 Phycovore Microlytogaster extera (Cresson) | | Phycovore Nostima picta (Fallén) 1 5 Phycovore Notiphila caudata (Fallén) 18 y) Scavenger, phycovore Notiphila solita Walker | | Scavenger Ochthera anatolikos Clausen O l Insect predator Pelina truncatula Loew 0) | Phycovore Philotelma alaskensis Cresson | l Phycovore Philygria debilis Loew 0) l Phycovore Scatella picea (Walker) | 0) Scavenger, phycovore Scatella stagnalis (Fallén) 0) 8 Scavenger, phycovore LAUXANIIDAE Camptoprosopella sp. Unknown Minettia lupulina (Fabricius) Scavenger MICROPEZIDAE Compsobata pallipes (Say) Scavenger OPOMYZIDAE Opomyza petrei Mesnil Stem borer SCIOMYZIDAE Atrichomelina pubera (Loew) 2 | Snail predator Dictya expansa Steyskal 7 8 Snail predator Dictya steyskali Valley 4 | Snail predator Elgiva solicita (Harris) 28 29 Snail predator Pherbellia nana (Fallén) 12 10) Snail predator Pteromicra pleuralis (Cresson) 7 4 Snail predator Sepedon armipes Loew | 10) Snail predator Sepedon borealis Steyskal 4 10) Snail predator Sepedon fuscipennis Loew 22 5) Snail predator Sepedon tenuicornis Cresson 168 44 Snail predator Tetanocera ferruginea Fallén 7 3 Snail predator Tetanocera fuscinervis (Zerrterstedt) l 0) Snail predator Tetanocera loewi Steyskal 2, | Snail predator Tetanocera plumosa Loew 0) Snail predator SEPSIDAE Enicita annulipes (Meigen) 0 2 Scavenger Enicomira minor (Haliday) 0 3 Scavenger Sepsis flavimana Meigen 10) 12 Scavenger a 170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Number Collected in Carex Carex Species lacustris stricta Trophic Guild SPHAEROCERIDAE Coproica urbana | 0) Scavenger Leptocera fontinalis (Fallén) 1] 178 Scavenger Limosina sp. 0) 2 Scavenger Rachispoda richardsi Sabrosky 0) 2 Scavenger Rachispoda spuleri Sabrosky 0) | Scavenger Trachyopella nuda Rohacek and Marshall | 0) Scavenger TEPHRITIDAE Euaresta bella (Loew) l 0 Seed predator Euaresta festiva (Loew) 2 O Seed predator ULIDUDAE Chaetopsis aenea (Wiedemann) 159 41 Scavenger Chaetopsis massyla (Walker) 161 15 Scavenger Number of species: 64 74 Number of individuals 1,454 | 32 Liohippelates pallipes (Loew).—This was a fairly common species in both spe- cies of Carex. Its larval feeding habits are unknown. Oscinella frit (Linnaeus).—Adults were collected in both species of sedge, although they were far more abundant in an adjacent stand of reed canary grass, Phalaris arun- dinacea L. Larvae are stem borers of a great variety of grasses (Ferrar 1987). Pseudopachychaeta approximatinervis (Zetterstedt)—Only two adults were taken in each species of Carex. The larvae are seed predators of spike-rush, Eleocharis smalliit (Valley et al. 1969), and other spe- cies of sedges. AB 25 — 3 20 = z 15 2) iS wv 10 mee =. 5 — ZO eal \o — AQ \o — wr \o = Ta) fo) N Son eR ieee SP OR” SS he aa 2 eer = = = = — = = eo meee BN Se ee a ne > > DATE Fig. 1. Seasonal occurrence of species of acalyptrate Diptera in Carex lacustris. VOLUME 106, NUMBER 1 171 Q 80 E 70 = 60 5 50 S 40 = 30 me 20 ad Z. 0 T bd T ee SiG re ate Ne aa ate aes EAS ES Ee DATE Fig. 2. Seasonal occurrence of Epichlorops exilis in Carex lacustris. Rhopalopterum carbonaria (Loew).— Ephydridae This was an abundant taxon in both species of Carex. Its larvae are secondary invaders of stems previously damaged by phytoph- agous insect larvae (Valley et al. 1969). Thaumatomyia glabra (Meigen).—Three adults of this species were taken in C. /a- custris, whereas eight were swept from C. stricta. Larvae are predators of root-inhab- iting aphids (Parker 1918). Family Drosophilidae Drosophila palustris Spencer.—Taken only in C. lacustris, adults of this marsh- inhabiting species were reared from larvae found in moist, decaying leaves and stems of various species of sedges. Drosophila quinaria Loew.—This 1s an- other species of Drosophila commonly found in freshwater marshes, although I collected only one specimen. Its larvae feed on decaying plant material. Scaptomyza pallida (Zetterstedt).—This was an abundant species throughout the collecting season, with greatest numbers being obtained in C. stricta. Its larvae are general scavengers of decaying plant ma- terial (Ferrar 1987). Coenia curvicauda (Meigen).—Usually this is a fairly common species in marshes, but I obtained only six specimens, mostly from C. lacustris. The larvae are general- ized feeders on organic particulate matter, including algae and decaying plant material (Foote 1990). Discocerina obscurella (Fallén).—Only three specimens of this normally common and widely distributed species were found in the stands of Carex. Its larvae are gen- eralized feeders of particulate organic mat- ter (Foote and Eastin 1974). Hyadina albovenosa Coquillett.—Un- common in both stands, larvae of this spe- cies feed on Cyanobacteria (Foote 1993). Hydrellia griseola (Fallén).—Uncom- mon in the sedge stands, larvae of this spe- cies feed as leaf miners on a great variety of grasses and other wetland monocots (Deonier 1971, 1978). Lytogaster excavata (Sturtevant and Wheeler).—Fairly common in the stand of C. stricta, larvae of this species consume Cyanobacteria, particularly species of soil- inhabiting Cylindrospermum (Foote 198 la). Nostima picta (Fallén).—Larvae of this 172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON S = 30 25 = 20 5 Sale x = 10 = 5S > (0 \o —) AQ ‘© —) wt oe) = la) io) = S = or. a in D “a car = coe oS BR BP Rr ee ee eee > > DATE Fig. 3. Seasonal occurrence of Sepedon tenuicornis in Carex lacustris. species are specialized consumers of the cy- anobacterial genus Oscillatoria (Foote 1983). Notiphila caudata Fallén.—This was an abundant species in both stands. Its larvae feed on particulate organic matter, including algal cells, on the surface of marsh soils (Eastin and Foote 1971). Pelina truncatula Loew.—An_ uncom- mon species in sedge marshes, this species is more regularly taken in stands of cattail (Typha spp.). Its larvae feed on such cy- anobacterial genera as Anabaena and Cylin- dropsermum (Foote 1981b). Philygria debilis Loew.—Only one spec- imen of this small, inconspicuous species was found in C. stricta. Adults are more commonly encountered in marshes and moist meadows dominated by grasses. Lar- vae feed on the cyanobacterial genus Os- cillatoria (Foote 1983). Family Micropezidae Compsobata pallipes (Say).—Eleven specimens were swept from C. stricta. Lar- vae of a closely related species have been reared from decaying plant material (Tes- key, 1972): Family Sciomyzidae Dictya expansa Steyskal, D. steyskali Valley.—Adults of both species were taken in both sedges throughout the summer months. Larvae prey on aquatic pulmonate snails (Valley and Berg 1977). Elgiva solicita (Harris).—A common species in both sedges throughout the warm season, this species has the interesting habit of overwintering as adults in marshes. Its larvae prey unselectively on aquatic pul- monate snails (Knutson and Berg 1964). Pherbellia nana nana (Fallén).—Adults were found in C. lacustris throughout the spring and summer months. Larvae of this species attack a great variety of small pul- monate aquatic snails that have been strand- ed by dropping water levels (Bratt et al. 1969). Sepedon borealis Steyskal, S. fuscipennis Loew, S. tenuicornis Cresson.—All three species of Sepedon were encountered re- peatedly in both sedge stands. Adults over- winter in marshes where their larvae prey on aquatic pulmonate snails (Neff and Berg 1966). Tetanocera ferruginea Fallén.—This was not a common species in the sedges stands, as the adults seemingly preferred habitats VOLUME 106, NUMBER 1 173 2 250 _ = LY 200 ae 5 150 s ~ 100 = S 50 = iil Z 0 \o S AN \o S Su io) _ va) ce) N RT gk Try oy Cia. Mangutt eke how) Sn, mebenis erage tes eee Sue ie So Sowa iene > > DATE Fig. 4. Seasonal occurrence of individuals of acalyptrate Diptera in Carex lacustris. possessing somewhat deeper water (e.g., cattail marshes). It was multivoltine, pro- ducing three or four generations a year. Its larvae feed on a variety of pulmonate aquatic snails (Foote 1999). Tetanocera loewi Steyskal.—An uncom- mon species in both Carex stands, adults appeared in mid-June, remained at low numbers throughout July and August, and disappeared from the marsh in late Septem- ber. It had a single generation a year, with overwintering occurring as unhatched first- instar larvae within the egg envelopes. Hatching took place in March, and most of the larval feeding on a variety of pulmonate aquatic snails (Physa, Lymnaea, Gyraulus, Helisoma) was completed during April and early May (Foote 1999). Family Sepsidae Sepsis flavimana Meigen.—This was the only species of Sepsidae that was taken re- peatedly in the sedge stands, and was par- ticularly abundant in C. stricta. Its larvae are unselective scavengers of decaying or- ganic matter, particularly dung (Ferrar 1987). Family Sphaeroceridae Leptocera fontinalis (Fallén)—This was the only abundant species of the six species of Sphaeroceridae encountered in the sedge stands, being particularly common in C. stricta. Larvae of all species are thought to be scavengers of decaying organic matter (Ferrar 1987). Family Ulidiidae Chaetopsis aenea (Wiedemann).—This was a common species in both stands of Carex, but showed a distinct preference for C. lacustris. It first appeared and reached peak abundance in early June, and largely disappeared by late July. Larvae feed as secondary consumers in stems of monocots that have been damaged by more phytoph- agous species (Allen and Foote 1992). Chaetopsis massyla (Walker).—This spe- cies was also recorded in both stands, with greatest abundance in C. lacustris in late June. Its larvae feed as secondary invaders of stems of damaged wetland plants (Allen and Foote 1992). OCCURRENCE IN THE TWO STANDS Carex stricta supported a somewhat greater number of acalyptrate taxa, with some 74 species being recorded. In con- trast, 64 species were taken in C. lacustris (Table 1). One possible explanation for the slightly greater species richness in C. stricta 174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON was the fact that this tussock sedge species has a clumped distribution with open spaces between clumps. This allowed light to reach the marsh floor, resulting in greater algal growth that served as larval food for some species of Ephydridae. In contrast, Carex lacustris is a rhizomatous species that formed a more closed canopy over the marsh substrate. There was considerable sharing of species among the two species of Carex, and Sgrenson’s Similarity Index (Sgrenson 1948) was 0.66 meaning that 66% of the acalyptrate species were found in both sedges. SEASONAL OCCURRENCE The number of species in both of the Carex stands peaked in June and remained fairly constant throughout the remainder of the summer (Fig. 1). Many species were found throughout the warm season, but a few species obviously were univoltine or bivoltine. For example, adults of Epichlo- rops exilis peaked in early June (Fig. 2). Adults of Sepedon tenuicornis overwin- tered in the marsh and were quite abundant again in late summer (Fig. 3). Total abun- dance of adult flies in the two stands peaked in early June (Fig. 4), and populations had largely collapsed by mid-October. GUILD STRUCTURE There was obvious partitioning of food resources in the two stands of Carex, with eight trophic guilds being recorded (Table 1). The largest guild involved some 23 spe- cies that fed as generalized scavengers of decaying organic matter. The phytophagous community consisted of a leaf-mining guild containing six species, a stem-boring guild of eight species, and a seed predator guild of three species. The guild of secondary in- vaders consisted of 13 species whose larvae fed as scavengers on stem tissues previous- ly macerated by more phytophagous spe- cies. An interesting guild consisted of seven species whose larvae consumed Cyanobac- teria. The snail predator guild contained 14 species, and three species preyed upon oth- er insects. The larval feeding habits of 15 species remain unknown. ACKNOWLEDGMENTS Appreciation is expressed to Barbara K. Andreas, Cuyahoga Community College in Cleveland, and Samuel J. Mazzer, Kent State University, for their aid in determin- ing the plant species occurring in the study marshes. LITERATURE CITED Allen, E. J. and B. A. Foote. 1992. Biology and im- mature stages of Chaetopsis massyla (Diptera: Otitidae), a secondary invader of herbaceous stems of wetland monocots. Proceedings of the Entomological Society of Washington 94: 320— 328. Beaulieu, FE and T. A. Wheeler. 2002. Insects (Diptera, Coleoptera, Lepidoptera) reared from wetland monocots (Cyperaceae, Poaceae, Typhaceae) in southern Quebec. Proceedings of the Entomolog- ical Society of Washington 104: 300-308. Bratt, A. D., L. V. Knutson, B. A. Foote, and C. O. Berg. 1969. Biology of Pherbellia (Diptera: Scio- myzidae). Memoir of the Cornell University Ag- ricultural Experiment Station 404: 1-247. Deonier, D. L. 1971. A systematic and ecological study of Nearctic Hydrellia (Diptera: Ephydridae). Smithsonian Contributions to Zoology 68: 1-147. . 1978. A Manual of the Common North Amer- ican Species of the Aquatic Leafmining Genus Hydrellia (Diptera: Ephydridae). Memoirs on En- tomology, International 12. Associated Publishers, Gainesville, FL, 354 pp. Eastin, W. C. and B. A. Foote. 1971. Biology and im- mature stages of Dichaeta caudata (Diptera: Ephydridae). Annals of the Entomological Society of America 64: 271—279. Ferrar, P. 1987. A Guide to the Breeding Habits and Immature Stages of Diptera Cyclorrapha. Ento- monograph 8, part 1, Text. E. J. Brill/Scandina- vian Press, Leiden, 478 pp. Foote, B. A. 198la. Biology and immature stages of Lytogaster excavata, a grazer of blue-green algae (Diptera: Ephydridae). Proceedings of the Ento- mological Society of Washington 83: 294-315. . 1981b. Biology and immature stages of Pelina truncatula, a consumer of blue-green algae (Dip- tera: Ephydridae). Proceedings of the Entomolog- ical Society of Washington 83: 607—619. . 1983. Biology and immature stages of Nos- tima approximata (Diptera: Ephydridae), a grazer of the blue-green algal genus Oscillatoria. Pro- ceedings of the Entomological Society of Wash- ington 85: 472—484. VOLUME 106, NUMBER 1 . 1990. Biology and immature stages of Coenia curvicauda (Diptera: Ephydridae). Journal of the New York Entomological Society 98: 93-102. . 1993. Biology of Hyadina albovenosa (Dip- tera: Ephydridae), a consumer of Cyanobacteria. Proceedings of the Entomological Society of Washington 95: 377-382. . 1999. Biology and immature stages of snail- killing flies belonging to the genus Tetanocera (Insecta: Diptera: Sciomyzidae). III. Life histories of the predators of aquatic snails. Annals of Car- negie Museum 68: 151-174. . In Press. Acalyptrate Diptera of a fen at the Herrick Preserve in Portage County, Ohio. Great Lakes Entomologist. Foote, B. A. and W. C. Eastin. 1974. Biology and im- mature stages of Discocerina obscurella (Diptera: Ephydridae). Proceedings of the Entomological Society of Washington 76: 401—408. Keiper, J. B. and W. E. Walton. 2000. Biology and immature stages of Brachydeutera sturtevanti (Diptera: Ephydridae), a hyponeustic generalist. Annals of the Entomological Society of America 93: 468-475. Keiper, J. B., P. L. Brutsche, and B. A. Foote. 1998. Higher Diptera associated with water willow, Jus- ticia americana (Acanthaceae). Proceedings of the Entomological Society of Washington 100: 576— 587. Keiper, J. B.. W. E. Walton, and B. A. Foote. 2002. Biology and ecology of higher Diptera from fresh- water wetlands. Annual Review of Entomology 47: 207-232. King, R. S. and J. C. Brazner. 1999. Coastal wetland insect communities along a trophic gradient in Green Bay, Lake Michigan. Wetlands 19: 426— 437. Knutson, L. V. and C. O. Berg. 1964. Biology and immature stages of snail-killing flies: The genus Elgiva (Diptera: Sciomyzidae). Annals of the En- tomological Society of America 57: 173-192. Larson, L. and B. A. Foote. 1997. Biology of four species of Notiphila Fallén (Diptera: Ephydridae) associated with the yellow water lily, Nuphar lu- teum (Nymphaeaceae). Proceedings of the Ento- mological Society of Washington 99: 541-559. Neff, S. E. and C. O. Berg. 1966. Biology and im- mature stages of malacophagus Diptera of the ge- nus Sepedon (Sciomyzidae). Bulletin of the Vir- ginia Agricultural Experiment Station 566: I—113. Parker, J. R. 1918. The life history and habits of Chlo- ropisca glabra Meig., a predacious Oscinid (Chlo- 175 ropid). Journal of Economic Entomology 11: 368— 380. Rogers, T. P, B. A. Foote, and J. L. Todd. 1991. Bi- ology and immature stages of Chlorops certimus and Epichlorops exilis (Diptera: Chloropidae), stem-borers of wetland sedges. Journal of the New York Entomological Society 99: 664-683. Sluss, T. P. and B. A. Foote. 1971. Biology and im- mature stages of Leucopis verticalis (Diptera: Chamaemyiidae). Canadian Entomologist 103: 1427-1434. . 1973. Biology and immature stages of Leu- copis pinicola and Chamaemyia polystigma (Dip- tera: Chamaemuiidae). Canadian Entomologist 105: 1443-1452. Sgrenson, T. 1948. A method of establishing groups of equal amplitude in plant sociology based on sim- ilarity of species content. Kongelige Danske Vi- denskabernes Selskab 5: 1—34. Spencer, K. A. and G. C. Steyskal. 1986. Manual of the Agromyzidae (Diptera) of the United States. United States Department of Agriculture, Agri- culture Handbook 638, 478 pp. Teskey, H. 1972. The mature larva and pupa of Comp- sobata univitta (Diptera: Micropezidae). Canadian Entomologist 104: 295-298. Todd, J. L. and B. A. Foote. 1987a. Spatial and tem- poral distribution of shore flies in a freshwater marsh (Diptera: Ephydridae). Proceedings of the Entomological Society of Washington 89: 448— 457. . 1987b. Resource partitioning in Chloropidae (Diptera) of a freshwater marsh. Proceedings of the Entomological Society of Washington 89: 803-8 10. Valley, K. R. and C. O. Berg. 1977. Biology and im- mature stages of snail-killing Diptera of the genus Dictya (Sciomyzidae). Search Agriculture 7: 1— 45. Valley, K. R. and B. A. Foote. 1996. Biology and im- mature stages of Eribolus longulus, with notes on E. nanus (Diptera: Chloropidae), secondary invad- ers of herbaceous wetland plants. Memoir of the Entomological Society of Washington 18: 273-— ATS) Valley, K. R., T. Wearsch, and B. A. Foote. 1969. Lar- val feeding habits of certain Chloropidae (Dip- tera). Proceedings of the Entomological Society of Washington 71: 29—34. Wearsch, T. K. and B. A. Foote. 1994. Biology and immature stages of Chloropidae (Insecta: Diptera) associated with spike-rushes (Cyperaceae: Eleo- charis). 1. Stem borers. Annals of Carnegie Mu- seum 63: 193-213. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 176-180 NEW NORTH AMERICAN RECORDS OF THE EUROPEAN BROOM PSYLLID ARYTAINA GENISTAE (LATREILLE) (STERNORRHYNCHA: PSYLLIDAE) A. G. WHEELER, JR. AND E. RICHARD HOEBEKE (AGW) Department of Entomology, Clemson University, Clemson, SC 29634-0365, U.S.A. (e-mail: awhlr@clemson.edu); (ERH) Department of Entomology, Cornell Uni- versity, Ithaca, NY 14853, U.S.A. (e-mail: erh2 @cornell.edu) Abstract.—The European psyllid Arytaina genistae (Latreille) has been unintentionally introduced into North America, probably with imported nursery stock of its principal host, Scotch broom (Cytisus scoparius; Fabaceae). Known previously in North America only from an early twentieth-century collection from Massachusetts and a record of its inter- ception in California in nursery stock originating in Washington State, A. genistae is reported from Nova Scotia, Canada, and from North Carolina, Oregon, and Washington in the United States. Morphological characters are provided that allow it to be separated from Arytainilla spartiophila (Forster), another European broom psyllid that also is es- tablished in eastern and western North America. Key Words: Scotch broom Scotch broom (Cytisus scoparius [L.] Link; Fabaceae) is a common northern and western European shrub that has been widely planted and become naturalized beyond its native range. This deciduous, perennial, yel- low-flowered legume was planted in Colonial American gardens (Leighton 1976) and was used as an ornamental in California and the Pacific Northwest by the mid-nineteenth cen- tury (Clark 1976, Andres and Coombs 1995). It also might have been accidentally intro- duced into North America with ship ballast (Lindroth 1957). In both eastern and western North America, Scotch broom has been used for erosion control along highways (Hitch- cock and Cronquist 1973, Pfeiffer 1986, Dirr 1998). It has been used in the Pacific North- west for stabilizing coastal sand dunes (Schwendiman 1977). This r-adapted shrub (Williams 1981) is considered an invasive plant in western North America and in Aus- Insecta, Psyllidae, Arytaina genistae, insect distribution, adventive species, tralia, Chile, Iran, New Zealand, and South Africa (Syrett et al. 1999). In Great Britain and continental Europe, a diverse arthropod fauna (>240 species) is associated with Scotch broom (Syrett et al. 1999). Long-term studies of its fauna were conducted at Silwood Park, Berkshire, Eng- land, by J.P. Dempster, O.W. Richards, N. Waloff, and their colleagues and students in the 1950s and 1960s (Waloff 1968). AI- though an Old World lyonetiid moth was introduced into the western United States in the early 1960s to help reduce densities of Scotch broom (Frick 1964, Andres and Coombs 1995), basic studies on the ecology of broom arthropods in England mostly pre- dated the main use of biological control against this plant in areas where it is not native (e.g., Rees and Paynter 1997, Syrett et al. 1999, Fowler et al. 2001). Among the insects of Scotch broom that VOLUME 106, NUMBER 1 have been accidentally introduced into North America are two species of Psyllidae, the bi- or trivoltine Arytaina genistae (La- treille) and the univoltine Arytainilla spar- tiophila (Forster). In North America, Ary- taina genistae is known only from Massa- chusetts (Crawford 1911, Hodkinson 1988) and Washington (California Department of Food and Agriculture 1993), whereas Ary- tainilla spartiophila is widely distributed in British Columbia, California, Oregon, and Washington (Waloff 1966, Syrett et al. 1999) and is known in the East from Vir- ginia (Pfeiffer 1986). Herein, we give the first records of Arytaina genistae for Can- ada and the southeastern United States. We also provide morphological characters that allow these adventive psyllids to be recog- nized in the Nearctic fauna. Arytaina genistae (Latreille) The first North American record of A. genistae was given by Crawford (1911) in describing the new species Psyllopa magna based on T.D.A. Cockerell’s collection of a large series of specimens from Spartinum [sic] sp. at Woods Hole, Massachusetts. The host plant likely was Scotch broom. Spar- tium scoparium not only is a former name for C. scoparius (Peterson and Prasad 1998), but this plant also was recorded from Woods Hole in the early twentieth century as a host for other Old World broom insects (Olsen 1918, Wheeler and Henry 1992). No collection date was given by Crawford (1911), but Cockerell, a professor at the University of Colorado, Boulder, likely col- lected the psyllid in 1911; that year he spent the summer at the Woods Hole Oceano- graphic Institution (Weber 1965). Crawford (1914) realized the psyllid from Massachu- setts that he had described as new was a previously described European species. He synonymized P. magna under A. genistae and noted that this adventive species prob- ably had been accidentally introduced into North America with nursery stock. Records of A. genistae from western North America have not appeared in the primary 177 literature. Waloff (1966) did not record it from Scotch broom in British Columbia or California, Hodkinson (1988) listed it only from Massachusetts in his checklist of Ne- arctic Psyllidae, Syrett et al. (1999) did not include it in a discussion of insects known from broom in North America, and Maw et al. (2000) did not include it in their checklist of Canadian Hemiptera. This psyllid, how- ever, was intercepted in Santa Clara Co., Cal- ifornia, on broom shipped from a nursery in Shelton, Washington (California Department of Food and Agriculture 1993; R.J. Gill, per- sonal communication). New records.—CANADA: Nova Scotia: Shelburne Co., Shelburne, nr. marine ter- minal, 19 July 1994, 21 3, 16 2; 7 Aug. 2001, 20 3, 16 2, ex Cytisus scoparius, E.R. Hoebeke & A.G. Wheeler. UNITED STATES: North Carolina: Buncombe Co., jet. U.S. Bus. Rt. 70 & I-40 (exit 55), east edge of Asheville, 4 July 2002, 9 3, 6 Q; 20 July 2002, 3 3d, 4 2, ex Cytisus scopar- ius, A.G. Wheeler; Haywood Co., nr. jet. US. Bus: Rt. 23 & SR-1801 (liner Cove Rd.), 4.5 km NNE of Waynesville, 29 June 2002;:8.:5% 82924 Julye200205e6 2679-13 Fully: “200257 StvGw Sr ile tex Ge seoparias A.G.W. Oregon: Marion Co., Aurora, 22 June 1999, 1 3, 4 9, ex C. scoparius, E.R.H. Washington: Clallam Co., nr. Crane, 91 Sept) 1999,..10 dy5 “Ps ex) scoparius A.G.W. & C.A. Stoops; Jefferson Co., Port Townsend, 9 Sept. 1999, 1 6, 4 2; ex C. scoparius, A.G.W. & C.A.S.; Kitsap Co., Gateway Park, Silverdale, 8 Sept. 1999, 3 6; 4 (Pex Ci scoparius, A:G. Wake CASS Mason Co., Rt. 3, Shelton, 10 July 1999, 2 6, ex C. scoparius, A.G.W.& C:A‘S.; Pierce Co., Purdy, 10 Sept. 1999, 2 9, ex C. scoparius,. Ac-G:W. & C_A°s. Voucher specimens have been deposited in the Cornell University Insect Collection (CUIC), Ithaca, NY. RECOGNITION FEATURES Arytaina genistae differs in several re- spects from Arytainilla spartiophila, the only other Scotch broom psyllid established 178 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. cells identified. in North America. In Arytaina genistae, the forewing is somewhat elliptical, broadest at the middle, with a dark brown to black pat- tern occupying cells r, and cu,, but also with small patches at the apices of cells m,, m,, and cu, (Fig. 1) (illustrated by Hodkin- son and White 1979: 38, fig. 109; Ossian- nilsson 1992: 95, fig. 406). In contrast, the forewing of Arytainilla spartiophila is ob- long-oval, broadest at the apical third, and entirely pale yellow throughout (Pfeiffer 1986: 215, fig. 1). The male and female ter- minalia of each species also are distinct, as illustrated by Hodkinson and White (1979) and Ossiannilsson (1992). DISCUSSION More specialist insect species are found on Scotch broom in the Pacific Northwest of North America than in any other region where this plant is adventive (Syrett et al. 1999). Most broom herbivores accidentally introduced into the Pacific Northwest and other regions either overwinter as eggs em- Arytaina genistae adult female, lateral view. y Scale line = 1.0 mm. Inset: Forewing with maculated bedded in year-old twigs or as larvae under the bark of host shoots (Waloff 1966). Ar- ytainilla spartiophylia, which overwinters in the egg stage and inserts its eggs into host shoots (Waloff 1968, Watmough 1968), fits the life-history pattern of most other introduced broom insects. In contrast, Arytaina genistae deposits its eggs super- ficially on leaves and other plant parts, and the adults overwinter on the host plant (Waloff 1968, Watmough 1968). It, there- fore, might not have been predicted to be introduced with nursery stock and to be- come established in North America. Yet re- cords of A. genistae from both the east and west coasts suggest multiple introductions from Europe. Alternatively, a single intro- duction and subsequent movement in North America with shipments of broom nursery stock could have resulted in the currently known distribution of this psyllid. ACKNOWLEDGMENTS We thank Raymond Gill (Plant Health and Pest Prevention Services, California VOLUME 106, NUMBER 1 Department of Food and Agriculture, Sac- ramento) for providing information on the interception record of A. genistae in Cali- fornia, Timothy Forrest (Department of Bi- ology, University of North Carolina at Asheville) for providing information on the occurrence of Scotch broom in western North Carolina, Craig Stoops (Department of Entomology, Clemson University) for helping to collect psyllids in Washington, Peter Adler (Department of Entomology, Clemson University) for providing useful comments on an earlier draft of the manu- script, and Kent Loeffler (Department of Plant Pathology, Cornell University) for photographing A. genistae. LITERATURE CITED Andres, L. A. and E. M. Coombs. 1995. Scotch broom, Cytisus scoparius (L.) Link; Fabaceae, pp. 303— 305. In Nechols, J. R., exec. ed. Biological Con- trol in the Western United States: Accomplish- ments and Benefits of Regional Research Project W-84, 1964-1989. University of California Divi- sion of Agriculture and Natural Resources Publi- cation 3361. California Department of Food and Agriculture. 1993. Exclusion: Genista psyllid. California Plant Pest and Disease Report 12(3—4): 88. Clark, L. J. 1976. Wild Flowers of the Pacific North- west from Alaska to Northern California. Gray’s Publishing, Sidney, B.C., Canada, 604 pp. Crawford, D. L. 1911. American Psyllidae V. Pomona Journal of Entomology 3: 628—632. . 1914. A monograph of the jumping plant-lice or Psyllidae of the New World. United States Na- tional Museum Bulletin 85. Government Printing Office, Washington, D.C., 186 pp. Dirr, M. A. 1998. Manual of Woody Landscape Plants: Their Identification, Ornamental Characteristics, Culture, Propagation and Uses, 5th ed. Stipes Pub- lishing, Champaign, Illinois, 1187 pp. Fowler, S. V., J. Memmott, Q. E. Paynter, A. W. Shep- pard, and P. Syrett. 2001. The scope and value of extensive ecological studies in the broom biolog- ical control programme, pp. 229-248. Jn Wajn- berg, E., J. K. Scott, and P. C. Quimby, eds. Eval- uating Indirect Ecological Effects of Biological Control. CABI Publishing, Wallingford, Oxford- shire, UK. Frick, K. E. 1964. Leucoptera spartifoliella, an intro- duced enemy of Scotch broom in the western United States. Journal of Economic Entomology 57: 589-591. Hitchcock, C. L. and A. Cronquist. 1973. Flora of the 179 Pacific Northwest: An Illustrated Manual. Univer- sity of Washington Press, Seattle, 730 pp. Hodkinson, I. D. 1988. The Nearctic Psylloidea (In- secta: Homoptera): An annotated check list. Jour- nal of Natural History 22: 1179-1243. Hodkinson, I. D. and I. M. White. 1979. Homoptera Psylloidea. Handbooks for the Identification of British Insects. Vol. I, Pt. Sa. Royal Entomolog- ical Society of London, London, 98 pp. Leighton, A. 1976. American Gardens in the Eigh- teenth Century: “For Use or for Delight.”” Hough- ton Mifflin, Boston, 514 pp. Lindroth, C. H. 1957. The Faunal Connections Be- tween Europe and North America. Wiley, New York, 344 pp. Maw, H. E. L., R. G. Foottit, K. G. A. Hamilton, and G. G. E. Scudder. 2000. Checklist of the Hemip- tera of Canada and Alaska. NRC Research Press, Ottawa, 220 pp. Olsen [C. E.]. 1918. [Exhibition of specimens.] Journal of the New York Entomological Society 26: 234. Ossiannilsson, E 1992. The Psylloidea (Homoptera) of Fennoscandia and Denmark. Fauna Entomologica Scandinavica. Vol. 26. E.J. Brill, Leiden, the Netherlands, 346 pp. Peterson, D. J. and R. Prasad. 1998. The biology of Canadian weeds. 109. Cytisus scoparius (L.) Link. Canadian Journal of Plant Science 78: 497—504. Pfeiffer, D. G. 1986. Arytainilla spartiophila (Foerster) (Homoptera: Psyllidae) on Cystisus scoparius (L.) Link in Virginia: First record in eastern North Amer- ica. Journal of Entomological Science 21: 214-218. Rees, M. and Q. Paynter. 1997. Biological control of Scotch broom: Modelling the determinants of abundance and the potential impact of introduced insect herbivores. Journal of Applied Ecology 34: 1203-1221. Schwendiman, J. L. 1977. Coastal sand dune stabili- zation in the Pacific Northwest. International Jour- nal of Biometeorology 21: 281—289. Syrett, P., S. V. Fowler, E. M. Coombs, J. R. Hosking, G. P. Markin, Q. E. Paynter, and A. W. Sheppard. 1999. The potential for biological control of Scotch broom (Cytisus scoparius) (Fabaceae) and related weedy species. Biocontrol News and In- formation 20: 17N—34N. Waloff, N. 1966. Scotch broom (Sarothamnus scopar- ius (L.) Wimmer) and its insect fauna introduced into the Pacific Northwest of America. Journal of Applied Ecology 3: 293-311. . 1968. Studies on the insect fauna on Scotch broom, Sarothamnus scoparius (L.) Wimmer. Ad- vances in Ecological Research 5: 87—208. Watmough, R. H. 1968. Population studies on two spe- cies of Psyllidae (Homoptera, Sternorhyncha) [sic] on broom (Sarothamnus scoparius (L.) Wimmer). Journal of Animal Ecology 37: 283-314. Weber, W. A. 1965. Theodore Dru Alison Cockerell, 180 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1866-1948. University of Colorado Studies. Se- America. Thomas Say Foundation Monograph ries in Bibliography No. 1. University of Colorado XV. Entomological Society of America, Lanham, Press, Boulder, 124 pp. Maryland, 282 pp. Wheeler, A. G., Jr. and T. J. Henry. 1992. A Synthesis Williams, P. A. 1981. Aspects of the ecology of broom of the Holarctic Miridae (Heteroptera): Distribu- (Cytisus scoparius) in Canterbury, New Zealand. tion, Biology, and Origin, with Emphasis on North New Zealand Journal of Botany 19: 31—43. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 181-191 POST-ADULT EMERGENCE DEVELOPMENT OF GENITALIC STRUCTURES IN SCHISTOCERCA STAL AND LOCUSTA L. (ORTHOPTERA: ACRIDIDAE) HoJUN SONG The Ohio State University, Department of Entomology, Museum of Biological Diver- sity, Columbus, OH 43212, U.S.A. (e-mail: song.131@osu.edu) Abstract.—The male genitalia and female ovipositor in Schistocerca americana (Drury) (Orthoptera: Acrididae: Cyrtacanthacridinae) continue to develop after adult emergence. The internal skeleton comprising the phallic complex is affected by cuticle deposition, resulting in qualitative shape changes during sexual maturation. Lateral apodemes of the ovipositor also grow in length and width during sexual maturation. Similar developmental patterns are found in the male genitalia of S. gregaria (Forskal) and Locusta migratoria (Linnaeus), suggesting the possibility that post-adult emer- gence genitalic development may be widespread within Acridoidea. Newly emerged individuals may be functionally incapable of copulation because the necessary struc- tures have not been fully matured. Taxonomic use of the genitalic structures is dis- cussed in light of the present finding. Key Words: The final molt in insect development re- sults in the adult instar responsible for the reproduction. While there are some groups that are capable of copulating immediately after emergence (Ridley 1989), many in- sects have a period of sexual maturation with the adult instar. The post-emergence maturation period of Schistocerca Stal (Or- thoptera: Acrididae) is about 30 days, which is very long compared to that of oth- er insects (Norris 1954, Ridley 1988, Weis- Fogh 1952). Historically, the gonads have been considered the only developing struc- tures responsible for delayed sexual matu- ration (Norris 1952, 1954, 1957). For ex- ample, Norris (1954) documented the growth of the egg-rudiments in females and underdeveloped receptaculum seminis in males of sexually immature locusts. How- ever, physiological studies revealed that de- velopmental changes at cuticular, muscular, and ultrastructural levels occur throughout post-adult emergence, cuticle development, genitalia, grasshopper, taxonomy the teneral period (Neville 1963a, Weis- Fogh 1952, Viscuso et al. 1985). Significant maturation in the adult stage is known from Orthoptera, Dermaptera, Odonata, Hemip- tera, Homoptera, Hymenoptera, and Diptera (Table 1). Structures that undergo this pro- cess include muscles, corpora allata, epithe- lium, and both internal and external skeletal elements. The period of development can be as long as 35 days (Table 1). The developmental changes documented until now are, however, quantitative chang- es that deal with the increase in thickness or volume. I demonstrate below that the genitalia undergo a qualitative change in shape during the entire period of the sexual maturation in Schistocerca americana (Drury). Also, I explore the taxonomic im- plications of this finding because the geni- talic characters have been used extensively in descriptive taxonomy (Dirsh 1973, Eber- hard 1985). ; Post-adult emergence development of different structures and their processes and timing along with orders known to have the phenomena. Table 1. Author Order Found g Process Post-emergence Timin Structure Neville (1963b) Neville (1983) Until sexual maturity Orthoptera, Dermaptera, Odon- Chitin layer deposition from Exoskeleton ata, Hemiptera, Homoptera, Hymenoptera Orthoptera Diptera epidermis Neville (1963a) Schlein (1972) One week 12 days Resilin deposition Resilin deposition Thoracic apodemes Genital apodemes Thoracic muscles Bursell (1961) Diptera, Orthoptera Until sexual maturity Increase in volume of mito- Finlayson (1975) chondria and contractile protein Development of nuclei and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Weis-Fogh (1952) Odhiambo (1966) Orthoptera Until sexual maturity Corpora allata allatum-cell cytoplasm Release of Golgi vesicles and Viscuso et al. (1985) Orthoptera Until sexual maturity Ejaculatory duct lysis of cells Resilin deposition Resilin deposition epithelium Male phallus This study This study Orthoptera 1-35 days 20-33 days N Orthoptera Lateral apodemes of female ovipositor MATERIALS AND METHODS From laboratory colonies, the last nymphal instars of both male and female Schistocerca americana were collected and transferred to a clean cage. When the nymphs molted to adults, they were placed in separate tubular cages, approximately 7 cm in diameter and 30 cm in height, with a steel mesh for locusts to rest on, and the date of molting was labeled on the cages. Each cage contained 3 males and | female, but if an individual died during the course of experiment, it was immediately removed. Locusts were daily fed fresh Romaine let- tuce and wheat bran, and maintained at 30°C, 12:12 Light:Dark cycle. One cage of insects was killed each day to sample de- velopment across 35 days of post-adult maturation. Insects were killed each day by freezing at —20°C for 30 minutes to collect developmental data. After the genitalia were dissected, specimens were stored in 95% ethanol. Ninety-five male genitalia and thirty-five ovipositors were examined in this study. Fifteen Schistocerca gregaria (Forskal) and fifteen Locusta migratoria (Linnaeus) (5 freshly molted, 5 2-week-old, and 5 sexually mature for each species) were also dissected. Sexual maturity for these species was determined by both time after hatching and yellow coloration (Ste- phen Roberts, personal communication). All three species were reared under crowd- ed condition to avoid any potential density- dependent effects known to exist in locusts. Male genitalia were extruded by inserting a probe under the supra-anal plate, using the technique described by Hubbell (1932). When the phallic complex was exposed, the surrounding membranes were removed. Ovipositors were dissected by making a slit at the distal part of abdomen. Brief descrip- tions of color and morphology were record- ed daily. Genitalia were placed in 10% KOH solution for several hours to dissolve muscles. Cleared genitalia were placed in a vial filled with glycerin, and each genitalic specimen was given an identification num- VOLUME 106, NUMBER | 183 Figs. 1-8. Dimensions of the Schistocerca americana phallic complex and ovipositor measured in the study. 1, Cingulum (RmW: width of rami; ApdR: width of apodeme ridge; CW: width of cingulum). 2, Dorsal view of epiphallus (EpW: horizontal width; EAp: dorso-ventral length of anterior projection). 3, Ventral view of epiphallus (B: dorso-ventral width of bridge; EpB: width of bulbous base). 4, Lateral view of endophallus (Ac1 and Ac2: length of arch of cingulum; Fx: dorso-ventral width of flexure; Gpr: width of gonopore process). 5, Ventral view of endophallus (Ap: length of apical valve; ApW: width of apical valve; Fx+Bp: longitudinal length of flexure + basal valve). 6, Dorsal view of endophallus (Ac3 and Ac4: width of arch of cingulum; Ar: width of arms to apical valve; Bp: lateral width of half of basal valve). 7, Ectophallic sclerite (EcH: horizontal width of half; EcC: horizontal width of midprojection). 8, Ovipositor (LaL: longitudinal length of lateral apo- deme; LaW: width of lateral apodeme). ber. Twenty different dimensions of male genitalia and length and width of the lateral apodemes of ovipositors were measured us- ing ocular micrometer attached to a dis- secting scope (Figs. 1-8). Because the phal- lic complex is comprised of several parts, the dimensions of the cingulum were mea- sured first, and the endophallus was dis- sected afterwards. Terminology used here followed that of Dirsh (1956). All measure- ments were logarithmically transformed to normalize variance and plotted against the days after emergence using MINITAB. RESULTS Developmental sequence of the genita- lia.—Differences between the genitalia of each specimen and those of a sexually ma- ture male were documented in order to de- scribe the developmental stage of a certain age. Although most individuals followed similar patterns, some seemed to develop more slowly than others. External oviposi- tors did not change, but internal lateral apo- demes muscles differentiated through time. However, it was not possible to document the cuticle deposition sequence grew as 184 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of these lateral apodemes, because they were still covered with muscles. Day | in- dicates the first day after the emergence. Day O-—1: No structure was sclerotized and the color of phallic complex was opaque and white. Epiphallus was fully formed, but the cuticle was very thin and white in color. Cingulum, ectophallic sclerite, and endo- phallus were not fully formed and appeared to be very fragile. The arch of cingulum in endophallus was absent. Day 2: All parts started to be sclerotized, and structures were light brown in color. Tracheae were found within the phallus. In cingulum, the lateral parts of rami were white, whereas others parts were all brown. Day 3: The lateral parts of rami began to be filled with cuticle and were brown. Day 4: All structures were more robust due to cuticle deposition. In one male, the distal projection of the arch of cingulum had start- ed to develop. It was very thin and cylin- drical and appeared to originate from below both zygoma and the base of arch of cin- gulum. Day 5: The structure found in one male on day 4 appeared in all males. Day 6-7: More cuticle deposition was in process, but not much different from the day 5. The amount of cuticle deposition varied as if certain individuals had devel- oped faster than others. Day 8-9: The arch of cingulum became gradually thicker and larger compared to the same structure of earlier days. Day 10: Cuticles became more rigid, and anterior projection of epiphallus was hard- ened. Day 11: The anterior projection of epi- phallus began to have double layers inter- nally. The hour-glass shaped portion below zygoma became smaller. The arch of cin- gulum was getting thicker, but still a cylin- drical shape. Day 12-13: The arch of cingulum became gradually thicker than the same structure of earlier days, and other parts became more robust. Day 14-16: Tip of anterior projection of epiphallus became darker, and the hour- glass shaped portion below zygoma became even smaller. Day 17-20: Morphology of phallus was similar to that of a fully mature male. The arch of cingulum became bulbous. External coloration of locusts was brown still indi- cating sexual immaturity. Day 21: Some locusts turned bright yellow, indicating sexual maturity. Day 22—30: Most locusts turned yellow and all of these had a fully mature phallus. Dur- ing this period, occasional copulation and Oviposition were observed. On day 30, there were still males with brown colora- tion, which did not have a fully mature phallus. Day 31-35: All the locusts were bright yel- low and the phallic complex was highly sclerotized and mature. Rate of development.—Cuticle deposi- tion occurred throughout the parts of phallic complex during sexual maturation, resulting in an increase in thickness (Figs. 9-12). Of twenty dimensions measured, most dimen- sions did not result in significant changes during development. For some dimensions, however, cuticle deposition was significant enough to change the shape of the structure. In cingulum, overall size remained constant throughout maturation period (CW in Fig. 9), but the apodemes of cingulum (ApdR in Fig. 9) became thicker and wider as the in- sect matured (Fig. 13). Likewise, overall size of epiphallus remained constant (EpW in Fig. 10), but there was a gradual increase in size of the bulbous base of epiphallus (EpB in Fig. 10, Fig. 14). Resulting devel- opmental patterns of dramatically changing dimensions in both cingulum and epiphal- lus may be represented as a curve where growth approaches an asymptote. Endo- phallus showed the most qualitative chang- es, resulting in dramatic morphological changes. Laterally, the distal projection of the arch of cingulum, which was absent for the first three days after emergence, sud- VOLUME 106, NUMBER 1 9 ZosePosogrergegegdortegeceeggecers cw e) 90920 fe) eae ogcerteoseses (are g 88 BR =o 2 ae ApdR Oo Log length | (e} @ @ ie} oo @ i am T T T T 0) 5 1 qd 2 255 ei) <6 Days after emergence 11 2.0 + Ap prseecengececes esos: cossgooessocs « 90880096880 9388s gs00Rabgggooe8e Aci ie) a eae Log length 3 aaealhs fo) us ais mil T T T T T T 0 5 10 15 20 25 30 35 Days after emergence Figs. 9-12. 10 | 8g089a 8080800896 °9088 9860089688909 98 EpWw Log length | T alas T a lem T T T 0) 5 10 iB 20 2 20 25 Days after emergence 12 22 SI 9908000 808888eeggoaess © 2.144 g® 38 o 008 8 £ fo) Qo 20 | 9988 8 Fx+Bp Pais 7 18 © 17 eset = OD Soars: otoece a | ey LOS A Rn H LO 5 ae « SAF! ost sos taal es 134 *,, 1.2 a ; Ti pam T T laa) & la ce 0 5 10 15720) 9250) eso Days after emergence Logarithmically transformed measurements were plotted against days after adult-emergence to show the developmental patterns of each measured structure. In each figure, dimensions from the same genitalic structure were plotted. 9, In cingulum, entire width of cingulum (CW) remains constant whereas width of apodeme ridge (ApdR) changes. ApdR starts to mature around 14 days after emergence. 10, In epiphallus, width (EpW) remains constant whereas width of bulbous base (EpB) gradually develops over time and starts to mature around 26 days after emergence. 11, In endophallus, length of apical valve (Ap) remains constant, whereas arch of cingulum (Ac1) suddenly appears at day 4 after emergence. Arch of cingulum does not grow lengthwise after it forms, but it gradually thickens until 20 days after emergence (not shown). 12, In endophallus, both width of basal valve (Bp) and length of flexure + basal valve (Fx+Bp) gradually increase and start to mature around 10 days after emergence. denly began developing four days after the emersence (Aci in Fig. 11, Fig. 15). It started as a thin cylindrical structure and became bulbous (Fig. 15). The only non- changing part was the length of apical valve of the penis (Ap in Fig. 11, Figs. 15, 16). Ventrally, the most significant morphologi- cal changes can be found in gonopore pro- cess and basal valve (Bp in Fig. 12, Fig. 16). These structures were not differentiated immediately after the adult emergence, but rapidly developed until ten days after emer- gence (Fig. 12). Overall, the size of endo- phallus increased during maturation mainly due to the growth of flexure and basal valve (Figs. 12, 15, 16). The distal part of endo- phallus remained relatively constant, and the proximal part elongated through cuticle deposition. The ejaculatory sac was present immediately after the emergence, but the cellular maturity was not measured (Figs. 15, 16). Ectophallic sclerite did not change in size except for the increase in cuticle thickness. In females, the lateral apodemes 186 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of ovipositor continued to develop in length and width after emergence (Fig. 17). The external valves did not change in size ex- cept that the muscle mass inside the valves increased. Post-emergence development of male genitalia in other species.—The phallic complexes of Schistocerca americana and S. gregaria are very similar, but the phallic complex of L. migratoria differ morpholog- ically. The size of the whole insect was sim- ilar between Schistocerca and Locusta, but L. migratoria has a phallic complex three times as large as that of Schistocerca. The entire phallic complex of L. migratoria is highly sclerotized immediately after emer- gence, indicated by the brown color of cu- ticle. Figures 18a and 19a show the develop- ment of the cingulum in S. gregaria and L. migratoria, based on specimens that were freshly molted, 2 weeks after molting, and 35 days after molting. In both cases, the apodemes of cingulum increased in width and length. The distal portion of cingulum also remained relatively unchanged, where- as the increase in cuticle deposition of the proximal portion was dramatic. In both Schistocerca species, width of the bulbous base of epiphallus in width through time. In L. migratoria, no structure increased in width, but thickness of cuticle increased through time. In both Schistocerca species, the arch of the cingulum was absent im- mediately after emergence and progressed from a cylindrical to a bulbous shape (Figs. 15, 18b). Both basal valve of penis and gonopore process increased in width and length. There seemed to be no equivalent structure to the arch of cingulum in L. mig- ratoria. The apical valve of penis remained constant, but the basal valve of penis went through dramatic structural changes (Fig. 19b). DISCUSSION Genitalia as apodemes.—This study sug- gests that adult grasshoppers are function- ally incapable of mating during sexual mat- uration. Sexual maturity is here defined as a period when both males and females can functionally copulate and produce viable offspring. Each component of the phallic complex experiences a structural change that seems to be closely associated with the muscles responsible for movement during copulation. Growth of the phallic apodemes and lateral apodemes of the ovipositor in Schistocerca americana is accompanied by an increase in muscle mass. Elastic nature and cuticle deposition patterns all indicate that the male genitalia and the lateral apo- demes of ovipositor probably contain resilin (Keffer and Babcock 1998, Neville 1963b, Tatham and Shewry 2002). In freshly emerged specimens, cuticle deposition has yet to occur and muscle cells have not dif- ferentiated. Timing of the complete devel- opment of genitalia and timing of sexual maturity coincide, suggesting that mature genitalia are the functional necessity of the copulation. In the colony where both sex- ually immature and mature specimens were reared together, copulation was observed only between bright yellow individuals, which always had fully mature genitalic structures. Until now, delayed mating has been explained only by the developmental time of soft reproductive structures such as epithelium, ejaculatory duct, and ovaries (Norris 1954, 1957). While these structures are crucial for reproduction, skeletally ma- ture structures that are responsible for cop- ulation, ejaculation, and oviposition are es- sential for organisms to function properly. This study thus proposes a new proximate causal reason for the delayed mating. Post-emergence genitalic development is widespread.—Post-adult emergence devel- opment of genitalic structures appears to be widespread, at least in Acrididae. Schisto- cerca and Locusta belong to two subfami- lies, Cyrtacanthacridinae and Oedipodinae, respectively, and their genitalic morpholo- gies differ greatly. However, both genera follow similar developmental patterns in terms of genitalia. For example, the epi- phallus, ectophallic sclerite, and cingulum VOLUME 106, NUMBER 1 187 16 Emergence Maturity Figs. 13-16. Graphical representation of qualitative developmental changes in Schistocerca americana phallic com- plex (left: immediately after emergence; middle: two weeks after emergence; right: sexual maturity). 13, Cingulum: overall size remains constant, but apodemes of cingulum becomes thicker and wider. Cuticles on rami become thicker and more evident. 14, Epiphallus: overall size remains constant, but base gradually becomes bulbous and sclerotized. 15, Endophallus (lateral): arch of cingulum and gonopore process are not present in freshly emerged individual, but they appear in later developmental stages. Arch of cingulum progresses from a cylindrical to a bulbous shape. 16, Endophallus (ventral): basal valve gradually becomes longer and wider. 188 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Emergence Fig. 17. Maturity Graphical representation of qualitative developmental changes in Schistocerca americana ovipositor (left: immediately after emergence; middle: two weeks after emergence; right: sexual maturity). External ovi- positor does not change, but lateral apodemes grow longer and wider. remain relatively constant in size, and the proximal portion of endophallus experienc- es dramatic structural changes. How widespread is the post-adult emer- gence genitalic development? Neville (1983) listed the adult insects shown to have daily growth layers in the cuticle, most of which were hemimetabolous. Al- though he did not specifically study the in- ternal apodemes, he demonstrated that gen- italic development could be found in many hemimetabolous insects. Ridley (1989) doc- umented the time of mating after emer- gence for the most insect orders. Although there are some exceptions, hemimetabolous insects seem to have longer post-emergence maturation periods compared to holometab- olous insects. Most holometabolous insects mate immediately after emergence (Ridley 1989), perhaps because all the necessary development occurs during the pupal stage. To date, however, there is no study that fo- cuses on the post-adult emergence genitalic development in any insect group. This ne- glected area of study deserves more atten- tion because the post-adult emergence gen- italic development may in fact be very widespread. Taxonomic use of genitalia.—Historical- ly, the male genitalia have been used in in- sect taxonomy extensively (Eberhard 1985). In Acridoidea, many species have a distinct male genitalic morphology (Hubbell 1932, 1960; Dirsh 1956, 1973; Cohn and Cantrall VOLUME 106, NUMBER 1 189 18a Emergence Maturity Figs. 18-19. Comparison of post-adult emergence developmental patterns between Schistocerca gregaria and Locusta migratoria (left: immediately after emergence; middle: two weeks after emergence; right: sexual maturity). 18a, S. gregaria cingulum. As in S. americana, apodemes of cingulum become thicker and wider. 18b, S. gregaria endophallus (lateral). As in S. americana, arch of cingulum and gonopore process are not present in freshly emerged individual, but they appear in later developmental stages. 19a, L. migratoria cingulum. Freshly emerged individuals have more sclerotized phallus than Schistocerca, but likewise apodemes of cingulum gradually develop. 19b, L. migratoria endophallus (lateral). Basal valves start as a small structure, but ventrally elongates over time. Arch of cingulum is not found in Locusta. 190 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1974). In the North American grasshoppers, Schistocerca alutacea (Harris), S. lineata Scudder, and S. rubiginosa (Harris), male genitalia are the only reliable characters for species differentiation (Hubbell 1960). De- spite its significance, however, the variabil- ity of the phallic complex within a species is rarely mentioned in the taxonomic liter- ature (Linsley 1939, Shapiro 1978), even though morphological characters generally require a study of large samples. Perhaps, practical issues such as destruction of spec- imens and laborious preparation prevent taxonomists from examining a large num- ber of specimens. Developmental aspects need to be con- sidered when using genitalia in taxonomy, especially in Acridoidea. Most taxonomic studies deal with museum specimens in which researchers cannot know the exact age of the specimens. Although there is a method to estimate the age based on the age ring in cuticles (Neville 1963c), it is rarely used. The fact that sexually immature grass- hoppers have less developed genitalia than sexually mature ones presents a potential danger in using genitalia in taxonomy. If a species is initially defined using the less de- veloped genitalia, a later taxonomist may interpret the same species to be different based on the fully developed genitalia. In fact, Dirsh (1974: 166) may have defined Schistocerca braziliensis based on imma- ture male genitalia. His drawing of the en- dophallus clearly showed the characteristic of undeveloped arch of cingulum. It is also dangerous to define a species based on the length and width of apodemes because the lateral apodemes of the ovipositor clearly grow during sexual maturation. In Micro- lepidoptera, Busck (1931) used the length and the sclerotization of apophyses in char- acterizing families, but there is a possibility that these apodemes can develop after adult emergence (Neville 1983). Though these observations are based on a small number of species, if the phenomenon is_ wide- spread, one needs to be cautious about us- ing genitalic characters, because arrested cuticle deposition in museum specimens could be misleading. ACKNOWLEDGMENTS The experiment was conducted at the United States Department of Agriculture, Agriculture Research Service (USDA, ARS) in Sidney, Montana, from June 28 to August 1, 2001. I am grateful to Dr. Greg Sword for the use of USDA facilities, as well as for his invaluable advice. I thank my advisors, Drs. Norman Johnson and John Wenzel, for guidance and numerous suggestions. I also thank Dr. Stephen Simp- son and Stephen Roberts (University of Ox- ford) for generously providing specimens. I appreciate useful comments on this manu- script from Dr. Greg Sword and one anon- ymous reviewer. This study is a portion of my Master’s Thesis at the Ohio State Uni- versity. LITERATURE CITED Bursell, E. 1961. Post-teneral development of the tho- racic musculature in tsetse flies. Proceedings of the Royal Entomological Society of London (A) 36: 69-74. Busck, A. 1931. On the female genitalia of the Micro- lepidoptera and their importance in the classifi- cation and determination of these moths. Bulletin of the Brooklyn Entomological Society 26: 199— 216. Cohn, T. J. and I. J. Cantrall. 1974. Variation and spe- ciation in the grasshoppers of the Conalcaeini (Or- thoptera: Acrididae: Melanoplinae): The lowland forms of Western Mexico, the genus Barytettix. San Diego Society of Natural History, Memoir 6, 131 pp. Dirsh, V. M. 1956. The phallic complex in Acridoidea (Orthoptera) in relation to taxonomy. Transactions of the Royal Entomological Society of London 108: 223-356. . 1973. Genital organs in Acridomorphoidea (Insecta) as taxonomic character. Zeitschrift fiir Zoologische Systematik und Evolutionsforschung 11: 133-154. . 1974. Genus Schistocerca (Acridomorpha, In- secta). Dr. W. Junk B.V. Publishers, The Hague, 238 pp. Eberhard, W. G. 1985. Sexual Selection and Animal Genitalia. Harvard University Press, Massachu- setts, 244 pp. Finlayson, L. H. 1975. Development and degeneration, VOLUME 106, NUMBER 1 pp. 75-149. In Usherwood, P. N. R., ed. Insect Muscle. Academic Press, London, 621 pp. Hubbell, T. H. 1932. A revision of the Puer group of the North American genus Melanoplus, with re- marks on the taxonomic value of the concealed male genitalia in the Cyrtacanthacridinae (Orthop- tera, Acrididae). Miscellaneous Publications, Mu- seum of Zoology, University of Michigan 23: 1— 64. 1960. The sibling species of the Alutacea group of the bird-locust genus Schistocerca (Or- thoptera, Acrididae, Cyrtacanthacridinae). Miscel- laneous Publications Museum of Zoology, Uni- versity of Michigan 116: 1-91. Keffer, S. L. and S. K. Babcock. 1998. Resilin in wa- terscorpion male genitalia (Insecta: Heteroptera: Nepidae). American Zoologist 38: 165A. Linsley, A. W. 1939. Variation of insect genitalia. An- nals of the Entomological Society of America 32: 173-176. Neville, A. C. 1963a. Daily growth layers in locust rubber-like cuticle influenced by an external rhythm. Journal of Insect Physiology 9: 177-186. . 1963b. Growth and deposition of resilin and chitin in locust rubber-like cuticle. Journal of In- sect Physiology 9: 265-278. . 1963c. Daily growth layers for determining the age of grasshopper populations. Oikos 14: 1-8. . 1983. Daily cuticular growth layers and the teneral stage in adult insects: A review. Journal of Insect Physiology 29: 211-219. Norris, M. J. 1952. Reproduction in the desert locust (Schistocerca gregaria Forskal) in relation to den- sity and phase. Anti-Locust Bulletin 13: 1—51. . 1954. Sexual maturation in the desert locust 19] (Schistocerca gregaria Forskal) with special ref- erence to the effects of grouping. Anti-Locust Bul- letin 18: 1—44. . 1957. Factors affecting the rate of sexual mat- uration of the desert locust (Schistocerca gregaria Forskal) in the laboratory. Anti-Locust Bulletin 28: 1-26. Odhiambo, T. R. 1966. Ultrastructure of the develop- ment of the corpus allatum in the adult male of the desert locust. Journal of Insect Physiology 12: 995-1002. Ridley, M. 1988. Mating frequency and fecundity in insects. Biological Review 63: 509-549. . 1989. The timing and frequency of mating in insects. Animal Behaviour 37: 535-545. Schlein, Y. 1972. Factors that influence the post-emer- gence growth in Sarcophaga falculata. Journal of Insect Physiology 18: 199-209. Shapiro, A. M. 1978. The assumption of adaptivity in genital morphology. Journal of Research on the Lepidoptera 17: 68-72. Tatham, A. S. and P. R. Shewry. 2002. Comparative structures and properties of elastic proteins. Phil- osophical Transactions of the Royal Society of London (B) 357: 229-234. Viscuso, R., G. Longo, and L. Sottie. 1985. Ultrastruc- tural modifications in the ejaculatory duct epithe- lium of Eyprepocnemis plorans (Charp.) (Orthop- tera: Acrididae) during sexual maturation. Inter- national Journal of Insect Morphology and Em- bryology 14: 163-177. Weis-Fogh, T. 1952. Fat combustion and metabolic rate of flying locusts (Schistocerca gregaria For- skal). Philosophical Transactions of the Royal So- ciety of London (B) 237: 1—36. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 192-198 TAXONOMIC NOTES ON NORDLANDIELLA DIAZ AND GANASPIDIUM WELD (HYMENOPTERA: FIGITIDAE: EUCOILINAE) MATTHEW L. BUFFINGTON Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: mbuff@citrus.ucr.edu) Abstract.—Nordlandiella semirufa (Kieffer), n. comb., is described and figured. This represents the second species of Nordlandiella Diaz, and extends the distribution of this genus into Mexico, Arizona, Hawaii, Texas, Nicaragua, Belize, and Chile. Nordlandiella semirufa is reported here as a parasitoid of Melanagromyza sp. (Diptera: Agromyzidae), feeding in flower heads of Bidens spp. (Asteraceae) in Hawaii. The genus Ganaspidium Weld is reviewed; Ganaspidium utilis Beardsley is a new synonym of Ganaspidium nigrimanus (Kieffer), n. comb. Nordlanderia merickeli Miller and Nordlanderia navajoe Miller are recognized as nomina dubia. All known species of Ganaspidium and Nordlan- diella are parasitoids of Agromyzidae (Diptera). Key Words: myzidae, Cynipoidea In late 1999, three specimens of an un- identified species of eucoiline were sent to me by the late J.W. Beardsley. I recognized that they belonged to Nordlandiella Diaz, 1982, and I matched these specimens to several specimens on loan from the Amer- ican Entomological Institute (AEICC), Gainesville, FL and the Cornell University Insect Collection: (CUE), Ithaca, NW. These specimens, however, did not agree with the description of N. abdominalis Diaz, 1982, the sonly, species, placed min Nordlandiella. The holotype of Cothonas- pis semirufa Kieffer, 1907, agrees precisely with that of the unidentified specimens col- lected and reared in Hawaii, as well as ma- terial from the AEICC and CUIC. Origi- nally C. semirufa was placed in Cothonas- pis Hartig, 1840 (Kieffer 1907); Weld (1952) moved this species to Trybliographa Forster, 1869. This species does not possess any of the diagnostic features of either Cothonaspis (Nordlander, 1976) or Tryblio- Nordlandiella, Ganaspidium, Melanagromyza, Eucoilinae, Figitidae, Agro- grapha (Nordlander, 1981), but does pos- sess the diagnostic features of Nordlandiel- la. Thus, Nordlandiella semirufa (Kieffer), n. comb., is proposed. Since the original description of C. semirufa lacks mention of a number of critical features, a redescription of the species is provided below. The first Hawaiian specimens of Nord- landiella semirufa were collected by J.W. Beardsley and W.D. Peneira on 20 February 1996 while sweeping miscellaneous weeds and low crops at the University of Hawaii Agricultural Experiment Station farm, Wai- manalo, Oahu, HI. The species was recog- nized by J.W. Beardsley as being a newly established immigrant to Hawaii, since no other eucoilines like it had been recorded from Hawaii previously in extensive sur- veys (Yoshimoto 1962; Beardsley 1986, 1988, 1989). Additional female specimens were collected by W.D. Peneira during May of 1996. Host associations were made pos- sible by M. Ramadan, Hawaiian Depart- VOLUME 106, NUMBER 1 ment of Agriculture, who, in April of 1996, reared several female N. semirufa from the puparia of an unidentified species of Me- lanagromyza Hendel (Diptera: Agromyzi- dae) feeding in the flower heads of Bidens pillosa L. (Asteraceae); these collections were made at Kunia, Oahu, HI. Additional reared material was made available for my examination by M. Trostle, Department of Entomology, Texas A&M University (Col- lege Station, TX), who reared several males and females from a species of Melanagro- myza feeding in the flower heads of Bidens sp. collected on Oahu, HI. This paper includes a review of Ganas- pidium Weld as a follow up to Beardsley’s (1986) review of the genus. Similarly to Nordlandiella, all species of Ganaspidium have been recorded as primary parasitioids of Agromyzidae (Beardsley 1986). Current data suggests that Ganaspidium and Nord- landiella are not closely related (Fontal-Ca- zalla et al. 2002; Buffington, unpublished data). Descriptive terminology follows that of Fontal-Cazalla et al. (2002) and Buffington (2002). All newly acquired specimens were either card mounted or point mounted. Ex- amination of specimens was conducted us- ing a Leica MZ8 stereomicroscope illumi- nated with fluorescent desk lamps. Scan- ning electron micrographs were made using a Phillips XL-30; a full set of images for Nordlandiella and Ganaspidium are avail- able at http://www.morphbank.com (Mor- phbank server operated by Fredrik Ronqu- ist, Uppsala University, Sweden). Digital stereoscope images were made using a JVC digital camera mounted on a Zeiss SV6 ste- reoscope; images were optimized using Au- tomontage© 4.0 software. Specimens were borrowed from the fol- lowing institutions: AEICC: American Entomological Insti- tute, Gainesville, FL, USA. BPBM: Bernice P. Bishop Museum, Honolulu, HI, USA. 193 CASCC: California Academy of Scienc- es, San) Francisco; CA USA CuUIc: Cornell University Insect Col- lection, Ithaca, NY, USA. EMEC: Essig Museum Entomology Collection, University of Cali- fornia, Berkeley, CA, USA. MLPA: Museo de La Plata, La Plata, Argentina. WERE: University of California, Riv- erside, Research Collection, Riverside, CA, USA. USNM: National Museum of Natural History, Smithsonian Institu- tion, Washington DC, USA. Nordlandiella Diaz Nordlandiella Diaz 1982: 323-325. Type species, Nordlandiella abdominalis Diaz, by original designation. Diagnosis.—Key features unique to members of this genus concern the mor- phology of the frons adjacent to the toruli, and the morphology of the scutellar plate: prominent single dorso-ventral groove pre- sent adjacent to outer margin of each toru- lus (Fig. 1, arrow); small to medium sized protuberance (Fig. 2, arrow) present in cen- ter of scutellar disk, with mid-pit (Fig. 3, arrow) located posteriorly; broad pits sur- round protuberance, each containing a sin- gle seta (Fig. 3). Included species.—N. abdominalis Diaz, 1982; N. semirufa (Kieffer), new combi- nation. Nordlandiella semirufa (Kieffer), new combination (Figs. 1—4) Cothonaspis (Anectoclis) semirufa Kieffer 1907: 137. Lectotype hereby designated. Trybliographa semirufa: Weld 1952: 222. Lectotype.—For the purposes of nomen- clatural stability, one of the three specimens included in Kieffer’s (1907) description is herein designated lectotype, and the re- maining two specimens designated paralec- totypes. LECTOTYPE, 2, San Marcos, 194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. cates presence of dorso-ventral groove adjacent to torulus. 2, Mesosoma, lateral view, arrow indicates distinct protuberance in center of scutellar plate. 3, Mesosoma, dorsal view, arrow indicates posterior positioning of scutellar mid-pit. 4, General habitus of N. semirufa. 5, Head, anterior view, arrow indicates presence of clypeal and malar conical protuberances. 6, Mesosoma, dorsal view, arrow indicates scutellar plate. Nicaragua. Coll. Baker. CASC #10604; the specimen is in good condition with the lo- cality data label followed by Kieffer’s de- termination label (large red label), deposi- tory label and my lectotype label. Two ad- ditional specimens, which correspond with two additional collection localities in Kief- fer’s (1907) description, are hereby desig- 1—4, Nordlandiella semirufa. 5—6, Ganaspidium nigrimanus. 1, Head, anterior view, arrow indi- nated as paralectotypes: BELIZE: #5640 (1 2) (paralectotype); NICARAGUA: San Marcos, Coll. Baker, 5654 (paralectotype); each are labeled “paralectotype’. Diagnosis.—Nordlandiella semirufa has shorter grooves adjacent to the antennal socket (not extending past the midline of eye) than N. abdominalis (extending ven- VOLUME 106, NUMBER 1 trally past midline of eye) and N. semirufa possesses an entirely reddish-colored me- tasoma (all black to chestnut colored in WN. abdominalis). Description.—Head: Nearly glabrous with a few scattered setae on anterior facet of mandibles and anterior margin of clyp- eus. Upper and lower face lacking sculp- ture, save for a short, deep furrow lateral to each antennal socket. Malar sulcus a single groove. Malar space smooth, lacking any protuberance. Gena smooth and rounded. Antenna: Female with 13 segments, mo- niliform, slightly clavate; segments 3—12 subequal in size; segment 13 about 1.5 length of segment 12. Male with 15 seg- ments, segment 3 modified, laterally exca- vated, curved outwardly. Pronotum: Pronotal plate narrow, with a few scattered setae present dorsally; dorsal- ly crested, bifurcate; pronotal fovea open. Junction between pronotum and mesoscu- tum smooth and lacking sculpture. Lateral aspect of pronotum smooth, with a few scattered setae (Fig. 2). Mesoscutum: Smooth with very few scattered setae; lacking sculpture entirely (Figs). Mesopectus: Upper part and lower part of mesopleuron smooth and glabrous. Me- sopleural triangle an indistinct shallow im- pression. Mesopleural carina simple; lower part of mesopleuron bordered by a simple precoxal carina. Scutellum: Scutellar plate small, with mid-pit situated posteriorly; center of plate with a distinct protuberance (Fig. 2, arrow); pits bearing setae present on dorsal surface; posterior margin rounded. Dorsal surface of scutellum reticulate with large, irregular fo- vea and setae; margined slightly posteriorly and laterally; projections absent. Metapectal-propodeal complex: Meta- pectus glabrous with a few scattered setae present posteriorly. Dorsal margin of spi- racular groove well defined, ventral margin indistinct. Posterior margin of metapectus with a thin ridge. No other metapectal ridg- es present (Fig. 2). Anteroventral cavity 195 semi-circular, setose. Propodeum with short, thin setae; propodeal carinae non-par- allel, bent at junction with auxillary pro- podeal carinae; auxillary propodeal carinae indistinct. Nucha glabrous, reticulate. Wings: Hyaline, asetose basally with an increase in setae distally; margin with dis- tinct setal fringe (Fig. 4). RI vein complete; radial cell always closed. Legs: Fore- and midcoxae about equal in size; hind coxa slightly larger. All coxae variably covered in setae; mid- and hind coxae with distinct setal patches (Fig. 2). Femora and tibiae sparsely setose; tarso- meres with distinct, appressed setae. Metasoma: Female: Distinctly larger than mesosoma. Thin hairy ring present at base of syntergum, remainder of metasoma glabrous. Micropunctures present on pos- terior %4 of syntergum, and on remaining terga. Terga posterior to syntergum directed posteriorly at about 70° angle (Fig. 4). Male: as in female, but terga posterior to syntergum directed ventrally at a 90° angle. Material examined:—BELIZE [see types]. ECUADOR: Rio Chota, 10.VI.1965, 1,800 m, Luis Pefia (1 2; AEIC). MEXICO: Yau- tepec, Canyon d Lobos 13.11.1959, 4,000 ft, HE Evans (1 2 1 3; CUIC); Yautepec, Can- yon d Lobos 7.1II.1959, 4,000 ft, HE Evans (1 2 2 6; CUIC); Morelos, Tlayacapan, 29.X.1982, screen sweeping, J.T. Huber (1 2; UCRC); Chiapas, Puebla Nuevo, 20.IIT.1958, RC Bechtel, E.I. Schlinger (1 2; EMEC). NICARAGUA: [see types]. USA: FLORI- DA, Paradise Key, 5.1V.1951, H&M Townes (1 2; AEIC); Florida City, 27.11.1936 (2 9; AEIC); HAWAII: Oahu, Waimanalo at UH Farm, el. 60-80 ft, 20.11.1996, sweeping weeds & crops. JW Beardsley & WD Per- reira, (3 2; BPBM, 3 2; UCRC); Oahu, Wai- manalo at UH Farm, el. 60-80 ft, 15- 22.V.1996, yellow sticky borad trap, WD Per- reira, (2 2; BPBM); Oahu, Tantalus Dr., el. 1500’, on Bidens pilosa 28.V.1997, WD Per- reira, (1 2; BPBM); Oahu, Kunia, 8.IV.1999, ex Melanagromyza sp. in Bidens pilosa flow- er heads, M. Ramadan/99-137 (3 2°; BPBM); Oahu, Honolulu Co., Pali Lookout, 8 mi W 196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of Jonet Hwy HI and Hwy 61, 25.VI.2001, ex Bidens sp., MK Trostle and SC Ruth (14 oWERE): Distribution.—Neotropical and Nearctic regions: Belize, Ecuador, Mexico, Nicara- gua, USA (Florida, Hawaii (Oahu)) (see above list of localities). Biology.—I have examined specimens reared from Melanagromyza spp. (Agro- myzidae) infesting flower heads of Bidens pilosa L. (J.W. Beardsley and W.D. Perrei- ra, personal communication), and Melana- gromyza spp. infesting flower heads of Bi- dens sp. (M.K. Trostle, personal communi- cation). Ganaspidium Weld Ganaspidium Weld 1955: 274. Type spe- cies, Ganaspidium pusillae Weld, by original designation. Diagnosis.—Prominent conical protuber- ances present on the clypeal and malar space (Fig. 5); notaulices completely lack- ing (Fig. 6); hairy ring on base of metasoma present and complete. Ganaspidium can be separated from Nordlanderia Quinlan by the latter having well-developed notaulices and lacking a hairy ring at the base of the metasoma. Discussion.—Weld (1955) proposed this genus to accommodate a new species of eu- coiline (G. pusillae) reared from leaf min- ing flies in the Winter Garden area of south- ern Texas. Beardsley (1986) reviewed the genus, described one new species (G. utilis Beardsley) and wrote a key to Hawaiian species. Preliminary phylogenetic data (Buffington, unpublished data) weakly sup- ports the inclusion of Ganaspidium within the Gronotoma group of genera (sensu Fontal-Cazalla et al. 2002), a basal clade of Eucoilinae that specialize on parasitizing Agromyzidae. Ganaspidium hunteri and G. nigrimanus share a number of synapo- morphic features (e.g., a broad, flat scutellar plate with a large central mid-pit; Fig. 6), whereas G. pusilla is highly autoapomorph- ic (€.g., possessing a small, narrow scutellar plate with a pair of distinct tubercles on ei- ther side of the mid-pit). Upon comparison of the types for Eu- coela nigrimanus Kieffer, 1907, and Gan- aspidium utilis Beardsley, 1988, it was clear that these two species are synonymous (synonymy above). Kieffer (1907) most likely placed E. nigrimanus in Eucoila (also spelled Eucoela) due to broadness of that genus concept at the time. Weld (1952) moved E. nigrimanus to Pseudoeucoila Ashmead, which is in itself a junior syno- nym of Leptopilina Forster (Nordlander 1980). Type specimens for all species ex- amined are in good condition. Distribution.—Neotropical Region: Chi- le, Argentina, Panama, Costa Rica, southern Mexico. Nearctic Region: Northern Mexi- co, continental United States, southern Can- ada (all three described species). Indo-Pa- cific Region: Hawai. Biology.—Several species in the agro- myzid genus Liriomyza have been recorded as hosts (Weld 1955, Harding 1965, Beardsley 1986, Johnson 1987, Hara and Matayoshi 1990, Acosta and Cave 1994). Ganaspidium hunteri and G. nigrimanus have been evaluated for their usefulness in biological control (Johnson 1987, Lynch and Johnson 1987, Mason and Johnson 1988, Rathman et al. 1991, Rathman et al. 1995). Petcharat and Johnson (1988) stud- ied the larval stages of Ganaspidium nigri- manus. Included species.—G. hunteri (Craw- ford). Eucoila hunteri Crawford 1913: 310, holotype in USNM; Ganaspidium hunteri: Beardsley, 1986. G. nigrimanus (Kieffer), new combination. Eucoela nigrimanus Kieffer 1907: 138, holotype in CASC (#10573). Pseudoeucoila nigrimanus: Weld 1952: 235. Ganaspidium utilis Beardsley 1988: 44-46, holotype and paratypes in BPBM, new synonymy. G. pusillae Weld 1955: 274, holotype and paratypes in USNM. Examination of the type series of Ganaspidium pusillae revealed that one paratype is Disorygma pacifica (Yoshimo- to), reared from Liriomyza pusilla; this VOLUME 106, NUMBER 1 specimen bears the label ““Disorygma’”’ in Nordlander’s handwriting. Notes on the Status of Nearctic Nordlanderia Miller (1989) described two species of eucoiline wasps that were placed in Nord- landeria Quinlan (Quinlan 1986). Though the location of the type specimens for these two nominal species is unknown (Miller, personal communication), it is clear from the scanning electron micrographs that ac- company the descriptions that these two species possess many of the diagnostic fea- tures of Ganaspidium but not all of the di- agnostic features of MNordlanderia (e.g., both species lack notaulices on the meso- scutum and possess a complete hairy ring at base of metasoma). Therefore, N. nava- joe Miller and N. merickeli Miller are re- garded as nomina dubia until the holotypes can be located and compared to species of Ganaspidium. ACKNOWLEDGMENTS I thank those individuals who _ reared specimens of Nordlandiella semirufa and made them available for my examination (J.W. Beardsley, W.D. Perreira, M.K. Tros- tle and R. Wharton). The holotypes for Ganaspdium pusillae and G. hunteri were kindly loaned to me by Cathy Apgar and Dave Smith, Systematic Entomology Lab- oratory, USDA (USNM); the holotype for N. abdominalis was kindly loaned to my for my examination by Juan Schnack of the Museo de La Plata, Argentina; the holotype for Cothonaspis semirufa Kieffer was kind- ly lent for my examination by Keve Ribar- do of the California Academy of Sciences. Additionally, I thank Ms. Peggy Beardsley for making available data from the late J.W. Beardsley’s laboratory in Arcadia, CA. I thank the Heraty and Pinto labs at UC Riv- erside for their support in this project, as well as critical reviews of early drafts of this paper. Finally, I dedicate this paper to the late J.W. Beardsley, with whom I started this project some months prior to his death. 197 LITERATURE CITED Acosta, N. M. and R. D. Cave. 1994. Survey of par- asitoids attacking Liriomyza spp. (Diptera: Agro- myzidae) in southern Honduras. Revista de Biol- ogia Tropical 42: 203-218. Beardsley, J. W. 1986. Taxonomic notes on the genus Ganaspidium Weld (Hymenoptera: Cynipoidea: Eucoilidae). Proceedings of the Hawaiian Ento- mological Society 26: 35—39. . 1988. Eucoilid parasites of agromyzid leaf- miners in Hawaii (Hymenoptera: Cynipoidea). Proceedings of the Hawaiian Entomological So- ciety 28: 33-49. 1989. Hawaiian Eucoilidae (Hymenoptera: Cynipoidea), key to genera and taxonomic notes on apparently non-endemic species. Proceedings of the Hawaiian Entomological Society 29: 165— 194. Buffington, M. L. 2002. Description of Aegeseucoela Buffington, new name, with notes on the status of Gronotoma Forster (Hymenoptera: Figitidae: Eu- coilinae). Proceedings of the Entomological So- ciety of Washington 104: 589-601. Crawford, J. C. 1913. Descriptions of new Hymenop- tera, No. 7. Proceedings of the United States Na- tional Museum 45: 309-317. Diaz, N. 1982. Il. Nuevo género y especie de Himen- O6ptero Cinipoideo parasitoide de Melanogromyza minimoides Spencer (Cynipoidea: Eucoilidae), pp. 325-328. In Valladares, G., N. B. Diaz and L. DeSantis. 1982. Tres notas sobre dipteros agrom- icidos de la Republica de Argentina y sus Himen- Opteros parasitoides (Insecta). Revista de la Socie- dad Entomologica Argentina 41: 319—330. Fontal-Cazalla, FE M., M. Buffington, G. Nordlander, J. Liljeblad, P. Ros-Farré, J. L. Nieves-Aldrey, J. Pujade-Villar, and E Ronquist. 2002. Phylogeny of the Eucoilinae (Hymenoptera: Cynipoidea: Fig- itidae). Cladistics 18: 154—189. Hara, A. H. and S. Matayoshi. 1990. Parasitoids and predators of insect pests on Chrysanthemums in Hawaii. Proceedings of the Hawaiian Entomolog- ical Society 30: 53-58. Harding, J. A. 1965. Parasitism of the leaf miner Lir- iomyza munda in the Winter Garden area of Texas. Journal of Economic Entomology 58: 442—443. Johnson, M. W. 1987. Parasitization of Liriomyza spp. (Diptera: Agromyzidae) infesting commercial wa- termelon plantings in Hawaii. Journal of Econom- ic Entomology 80: 56—61. Kieffer, J. J. 1907. Beschreibung neuer parasitischer Cynipiden aus Zentral- und Nord-Amerika (En Aleman). Entomologische Zeitschrift 21: 70-162. Lynch, J. A. and M. W. Johnson. 1987. Stratified sam- pling of Liriomyza spp. (Diptera: Agromyzidae) and associated hymenopterous parasites on water- melon. Journal of Economic Entomology 80: 1254-1261. 198 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mason, G. A. and M. W. Johnson. 1988. Tolerance to permethrin and fenvalerate in hymenopterous par- asitoids associated with Liriomyza spp. (Diptera: Agromyzidae). Journal of Economic Entomology 81: 123-126. Miller, T. D. 1989. First Nearctic record of the genus Nordlanderia (Hymenoptera: Eucoilidae) with de- scriptions of two new species. Proceedings of the Entomological Society of Washington 91: 158— 163. Nordlander, G. 1980. Revision of the genus Leptopi- lina Forster, 1869 with notes on the status of some other genera (Hymenoptera: Cynipoidea: Eucoili- dae). Entomologica Scandinavica 11: 428—453. Petcharat, J. and M. Johnson. 1988. Biology of the leafminer parasitoid Ganaspidium utilis Beardsley (Hymenoptera. Eucoilidae). Annals of the Ento- mological Society of America 81: 477—480. Quinlan, J. 1986. A key to the Afrotropical genera of Eucoilidae (Hymenoptera), with a revision of cer- tain genera. Bulletin of the British Museum of Natural History, Entomology 52: 243-366. Rathman, R. J., M. W. Johnson, and B. E. Tabashnik. 1991. Production of Ganaspidium utilis (Hyme- noptera: Eucoilidae) for biological control of Lir- iomyza spp. (Diptera: Agromyzidae). Biological Control 1: 256—260. Rathman, R. J.. M. W. Johnson, B. E. Tabashnik, and K. H. Spollen. 1995. Variation in susceptibility to insecticides in the leafminer parasitoid Ganaspi- dium utilis (Hymenoptera, Eucoilidae). Journal of Economic Entomology 88: 475-479. Weld, L. H. 1952. Cynipoidea (Hym.) 1905-1950. Pri- vately Printed, Ann Arbor, Michigan, 351 pp. 1955. A new genus and species of North American Cynipoidea (Hymenoptera). Proceed- ings of the Entomological Society of Washington 57: 274. Yoshimoto, C. 1962. Revision of the Hawaiian Eu- coilinae. Pacific Insects 4: 799-845. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 199-201 POLLEN TRANSPORT BY NORTH AMERICAN TRICHODES HERBST (COLEOPTERA: CLERIDAE) JONATHAN R. MAWDSLEY Department of Systematic Biology, Entomology Section, National Museum of Natural History, Smithsonian Institution, PO. Box 37012, Washington, DC 20013-7012, U.S.A. (e-mail: mawdsley.jonathan @nmunh.si.edu) Abstract.—Adults of the 11 North American species in the genus Trichodes Herbst (Coleoptera: Cleridae) readily acquire pollen while visiting flowers. On museum speci- mens, pollen grains are concentrated primarily on the vestiture of the pronotum and legs, although pollen is also found on the pubescence at the base of the elytra, the frons, and the ventral surface. Field observations of Trichodes ornatus (Say) in Colorado indicate that adults often move between flowers, and suggest that individual populations of these clerids may show some fidelity to particular plant species. Trichodes species may be important pollinators of native plant species in North America. Key Words: Beetles in the genus Trichodes Herbst (Coleoptera: Cleridae) are widely known as flower visitors (Corporaal 1950, Foster 1976, Arnett and Jacques 1981). At flowers, these large, colorful clerids feed on pollen, prey on other insects, mate, and oviposit (Foster 1976; Mawdsley, unpublished ob- servations). Although most Trichodes spe- cies are densely pubescent, the possibility that these beetles may pollinate flowers has not been previously investigated. Species in the related family Melyridae are known to be important pollinators (Grant and Grant 1965), and may be more efficient and ef- fective than honeybees under favorable cir- cumstances (du Toit 1990). As a part of my ongoing investigations into the pollination ecology of beetles in the superfamily Cleroidea, I examined pinned adult specimens of North American Tricho- des species in the collection of the National Museum of Natural History, Smithsonian Institution (NMNH). It quickly became ap- parent that almost every specimen in this Trichodes, Cleridae, Coleoptera, pollen, pollination, pollinator large collection had pollen grains present in its vestiture. Many of the specimens were densely coated with pollen. Data from these museum specimens, combined with my own field observations on Trichodes orna- tus (Say) in 1999, suggest that Trichodes species may be significant pollinators of na- tive plant species in North America. EXAMINATION OF MUSEUM SPECIMENS I examined the entire collection of North American Trichodes in NMNH. Almost all of the specimens in this collection (includ- ing specimens of all 11 currently-recog- nized North American species of Tricho- des) had at least some pollen grains trapped in the dorsal vestiture. Of the 140 speci- mens that were labeled as having been col- lected on particular species or genera of plants, all 140 bore pollen. In Table 1, I provide a list of the flowers on which these pollen-bearing clerids were collected. Trichodes species are densely pubescent. Most specimens examined had _ pollen 200 Table 1. Pinned specimens of North American 77ri- chodes in NMNH were collected on flowers of the following plant species. All of these specimens bore pollen grains. NMNH specimens of 7. bicinctus Greene, 7. bimaculatus LeConte, and T. nexus Wolcott also bore pollen grains, but these specimens lacked information about associated plant species. Informa- tion in parentheses includes state, locality, and number of beetle specimens. Botanical nomenclature follows the online database of the Missouri Botanical Garden (http://mobot.mobot.org/W3T/Search/vast.html). Trichodes apivorus Germar Carduus sp. (VA: Augusta Co., 1) Ceanothus sp. (VA: Falls Church, 3; Glencarlyn, 5) Erigeron sp. (FL: Liberty Co., Rt. 270, 5 miles W Regi2 ss) Trichodes bibalteatus LeConte Acacia greggti Gray (TX: Midland, 3) Castilleja sulphurea Rydberg (TX: 5 miles E Greg- ory, 1) Condalia obtusifolia Hooker (TX: Starr Co., 8) Coreopsis cardaminefolia (DeCandolle) Nuttall (TX: San Antonio, 1) Hedeoma sp. (TX: Cotulla, 1) Hymenocarpus sp. (TX: near Uvalde, 5) Monarda citriodora Cervantes (TX: Clarendon, 1) Opuntia sp. (TX: Cotulla, 8) Prosopis glandulosa Torrey (TX: Scarry Co., Ful- ler Ranch 1) Prosopis juliflora (Swartz) DeCandolle (TX: Starr Co., 4) Prosopis sp. (TX: Guthrie Co., 1; Lubbock Co., Lubbock, Couth Ranch, 1) Ratibida columnaris (Pursh) D. Don (TX: Cotulla, 3) Rudbeckia amplexicaulis Vahl (TX: Denton, 1) Rudbeckia sp. (TX: Dallas, 1) Verbesina encelioides (Cavanilles) Gray (TX: Co- tulla, 2) Trichodes nutalli Kirby Chrysanthemum leucanthemum L. (NY: Virgil, 1) Heracleum lanatum Michaux (CO: Boulder, 1) Solidago sp. (WI: Waupaca, 1) Trichodes oregonensis Barr Achillea millefolium L. (OR: Jackson Co., 12 miles NE Ashland, 9; 15 miles NE Ashland, 3; 21 miles NE Ashland, 11) Trichodes oresterus Wolcott Condalia obtusifolia Hooker (TX: Starr Co., 1) Opuntia sp. (TX: Uvalde, 1) Prosopis sp. (TX: Presidio, 1) Trichodes ornatus (Say) Achillea millefolium L. (1D: Spencer, 1) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Adenostema sp. (CA: West fork of San Gabriel River, 2) Aronia sp. (MT: Missoula, 1) Asclepias sp. (C: Hotchkiss, Rogers Mesa, 1) Brassica nigra (L.) Koch (OR: Talent, 1) Ceanothus fendleri Gray (AZ: Flagstaff, Fort Val- ley, 4) Ceanothus sp. (OR: Butte Falls, 1) Eucnide urens (Gray) Parry (AZ: Mohave Co., 2.8 miles E Willow Beach, 1; Kingman Wash, 1; Willow Beach, 2. CA: San Bernardino Co., 1.7 miles W Parker Dam, 4; 11 miles N Earp, 1. NV: Clarke Co., 2.3 miles NW Hoover Dam, 7) Fallugia sp. (AZ: Grand View, Grand Canyon, 1) Heracleum lanatum Michaux (ID: Lakeview, 4) Tris sp. (AZ: Flagstaff, 10) Mentzelia involucrata Watson (CA: Riverside Co., Desert Hot Springs, 1) Opuntia sp. (AZ: Sabino Canyon, 10) Prunus demissa (Nuttall) Walpers (OR: Talent, 1) Rosa sp. (NV: Baker, 1) Rubus sp. (CO: Longview, 1) Sphaeralcea sp. (NV: Mercury, Nevada Test Site, 1) Taraxacum officinale Weber (WY: Molson, 1) Trichodes peninsularis horni Wolcott and Chapin Chrysothamnus sp. (AZ: Maricopa Co., Phoenix, Upper Moon Valley, 1) Desmanthus sp. (AZ: Nogales, 1) Gutierrezia sp. (AZ: Douglas, 1) Verbesina encelioides (Cavanilles) Bentham and Hooker (AZ: Douglas, 1) Trichodes simulator Horn Chrysothamnus nauseosus (Pallas) Britton (UT: Beaver, 1; Iron Co., 3 miles E of Cedar City, 2) Chrysothamnus sp. (UT: Ft. Duchesne, 1) grains adhering to the dense pronotal hairs and to the shorter hairs on the ventral sur- face of the tibiae. However, pollen grains also commonly adhered to the vestiture of the frons, the base of the elytra, the venter, and the rest of the legs, and many speci- mens were uniformly covered with a dense coat of pollen. FIELD OBSERVATIONS I collected adults of Trichodes ornatus from flowers in open woodlands of Pinus ponderosa Douglas in Estes Park, Larimer County, Colorado, during the last week of VOLUME 106, NUMBER 1 June, 1999. My first collecting site was de- scribed elsewhere (Mawdsley 1999): anoth- er study area of comparable size was iden- tified in similar forest on private land in an- other portion of the Estes Park valley. Adults at both sites showed a strong pref- erence for flowers of Potentilla diversifolia Lehmann. At the first site, 15 adults of 7. ornatus were collected on flowers of P. div- ersifolia, while 2 adults were found on Ru- bus parviflorus Nuttall. At the second site, 12 adults of 7. ornatus were found on P. diversifolia, while 1 adult was found on R. parviflorus. At both sites, clerids were not found on other seemingly suitable nectar and pollen sources, including flowers of Achillea lanulosa Nuttall, Gaillardia spp., and Potentilla fruticosa L., during the seven days of observation. Adults of 7. ornatus at both sites actively moved about the flowers and were often ob- served flying between flowers. W. EF. Barr (in litt.) also indicated that Trichodes spe- cies frequently move about and between flowers while feeding. DISCUSSION My examinations of museum specimens indicate that North American Trichodes readily acquire pollen grains while visiting flowers. Given the large quantities of pollen observed on some museum specimens, and given that adult Trichodes often move be- tween flowers, it seems likely that these clerids are responsible for at least some pol- lination of the flowers that they visit. It also appears that individual popula- tions of these beetles may preferentially vis- it certain flower species, as suggested by my observations on Trichodes ornatus in the Estes Park valley. Some of the data in Table 1 provide additional support for this hypothesis. There is, for example, an evi- dent association of northern Virginia pop- ulations of T. apivorus with Ceanothus sp.; an association of T. bibalteatus with Con- 201 dalia obtusifolia and Prosopis juliflora in Starr Co., Texas, and with Opuntia sp. at Cotulla, Texas; an association of T. orego- nensis with Achillea millefolium; an asso- ciation of 7. ornatus with Eucnide urens in the Mojave Desert and with Opuntia sp. in Sabino Canyon, Arizona; and an associa- tion of 7. simulator with Chrysothamnus nauseosus in Utah. Fidelity to individual plant species, even if on a local population level, would mean that these clerids are probably more effective pollinators than other, more generalist beetle species. ACKNOWLEDGMENTS I thank Terry L. Erwin for sponsoring my continuing work at the Smithsonian Insti- tution as a Research Associate and also thank Natalia J. Vandenberg for making the NMNH clerid collection available for study. William E Barr provided valuable infor- mation on Trichodes biology during my earlier studies of mimicry in Cleridae. Charles A. and Katherine S. Johnson pro- vided assistance with field studies in Col- orado. LITERATURE CITED Arnett, R. H. and R. L. Jacques. 1981. Simon and Schuster’s Guide to Insects. Simon and Schuster, New York, 511 pp. Corporaal, J. B. 1950. Coleopterorum Catalogus, Pars 23 (Editio Secunda), Cleridae. W. Junk, The Hague, 373 pp. du Toit, A. P. 1990. The importance of certain insects as pollinators of sunflower (Helianthus annuus L.). South African Journal of Plant and Soil 7(3): 159-162. Foster, D. E. 1976. Revision of North American 77ri- chodes (Herbst) (Coleoptera: Cleridae). Special Publications of the Museum, Texas Tech Univer- sity 11: 1-86. Grant, V. E. and K. A. Grant. 1965. Flower pollination in the phlox family. Columbia University Press, xi + 180 pp. Mawdsley, J. R. 1999. Redescription and notes on the biology of Amecocerus senilis (LeConte) (Cole- optera: Melyridae: Dasytinae). Journal of the New York Entomological Society 107(1): 68-72. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 202-216 SEVEN NEW SPECIES OF POLYPLECTROPUS ULMER (TRICHOPTERA: POLYCENTROPODIDAE) FROM COSTA RICA MARIA LOURDES CHAMORRO-LACAYO AND RALPH W. HOLZENTHAL Department of Entomology, University of Minnesota, 1980 Folwell Ave., Room 219 Hodson Hall, Saint Paul, MN 55108, U.S.A. (e-mail: cham0138 @umn.edu) Abstract.—Seven new species of Polyplectropus collected during an inventory of the Trichoptera of Costa Rica are described and illustrated: Polyplectropus clauseni, Poly- plectropus exilis, Polyplectropus hymenochilus, Polyplectropus kylistos, Polyplectropus paradelphae, Polyplectropus perpendicularis, and Polyplectropus yolandae. Key Words: Trichoptera, Polyplectropus, Polycentropodidae, Costa Rica, new species, taxonomy, systematics, Neotropics, biodiversity The genus Polyplectropus Ulmer 1905 is distributed worldwide, with representatives in Africa, the Orient, New Zealand, and tropical America (Morse 2001). The defin- ing adult character for the genus is the ab- sence of R2 (Fork I) in the hind wings. However, apparent differences in genitalia and larval characters between Old and New World species of Polyplectropus, leave un- certainties regarding the monophyly of Po- lyplectropus (Flint et al. 1999). The senior author will be addressing this issue in the future. Notwithstanding, the genus Polyplectro- pus presently has 138 species (Morse 2001), 93 in the Old World and 45 in the New World, mainly in the Neotropics (Flint et al. 1999). Earlier authors, including Ross (1941, 1947), Denning (1962), and Yama- moto (1966, 1967), described several New World species under the genus Polycentro- pus Curtis 1835, which were later trans- ferred to Polyplectropus by Flint (1968). Flint further synonymized the genera Ec- nomodellina Ulmer 1962 (= Ecnomodes Ulmer 1911) (Flint 1968) and Cordillop- syche Banks 1913 (Flint 1967) with Poly- plectropus. Bueno-Soria (1990) described 16 new species and revised the Mexican and Central American Polyplectropus spe- cies and found these to be monophyletic. Earlier and ongoing Trichoptera inven- tories in Costa Rica, Nicaragua, Venezuela, and Brazil by the authors and their col- leagues have yielded numerous additional new Polyplectropus species. Seven of these new species, all from Costa Rica, are de- scribed below. Previously, eight species of Polyplectropus were recorded from Costa Rica (Flint et al. 1999). Types are deposited in the University of Minnesota, Saint Paul, Minnesota (UMSP), the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (NMNH), and the Instituto Nacional de Biodiversidad, Heredia, Costa Rica (IN- BIO). The terminology used in this paper has been adapted from Hamilton (1986). Polyplectropus clauseni Chamorro-L. and Holzenthal, new species (Fig. 1) Polyplectropus clauseni is a member of the thilus group of Yamamoto (1967) based on its similarities to Polyplectropus deltoi- des (Yamamoto) 1967, especially the shape VOLUME 106, NUMBER 1 Fig. 1. pendage, caudal view. of the intermediate appendage, the dorso- lateral and the mesolateral processes of the preanal appendage, and the dorsolateral and ventromesal lobes of the inferior append- age. Polyplectropus clauseni can be distin- guished from P. deltoides by the shape of 203 Polyplectropus clauseni, male genitalia. A, Lateral view. B, Dorsal view. C, Ventral view. D, Caudal view of preanal appendage. E, Phallus. EK Apex of phallus, dorsal view. G, Ventromesal lobe of inferior ap- the ventromesal process of the preanal ap- pendage, which in lateral view appears bi- partite, with the dorsal lobe directed dor- sally and the ventral lobe directed ventrally, both similar in size and shape and bearing robust, sclerotized spines. It is also distin- 204 guished by the presence of median ventro- lateral spines on the phallus. Male.—Length of forewing 5.1 mm. Col- or of body and antennal scape yellowish brown; dorsum of head, pronotum, anterior half of mesothorax, tegula, and forewing base with long, erect brown and white se- tae; forewing with patches of brown, white, and golden setae, anterior edge bearing only patches of thick brown setae. Genitalia as in Fig. 1. Sternum IX, in lateral view with ventral margin nearly straight, posterior margin dorsally narrow, expanded medially, and becoming notched subventrally, ante- rior margin sinuate; in ventral view rect- angular, posterior margin with median hump flanked by lateral indentations. Ter- gum X membranous, broad, elongate, bear- ing dorsal microsetae. Intermediate append- age setose, in lateral view semicircular, ex- panding posteroventrally into lobe bearing apical setae; in dorsal view elongate, nar- row, slightly expanded basomesally. Pre- anal appendage apparently tripartite; dor- solateral process, heavily sclerotized, elon- gate, originating from dorsum of mesola- teral process, directed anteromesally, recurved posterolaterally, tapering mesally to acute apex; mesolateral process setose, in dorsal view digitate, apex rounded and di- rected laterally, in lateral view circular, ap- parently expanded into ventromesal pro- cess; ventromesal process, in lateral view appearing bipartite, with dorsal lobe direct- ed dorsally and ventral lobe directed ven- trally, both lobes similar in size and shape and bearing robust, sclerotized spines and pointed apices; in caudal view trapezoidal, with mesoventral corner bearing cluster of robust, sclerotized spines, ventrolateral margin rounded with additional robust, sclerotized spines along venter. Inferior ap- pendage divided into two lobes; dorsolat- eral lobe setose, in lateral view oval, broad, with posterodorsal margin rounded, in ven- tral view with lateral margin rounded, pro- duced into narrow apex, mesal margin nar- row, expanding into ventromesal lobe; ven- tromesal lobe short, setose, bearing cluster PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of robust, sclerotized spines, in lateral view with ventral margin nearly straight, poste- rior margin truncate, lobe barely exceeding origin of posterior margin of dorsolateral lobe, in ventral view triangular, mesal mar- gin straight, with semicircular ventromesal hump arising from base of inferior append- age. Phallobase short, narrow, with apico- ventral spines; phallic sclerite cylindrical with apex truncate, apex in dorsal view with median trough. Female.—Unknown. Type material.—Holotype, 6: COSTA RICA: Alajuela: Cerro Campana, Rio Bochinche, tributary, 6 km (air) NW Dos Rios, 10.945°N, 85.413°W, el. 600 m, 22— 23.vii.1987, Holzenthal, Morse, Clausen (UMSP) (UMSP000060983). Paratypes: Guanacaste: Parque Nacional Rinc6én de la Vieja, Quebrada Agua Apinolada, LO-7S9°N, 286.292- Ws. scl. 34/9 Siaeame 25.vi.1996, Holzenthal, Heyn, Armitage, 1 6 (UMSP); Guanacaste: Rio Los Aho- gados, 11.3 km, ENE Quebrada Grande, 10.865°N, 85.423°W, el. 470 m, 7.111.1986, Holzenthal & Fasth, | d (NMNH). Etymology.—Named in honor of Dr. Philip Clausen, Curator, University of Min- nesota Insect Collection, in recognition of his many years of service in caring for the collection. Polyplectropus exilis Chamorro-L. and Holzenthal, new species (Fig. 2) Polyplectropus exilis is a member of the charlesi group, as defined by Bueno-Soria (1990), and appears to be most closely re- lated to Polyplectropus beutelspacheri Bueno-Soria 1990, based on similarities in the shape of the preanal appendage and in the overall shape of the inferior appendage. Polyplectropus exilis differs from P. beu- telspacheri in the posteroventrally extended margin of the inferior appendage when viewed laterally, in the shorter and narrow- er ventromesal lobe of the inferior append- age, which is devoid of robust sclerotized apical spines, in the presence of a long, nar- VOLUME 106, NUMBER 1 Fig. 2. Polyplectropus exilis, male genitalia. A, Lateral view. B, 2C Ventral view. C, Dorsal view. D, Caudal view of preanal and intermediate appendages. E, Phallus and phallic spines. G, Apex of phallus, dorsal view. row, transparent, posteriorly directed pro- jection originating equidistantly from both lobes of the inferior appendage, and in the mesoventrally expanded sternum IX. Male.—Length of forewing 5.0—5.9 mm. Color of body yellowish brown, legs with long, yellowish-brown setae on femora and tarsi; dorsum of head, pronotum, and tegula with long, erect brown setae, apex of head with cluster of whitish, stout setae; fore- wing covered by brown setae and scattered patches of golden setae. Genitalia as in Fig. 206 2. Sternum IX, in lateral view with ventral margin straight, expanded posteriorly, pos- terior margin nearly straight, curving ven- trally, ventral margin slightly produced an- teriorly; in ventral view, nearly rectangular, posterior margin produced mesoventrally. Tergum X membranous, short. Intermediate appendage digitate, setose, not exceeding tergum X, produced posteromesally in dor- sal view. Preanal appendage bipartite; me- solateral process deltoid, setose, produced into mesoventral process with small papil- late lobe originating midway between both processes; mesoventral process basally broad, curving posteroventrally into acute apex. Inferior appendage divided into two lobes and medial process; dorsolateral lobe, in lateral view, club-shaped, setose, expand- ed posteroventrally, in ventral view elon- gate and narrow; ventromesal lobe setose, short, one third size of dorsolateral lobe, in lateral view narrow, tapering into rounded apex, in ventral view deltoid, directed pos- terolaterally; medial process arising equi- distant from both lobes, lightly sclerotized, tapering into acute apex, directed postero- mesally, longer than ventromesal lobe (me- dial process broken in some paratypes). Phallobase long, apically membranous; phallic sclerite cylindrical, sclerotized, nar- rowing into rounded apex, constricted sub- apically; phallic apparatus with approxi- mately six small phallic spines, unclear if embedded in phallic sclerite or endothecal membrane. Female.—Unknown. Type material.—Holotype: 6. COSTA RICA: Cartago: Reserva Tapanti, Quebrada Segunda at administration building, OGLE SS AST We rele 2508 mo = 10.v.1990, Holzenthal and Blahnik (UMSP) (UMSP000060974). Paratypes: Alajuela: Reserva Forestal San Ram6n, Rio San Lor- encito and tributaries, 10.216°N, 84.607°W, el. 980 m, 30.111—1.1v.1987, Holzenthal, Hamilton, Heyn, 2 ¢ (UMSP); Puntarenas: Rio Guineal, ca. 1 km (air), E. Finca He- lechales, 9.076°N, 83.092°W, el. 840 m, 4.vili. 1987, Holzenthal, Morse, Clausen, 3 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 6 (INBIO); same data as _ previous, 22.11.1986, Holzenthal, Morse, Fasth, 4 6 (NMNH); Puntarenas: Rio Cotén in Las Al- turas, 8.938°N, 83.826°W, el. 1,360 m, 18.11.1991, Holzenthal, Munoz, Huisman, 1 6 (UMSP); Puntarenas: Rio Singri, ca. 2 km (air) S. Finca Helechales, 9.057°N, 83.082°W, el: 720° m, 21.11:1986, Holzen- thal, Morse, Fasth, 6 ¢d (UMSP). Etymology.—Evxilis is the Latin word for weak or thin, which describes the lightly sclerotized medial processes arising from both lobes of the inferior appendages of the genitalia. Polyplectropus hymenochilus Chamorro-L. and Holzenthal, new species (Fig. 3) Polyplectropus hymenochilus belongs to the charlesi group, based on similarities with Polyplectropus mignonae Bueno-Soria 1990, especially the shape of the ventro- mesal lobe of the inferior appendage and the mesolateral and mesoventral processes of the preanal appendage. However, P. hy- menochilus can be easily distinguished from P. mignonae and other members of this group by the presence of a recurved dorsolateral process on the preanal append- age, by the membranous posteroventral ex- tension of the intermediate appendage, which surrounds the phallus, and by the rod-like dorsolateral lobe of the inferior ap- pendages, bearing a single apical spine. Male.—Length of forewing 4.5—4.7 mm. Color of body brown, antenna and legs yel- lowish brown; dorsum of head, pronotum, anterior half of mesothorax, and tegula with long, erect brown and white setae, anterior edge of forewing covered with dark brown setae, remainder of wing covered with gold- en setae and patches of white setae. Geni- talia as in Fig. 3. Sternum IX, in lateral view with ventral margin straight, posterior margin slightly sinuate, anterior margin slightly produced submedially; in ventral view, nearly rectangular with rounded an- terolateral margins. Tergum X membranous and bearing microsetae on dorsum, in lat- VOLUME 106, NUMBER 1 Biss 3: Caudal view of preanal and intermediate appendages. E, age, dorsal view. eral view elongate, in dorsal view with bi- lobed apex. Intermediate appendage digi- tate, setose, expanding posteroventrally into a membranous lip surrounding the phallus. Preanal appendage tripartite; dorsolateral process heavily sclerotized, elongate, orig- 207 3D Polyplectropus hymenochilus, male genitalia. A, Lateral view. B, Ventral view. C, Dorsal view. D, Phallus. EK Ventromesal process of the preanal append- inating from dorsum of mesolateral process, directed anteromesally, recurved posterolat- erally, tapering mesally to acute apex; me- solateral process setose, thinly sclerotized, in dorsal view digitate, directed posterolat- erally, in lateral view subquadrate, broad, 208 produced into setose ventromesal process; ventromesal process directed posteroven- trally, with highly sclerotized apex, pointed in lateral view, truncate in caudal view. In- ferior appendage divided into two lobes; dorsolateral lobe setose, in lateral view rod- like, base slightly broader than remainder two thirds, apex narrower and directed ven- trally, bearing single highly sclerotized api- cal spine, apex directed posteromesally; ventromesal lobe short, setose, half the size of dorsolateral lobe, in lateral view rectan- gular, basally narrow, posterior margin trun- cate and bearing six highly sclerotized spines (some specimens may have less); ventromesal lobe, in ventral view, quadrate, posterior margin transverse with many vis- ible spines, mesal margin broadening ba- sally and becoming pleated at base. Phallus short, ventrally membranous; phallic scler- ite cylindrical, narrowing to rounded apex, curved ventrally; endothecal membrane with approximately 20 small embedded phallic spines. Female.—Unknown. Type material—Holotype: 6. COSTA RICA: Guanacaste: Parque Nacional Guan- acaste, El Hacha, Quebrada Alcornoque, LLO09INS FS9:577° W:: el! 250 my! 2orvit. 1987, Holzenthal, Morse, Clausen (UMSP) (UMSP000060976). Paratypes: Same data as holotype, 1 d (INBIO), 1 gd (NMNH); Guanacaste: Parque Nacional Guanacaste, El] Hacha, Quebrada Pedregal, 10.983°N, 85.539°W, el. 300 m, 27.vii.1987, Holzen- thal, Morse, Clausen, 1 d6 (UMSP). Etymology.—Hymenochilus comes from the Greek words for membrane “‘hymeno”’ and lip or rim “*-chilus,”? which describes the weak membranous posteroventral ex- tension of the intermediate appendage, which surrounds the phallus. Polyplectropus kylistos Chamorro-L. and Holzenthal, new species (Fig. 4) Polyplectropus kylistos is closest to Po- lyplectropus charlesi (Ross) 1941 of the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON charlesi group based on similarities in the elongate posteromesal expansion of ster- num IX, the overall shape of the ventro- mesal lobe of the inferior appendage, and the presence of a pair of posterolaterally di- rected spines embedded in the endothecal membrane of the phallus. Polyplectropus kylistos can be distinguished from P. char- lesi by the semicircular shape and pointed apices of both lobes of the inferior append- age, and by the presence of a membranous projection bearing a single, robust, sclero- tized spine between the lobes of the inferior appendage. Male.—Length of forewing 4.6 mm. Col- or in alcohol of body, legs and antenna yel- lowish brown; dorsum of head, pronotum, and tegula with long, erect brown setae. Genitalia as in Fig. 4. Sternum [X with ven- tral margin nearly straight, produced pos- teromesally, length of expansion almost half length of inferior appendage; anterior margin in lateral view rounded, posterior margin sinuate, anterior margin in ventral view concave. Tergum X membranous, trapezoidal with dorsal microsetae. Inter- mediate appendage with digitate, setose posterodorsal lobe expanded mesally and ventrally into membrane; posteromedial lobe with apical setae. Preanal appendage bipartite; mesolateral process setose, in dor- sal view digitate, in lateral view nearly cir- cular, produced ventrally into hooked ven- tromesal process; ventromesal process se- tose, in lateral view with apex rounded and produced into ventral point, ventral margin straight, in caudal view with apex concave, with lateral and mesal sclerotized points. Inferior appendage divided into two lobes; dorsolateral lobes setose, in lateral view semicircular tapering apically into poster- oventrally directed spine, in ventral view produced basolaterally, tapering mesally into apical spine; ventromesal lobe broad, ventral margin rounded, produced into pos- teromesally directed point, basodorsal mar- gin membranous, bearing a membranous lobe with a robust sclerotized apical spine; mesoventral lobe variable, apex truncate, VOLUME 106, NUMBER 1 Fig. 4. 209 Polyplectropus kylistos, male genitalia. A, Lateral view. B, Ventral view. C, Dorsal view. D, Caudal view of preanal and intermediate appendages. E, Phallus, lateral view. F, Phallus, dorsal view. sometime bearing a spine on posterodorsal corner, in ventral view triangular, basally broad, with posteromesally directed, point- ed apex. Phallus long; phallic sclerite in lat- eral view cylindrical, narrowing apically and bent dorsally, apex with ventrally di- rected, highly sclerotized point; phallic sclerite in dorsal view rectangular, apex with posteromedial projection; endothecal membrane with pair of elongate, sclero- tized, posterolaterally directed spines orig- inating at base. 210 Female.—Unknown. Type material—Holotype, ¢d. COSTA RICA: Guanacaste: Parque Nacional Guan- acaste, Estacion _Pitilla, Rio Orosi, 10.991°N, 85.428°W, el. 700 m, 22- 25.v.1990, Holzenthal, Blahnik (UMSP) (UMSP000067800). Paratypes: Alajuela: Rio Pizote, ca. 5 km (air), S. Brasilia, 10.972°N, 85.345°W, el. 390 m, 12.111.1986, Holzenthal & Fasth, 1 ¢d (UMSP), 1 ¢ (NMNH). Etymology.—Kylistos comes from the Greek word for rolled or turned, which de- scribes the semicircular shape of the infe- rior appendages. Polyplectropus paradelphae Chamorro-L. and Holzenthal, new species (Fig. 5) Polyplectropus paradelphae is very sim- ilar to Polyplectropus yolandae n.sp. in the digitate ventromesal lobe and basally nar- row and apically broad dorsolateral lobe of the inferior appendage, in the mesoventrally directed digitate dorsolateral process of the intermediate appendage, and in the general form of the tripartite preanal appendage. Polyplectropus paradelphae can be distin- guished from P. yolandae by the broader ventromesal processes of the preanal ap- pendages, which do not meet mesally. In- stead, the mesal margins form an arc. It is also distinguished by the absence of a me- dian lobe on the intermediate appendage, by the sinuate dorsolateral process of the pre- anal appendages, by the more elongate dor- solateral lobe of the inferior appendage, and by the wider phallus, which has an apically narrow phallic sclerite. Male.—Length of forewing 5.0 mm. Col- or in alcohol of body yellowish brown. Genitalia as in Fig. 5. Sternum IX, in lateral view deltoid, ventral margin nearly straight, posterior margin sinuate. Tergum X mem- branous with microsetae on dorsum, in dor- sal view trapezoidal. Intermediate append- age digitate, setose, directed mesoventrally. Preanal appendage tripartite; dorsolateral process heavily sclerotized, elongate, sinu- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ate, directed anteromesally, recurved pos- terolaterally, curving ventrally, finally straightening posteriorly and tapering into acute apex; mesolateral process setose, in dorsal view digitate, directed posteromesal- ly, in lateral view nearly triangular, broad basally, produced into setose ventromesal process; ventromesal process, in lateral view broad, ending in pointed anteroven- trally directed apex, in caudal view digitate, directed ventrally, mesal margin forming an arc. Inferior appendage divided into two lobes; dorsolateral lobe setose, elongate, narrow basally, broadening apically, poste- rior margin rounded, in ventral view with anterodorsal margin bent mesally, exposing internal margin, medial setose lobe origi- nating from interior mesal margin; ventro- mesal lobe setose, narrow, digitate, less than one third size of dorsolateral lobe, with robust, sclerotized spines present on baso- mesal margin; base of inferior appendage elongate, narrow, produced anteriorly. Phal- lus membranous, stout; phallic sclerite curved ventrally, narrowing apically, mem- brane present dorsally. Female.—Unknown. Type material.—Holotype: ¢. COSTA RICA: Lim6n: Reserva Biolégica Hitoy- Cerere, Rio Cerere,; 9.67 1°N2283.0238; Weel: 90 m, 23-—24.111.1987, Holzenthal, Hamil- ton, Heyn, 1 d (UMSP). Etymology.—Paradelphae comes from the Greek words for beside “‘para-”’ and sis- ter ‘‘-adelphe.”’ This species was concealed among the specimens of Polyplectropus yo- landae, until it was discovered by the senior author after closer examination. Polyplectropus perpendicularis Chamorro-L. and Holzenthal, new species (Fig. 6) Polyplectropus perpendicularis belongs to the charlesi group with similarities to Polyplectropus mignonae Bueno-Soria 1990, especially the long and digitate inter- mediate appendage, the absence of dorso- lateral process, the overall shape and round- VOLUME 106, NUMBER 1 9D Fig. 5. Polyplectropus paradelphae, male genitalia. A, Lateral view. B, Ventral view. C, Dorsal view. D, Caudal view preanal and intermediate appendages. E, Phallus. ed mesolateral process of the ventromesal process of the preanal appendage, and the elliptical ventromesal lobe of the inferior appendage, which bears many robust, scler- otized spines on its apex. Polyplectropus perpendicularis can be distinguished from P. mignonae by the shape of the dorsolat- eral lobe of the inferior appendage, which is narrow, dorsoventrally elongate, perpen- dicularly produced posterad, and has a trun- cate apex. Male.—Length of forewing 5.8 mm. Col- 212 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Polyplectropus perpendicularis, male genitalia. A, Lateral view. B, Dorsal view. C, Ventral view. D, Caudal view of preanal and intermediate appendages. E, Phallus. K Apex of left dorsolateral lobe of inferior appendage, lateral view. G, Ventromesal lobe with robust setae of inferior appendage, caudal view. or of body dark brown, legs and antenna gitudinal w-shaped patch devoid of setae in yellowish brown; dorsum of head, prono- the region of the anal veins. Genitalia as in tum, tegula, and forewing base with long, Fig. 6. Sternum IX, in lateral view, deltoid, erect brown setae and a few white setae; posterior margin sinuate, anterior margin forewing with golden-brown setae, and lon- produced submedially, ventral margin near- VOLUME 106, NUMBER 1 ly straight; sternum in ventral view ovoid, lateral margin rounded, anterior margin concave, posterolateral corners produced posterad, posterior margin semicircular. Tergum X membranous, short, in dorsal view subquadrate. Intermediate appendage digitate, elongate, broad basally, narrow medially, truncate apex each bearing two setae, as long as or longer than inferior ap- pendage. Preanal appendage bipartite; me- solateral process setose, in lateral view cir- cular, slightly expanded posterad, in dorsal view nearly oval, slightly expanded later- ally, produced ventrally into ventromesal process; ventromesal process setose, in lat- eral view rectangular, with posterior margin truncate, expanded ventrally into sclero- tized point, ventral margin nearly straight; ventromesal process, in caudal view trian- gular, with dorsal setose lobes, lateral and mesal margins nearly straight, meeting pos- teriorly in a sclerotized point. Inferior ap- pendage divided into two lobes; dorsolat- eral lobe setose, in lateral view elongate dorsoventrally, perpendicularly bent poste- riorly, width continuous throughout length, apex truncate and bearing two robust, scler- otized spines, originating on corner, some- times subapically; dorsolateral lobe in ven- tral view flattened, expanded laterally, me- sal margin concave, subapically curved me- soventrally; ventromesal lobe setose, in lateral view elliptical, with elongate medial flange, apex rounded and bearing dorsal cluster of robust, sclerotized spines, in ven- tral view rectangular, lateral margin con- cave, posterior margin transverse, with ro- bust, sclerotized spines, six visible in ven- tral view, nine in caudal view, mesal margin nearly straight, broadening basally. Phallus long, largely membranous, narrowing api- cally; phallic sclerite cylindrical, apex near- ly truncate. Female.—Unknown. Type material.—Holotype, d. COSTA RICA: Puntarenas: Tributary to Rio Bella- vista in Las Alturas (road to quary) 8.952°N, 82.848°W, el. 1480 m, 19.111. 213 1991, Holzenthal, Munoz, Huisman (UMSP) (UMSP000060984). Etymology.—Perpendicularis comes from the Latin word for upright or at right angles, which describes the dorsolateral lobe of the inferior appendage, which is narrow, dorsoventrally elongate and_per- pendicularly produced. Polyplectropus yolandae Chamorro-L. and Holzenthal, new species (Fig. 7) Polyplectropus yolandae belongs to the thilus group based on similarities with Po- lyplectropus carolae Bueno-Soria 1990 and Polyplectropus denticulus Bueno-Soria 1990, especially the broad dorsal lobe and reduced, narrow ventromesal lobe of the in- ferior appendage. These species are also similar in the presence of the highly scler- otized, recurved dorsolateral process of the preanal appendage and in the overall shape of the preanal appendage and phallus. Po- lyplectropus yolandae can be distinguished from P. carolae and P. denticulus, and oth- er members of the thilus group by the me- soventrally directed, digitate lobe of the in- termediate appendage, by the more pro- longed and rounded posterior margin of the dorsolateral lobe of the inferior appendage, and by the row of robust, sclerotized spines along the dorsal margin of the ventromesal lobe of the inferior appendage. Male.—Length of forewing 4.8—5.0 mm. Color of body and antenna yellowish brown, legs with brown setae; dorsum of head, pronotum, anterior half of mesotho- rax, and tegula with long, erect brown, white, and golden setae; forewing with patches of brown, white, and golden setae, anterior edge bearing only brown setae. Genitalia as in Fig. 7. Sternum IX, in lateral view deltoid, produced anteroventrally, pos- terior and ventral margins straight; in ven- tral view nearly rectangular, posterior mar- gin nearly straight, narrowing laterally, an- teroventral corners rounded. Tergum X membranous, broad and elongate with mi- crosetae on dorsum. Intermediate append- 214 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Riga: age bipartite; dorsolateral process digitate, setose, directed mesoventrally, produced into a small median lobe flanking the phal- lus laterally and bearing apical setae. Preanal appendage tripartite; dorsolateral process heavily sclerotized, elongate, orig- Polyplectropus yolandae, male genitalia. A, Lateral view. B, Ventral view. C, Dorsal view. D, Caudal view of preanal and intermediate appendages. E, Phallus. inating from dorsum of mesolateral process, directed anteromesally, recurved posterolat- erally, tapering mesally to acute apex; me- solateral process setose, in dorsal view dig- itate, directed posteromesally, in lateral view oval, broad basally, produced into se- VOLUME 106, NUMBER 1 tose ventromesal process; ventromesal pro- cess, in lateral view, broad basally, tightly bent ventrally, narrowing medially, slightly broadening subapically, finally tapering to pointed apex; ventromesal process, in cau- dal view digitate, directed mesally, apex weakly recurved laterally. Inferior append- age divided into two lobes; dorsolateral lobe setose, basally narrow broadening api- cally, posterior margin rounded and slightly produced (in some specimens margin even- ly rounded); ventromesal lobe setose, nar- row, digitate, less than half length of dor- solateral lobe with robust, highly sclero- tized spines present on entire length of dor- sal margin; base of inferior appendage elongate, narrow, produced anteriorly. Phal- lus short, weakly sclerotized; phallic scler- ite robust, cylindrical, narrow subapically, broadening apically, apex truncate. Female.—Unknown. Type material.—Holotype: ¢. COSTA RICA: Guanacaste: Parque Nacional Guan- acaste, Estacion Maritza, Rio Tempisquito, 1OPS58°N, 85:497°W, el. 550 m,~ 30= 31.viii.1990, Huisman, Blahnik, Quesada (UMSP) (UMSPO000060980). Paratypes: Same data as holotype, 8 d (UMSP); same data as holotype, 19—20.vii.1987, el. 1,550 m, Holzenthal, Morse, Clausen, 2 ¢d (NMNH); Guanacaste: P.N. Guanacaste, Es- tacion Pitilla, Rio Orost, 85.428°W, el. 700 m, 22—25.v.1990, Hol- zenthal & Blahnik, 1 6 (UMSP); Guana- caste: PN. Guanacaste, Maritza, Rio Tem- pisquito Sur, 10.95°N, 85.48°W, el. 600 m, 30. viii. 1990, Huisman & Quesada, 3 d (IN- BIO); Guanacaste: PN. Guanacaste, ca. 0.7 km N. Estaci6n Maritza, 10.96°N, 85.50°W, el. 550 m, 30.vili.1990, Huisman & Que- sada, 1 6 (UMSP); Alajuela: Reserva For- estal San Ramon, Rio San Lorencito and tributaries, 10.216°N, 84.606°W, el. 980 m, 6—10.iii.1991, Holzenthal, Munoz, Huis- man, 2 6 (UMSP); Alajuela: P.N. Rincon de la Vieja, Quebrada Provisién, 10.769°N, 85.281°W, el. 810 m, 4.iii.1986, Holzenthal & Fasth, 1 6 (UMSP); Heredia: Rara Avis Biological Station, Quebrada Chiquiza, 10,991P°N, 10.229°N, 84.032°W, 31.i11.1989, el. 550 m, Blahnik & Solis, 1 ¢6 (UMSP). Etymology.——Named in honor of Jolanda Huisman for her contributions to the study of Trichoptera and for her collaboration in the Costa Rican inventory. ACKNOWLEDGMENTS We thank Dr. Oliver S. Flint, Jr., Smith- sonian Institution, for providing specimens and for advice regarding the manuscript. Thanks are also extended to Dr. Roger Blahnik for his advice and assistance. This material is based upon work supported by National Science Foundation Grants BSR- 8512368, BSR-8917684, and DEB- 9400632 (for field work in Costa Rica) and DEB-9977939 (a PEET grant to study Neo- tropical Trichoptera taxonomy). The man- uscript was completed while the senior au- thor held a research fellowship from the Smithsonian Institution. This support is gratefully acknowledged. LITERATURE CITED Banks, N. 1913. Synopses and descriptions of exotic Neuroptera. Transactions of the American Ento- mological Society 39: 201—242. Bueno-Soria, J. 1990. Estudios en Insectos Acuaticos VIII. Revisi6n para México y Centro América del Género Polyplectropus Ulmer (Trichoptera: Poly- centropodidae). Anales del Instituto de Biologia, Universidad Nacional Autonoma De México, Se- ries Zoologica 61(3): 357—404. Denning, D. G. 1962. New Trichoptera from México. Journal of the Kansas Entomological Society 35(4): 402-408. Flint, O. S. 1967. Studies of Neotropical caddisflies, V: Types of species described by Banks and Ha- gen. Proceedings of the United States National Museum 123 (3619): 1-37. . 1968. Bredin-Archbold-Smithsonian Biologi- cal Survey of Dominica, 9. The Trichoptera (Cad- disflies) of the Lesser Antilles. Proceedings of the United States National Museum 125(3665): 1—86. Flint, O. S. Jr, R. W. Holzenthal, and S. C. Harris. 1999. Catalog of the Neotropical caddisflies (In- secta: Trichoptera). Special Publication, Ohio Bi- ological Survey, Columbus, Ohio, 239 pp. Hamilton, S. W. 1986. Systematics and biogeography of the New World Polycentropus sensu stricto (Trichoptera: Polycentropodidae). Ph.D. Disserta- tion, Clemson University, Clemson, 257 pp. 216 Morse, J. C., Clemson University. 2001. Trichoptera World Checklist. — http://entweb.clemson.edu/ database/trichopt/index.htm Ross, H. H. 1941. Descriptions and records of North American Trichoptera. Transactions of the Amer- ican Entomological Society 67: 35—126. 1947. Descriptions and records of North American Trichoptera, with synoptic notes. Trans- actions of the American Entomological Society 73: 125-168. Ulmer, G. 1905. Zur Kenntniss Aussereuropaischer Trichopteren. Stettiner Entomologische Zeitung 66: 3-119. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 1911. Einige Siidamerikanische Trichopteren. Annals de la Société Entomologique du Belgique 55: 15-26. . 1962. Ein neuer name fiir Ecnomodes Ulm. (Trichoptera). Mitteilungen der Deutschen Ento- mologisch Gessellschaft 21: 5. Yamamoto, T. 1966. Five new species of the Caddisfly genus Polycentropus from South America (Tri- choptera: Polycentropodidae). Canadian Entomol- ogist 98: 908-912. . 1967. New species of the Caddisfly genus Po- lycentropus from Central America (Trichoptera: Polycentropodidae). Journal of the Kansas Ento- mological Society 40: 127—132. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 217-232 THE MOTHS (LEPIDOPTERA) AND ASSOCIATED FLORA OF KELLEYS ISLAND, LAKE ERIE B. K. COLEMAN, J. K. BISSELL, J. Crna, P MACKEIGAN, AND J. B. KEIPER The Cleveland Museum of Natural History, | Wade Oval Drive, Cleveland, OH 44106, U.S.A. (address correspondence to J. B. Keiper, e-mail: jkeiper@cmnh.org) Abstract.—We surveyed the nocturnal Lepidoptera of Kelleys Island (Ohio, Erie Coun- ty) within Lake Erie from 1999-2001, and found 216 species from 20 families. A loga- rithmic growth curve applied to our data suggests that the number of species obtained represents 65% of the number that would theoretically be captured after ten years of effort. We also list the 212 plant species found on the island to aid future efforts of moth/ host plant associations. Key Words: With more than 11,250 species recorded for North America (Hodges et al. 1988), moths (Insecta: Lepidoptera) provide an abundant food source as adults, larvae, and pupae to other trophic levels (1.e., predators and parasites), and constitute a significant proportion of terrestrial herbivores (Price 1997). The Ohio moth fauna has received a good deal of attention during the past two decades. Regional species lists have been compiled that give excellent data on the adult distribution and flight periods (e.g., Rings et al. 1987, Rings and Metzler 1989, Summerville et al. 1999, Teraguchi and Lu- blin 1999a-f, Rings and Downer 2001). Kelleys Island is one of the Lake Erie islands situated off the north coast of Ohio. A portion of the island owned by The Cleveland Museum of Natural History is known as the North Coleman Tract, and has been designated a conservation area. How- ever, other areas of the island are under de- velopment. We began a survey of the moths of Kelleys Island in an attempt to obtain data on this species-rich group before fur- ther environmental deterioration from hu- man activities occurs. The only moth spe- moths, Lepidoptera, vegetation, survey, Ohio cies list was prepared by Nault et al. (1989), when they captured |1 species from three families during sweep netting. Use of light traps is a commonly used and effective technique for collecting nocturnal Lepidop- tera, but transient species might be captured that do not breed in the trap area. Light trapping terrestrial insects on an island pro- vides a more accurate look at the number of species per unit area and habitat use, as it is reasonable to assume that all specimens captured originated from the island. We present the results of three years of light trapping for adult moths on the North Coleman Tract. Because of the strong tie between plants, moth oviposition, and lar- val development, we also give a listing of the vascular plants found on the island to aid future efforts of moth/host plant asso- ciations. MATERIALS AND METHODS Kelleys Island (OH, Erie County) is a 4.8 x 6.4 km island, positioned approximately 5 km north of the mainland city of Marble- head (Ottawa County). Moths were collect- ed with a BioQuip® 15-watt black light trap 218 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Moth taxa obtained in light trap from Kelleys Island, including Hodges’ number, the months col- lected, and the proportion of the total catch each species represented. A plus (+) indicates that the species listed most closely approximates the actual specimen taken. Hodges # Taxa Month(s) Collected Proportion TINEIDAE 372. Acrolophus plumifrontella (Clemens 1859) July <0.01 373 Acrolophus popeanella (Clemens 1859) July—August <0.01 OECOPHORIDAE 1014 + Antaeotricha leucillana (Zeller 1854) May—August <0.01 1046 Callima argenticinctella Clemens 1860 June—August <0.01 COLEOPHORIDAE 1387 Coleophora spissicornis (Haworth 1828) June—July <0.01 COSMOPTERIGIDAE 1615 Walshia sp. June, August 0.03 GELECHIIDAE 2289 Trichotaphe alacella Clemens 1862 July <0.01 2295 Trichotaphe flavocostella (Clemens 1860) June—July <0.01 PLUTELLIDAE 2366 Plutella xylostella (Linnaeus 1758) April <0.01 YPONOMEUTIDAE 2401 Atteva punctella (Cramer 1781) June—August <0.01 TORTRICIDAE 2927 Phaneta ochrocephala (Walsingham 1895) August <0.01 293i) Phaneta parmatana (Clemens 1860) August <0.01 3116 Eucosma dorsisignatana (Clemens 1860) August <0.01 3120 Eucosma derelecta Heinrich 1929 August <0.01 3202 Epiblema otiosana (Clemens 1860) August <0.01 8235 Proteoteras moffatiana Fernald 1905 June <0.01 3404 Dichrorampha simulana (Clemens 1860) June <0.01 3492 Cydia pomonella (Linnaeus 1758) June—August <0.01 3597 Argyrotaenia velutinana (Walker 1863) August <0.01 3635 Choristoneura rosaceana (Harris 1841) June, August <0.01 3684 Clepsis clemensiana (Fernald 1879) June <0.01 3693 Xenotemna pallorana (Robinson 1869) August <0.01 3701 Sparganothis pulcherrimana (Walsingham 1879) June <0.01 S752 Platynota flavedana Clemens 1860 August <0.01 ZY GAENIDAE 4624 Harrisina americana (Guerin 1824) June <0.01 LIMACODIDAE 4661 Packardia elegans (Packard 1864) June <0.01 4665 Lithacodes fasciola (Herrich-Schaffer 1865) June <0.01 4697 Euclea delphinii (Boisduval 1832) June <0.01 CRAMBIDAE 4743 Neocataclysta magnificalis (Hubner 1796) July <0.01 4755 Synclita obliteralis (Walker 1859) June—August <0.01 4895 Chalcoela iphitalis (Walker 1859) June, August <0.01 4944 Crocidophora serratissimalis Zeller 1872 August <0.01 4949 Ostrinia nubilalis (Hubner 1796) June, August <0.01 5071 Pyrausta acrionalis (Walker 1859) July—August <0.01 5079 Udea rubigalis (Guenee 1854) April—September 0.02 5142 Diacme elealis (Walker 1859) July <0.01 VOLUME 106, NUMBER 1 219 Table 1. Continued. ee ee Hodges # Taxa Month(s) Collected Proportion 5156 Nomophila nearctica Munroe 1973 June—September <0.01 Silp9 Desmia funeralis (Hubner 1796) June, August <0.01 5226 Palpita magniferalis (Walker 1861) May <().01 5280 Herpetogramma aeglealis (Walker 1859) August <0.01 5357 Crambus leachellus (Zincken 1818) September <0.01 5361 Crambus albellus Clemens 1860 June 0.01 5362 Crambus agitatellus Clemens 1860 June—August 0.12 5378 Crambus laqueatellus Clemens 1860 June <0.01 5/9 Crambus luteolellus Clemens 1860 July 0.03 5403 Agriphila vulgivagella (Clemens 1860) August-September <0.01 5413 Pediasia trisecta (Walker 1856) July—September <0.01 5420 Microcrambus elegans (Clemens 1860) July—August 0.06 5435 Fissicrambus mutabilis (Clemens 1860) August <0.01 5464 Urola nivalis (Drury 1773) June—July <0.01 5492 Eoreuma densella (Zeller 1881) July <0.01 5524 Hypsopygia costalis (Fabricius 1775) September <0.01 PYRALIDAE 5566 Arta statalis (Grote 1875) July <0.01 5579 Epipaschia zelleri (Gorte 1876) June <0.01 5799 Nephopterix basilaris Zeller 1872 August <0.01 5804 Nephopterix rubrisparsella (Ragonot 1887) June, August <0.01 5944 Homoeosoma deceptorium Heinrich 1956 July <0.01 5946 Phycitodes albatella (Ragonot 1887) June <0.01 6005 Moodna ostrinella (Clemens 1860) August <0.01 PTEROPHORIDAE 6204 Oidaematophorus elliottii (Fernald 1893) June <0.01 THYATIRIDAE 6235 Habrosyne scripta (Gossbeck 1840) July <0.01 GEOMETRIDAE 6270 Protitame virginalis (Hulst 1900) June <0.01 6322 Mellilla xanthometata (Walker 1862) August <0.01 6353 Semiothisa multilineata (Packard 1873) July <0.01 6362 Semiothisa continuata (Walker 1862) July <0.01 6419 Enconista dislocaria (Packard 1876) May—July <0.01 6590 Anavitrinela pampinaria (Guenee 1857) August <0.01 6640 Biston betularia (Linnaeus 1758) August <0.01 6654 Hypagyrtis unipunctata (Haworth 1809) June <0.01 6667 Lomographa vestaliata (Guenee 1857) June <0.01 6720 Lytrosis unitaria (Herrich-Schaffer 1854) June <0.01 6724 Euchlaena serrata (Drury 1770) June <0.01 6725 Euchlaena muzaria (Walker 1860) June <0.01 6726 Euchlaena obtusaria (Hubner 1807-13) August <0.01 6729 Euchlaena johnsonaria (Fitch 1869) June, August <0.01 6740 + Xanthotype sp. August <0.01 6753 + Pero sp. May—August <0.01 6822 Metarranthis duaria (Guenee 1857) June <0.01 6826 Metarranthis hypocharia (Herrich-Schaffer 1854) May—June <0.01 6941 Eusarca confusaria Hubner 1813 May—September 0.03 6964 Tetracis cachexiata Guenee 1857 June <0.01 6982 Prochoerodes transversata (Drury 1770) September <0.01 6987 Antepione thisoaria (Guenee 1857) May <0.01 7009 Nematocampa limbata (Haworth 1807) June <0.01 7053 Dichorda iridaria (Guenee 1857) May—June <0.01 a N 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Hodges # Taxa Month(s) Collected Proportion 7132 Pleuroprucha insulsaria (Guenee 1857) June, August <0.01 7146 Haematopis grataria (Fabricius 1798) September <0.01 7159 Scopula limboundata (Haworth 1809) June, August <0.01 7169 Scopula inductata (Guenee 1857) June—July <0.01 WSO ar Eulithis sp. June, July, September <0.01 7414 Orthonama obstipata (Fabricius 1794) April, June, July, September 0.02 7416 Orthonama centrostrigaria (Wollaston 1858) June—August 0.01 7445 Horisme intestinata (Guenee 1857) August <0.01 TATA + Eupithecia sp. April—September 0.05 LASIOCAMPIDAE 7670 Tolype velleda (Stoll 1791) September <0.01 SATURNIIDAE 7746 Automeris io (Fabricius 1775) June <0.01 SPHINGIDAE 7871 Deidamia inscripta (Harris 1839) April-May <0.01 7885 Darapsa myron (Cramer 1780) June <0.01 NOTODONTIDAE 7896 Clostera inclusa (Hubner 1829-31) May <0.01 7901 Clostera apicalis (Walker 1855) July <0.01 7902 Datana ministra (Drury 1773) June—July <0.01 7904 + Datana sp. June—July <0.01 7906 Datana contracta Walker 1855 June—August <0.01 7907 Datana integerrima Grote and Robinson 1860 July <0.01 7908 Datana perspicua Grote and Robinson 1865 July <0.01 7915 Nadata gibbosa (Smith 1797) June, August <0.01 7920 Peridea angulosa (Smith 1797) June <0.01 7929 Nerice bidentata Walker 1855 June <0.01 7930 Ellida canaplaga (Walker 1856) April, August <0.01 7931 Gluphisia septentrionis Walker 1855 August <0.01 7985 Heterocampa subrotata Harvey 1874 June—July <0.01 7994 Heterocampa guttivitta (Walker 1855) June—July <0.01 71995 Heterocampa biundata Walker 1855 May—June <0.01 7999 Lochmaeus bilineata (Packard 1864) August <0.01 8007 Schizura unicornis (Smith 1797) June, August <0.01 ARCTIIDAE 8045.1 Crambidia pallida Packard 1864 August <0.01 8089 Hypoprepia miniata (Kirby 1837) June, August 0.01 8121 Holomelina aurantiaca (Hubner 1827-31) August <0.01 8129 Pyrrharctia isabella (Smith 1797) June, August <0.01 8133 Spilosoma latipennis Stretch 1872 May—June <0.01 8137 Spilosoma virginica (Fabricius 1798) June—August <0.01 8140 Hyphantria cunea (Drury 1773) June <0.01 8146 Ecpantheria scribonia (Stoll 1790) June <0.01 8169 + Apantesis sp. June, August 0.01 8203 Halysidota tessellaris Smith 1797 June—July <0.01 8203 + Halysidota sp. July <0.01 8211 Lophocampa caryae Harris 1841 May—June <0.01 8230 Cycnia tenera Hubner 1818 June <0.01 8231 Cycnia oregonensis (Stretch 1873) June <0.01 8238 Euchaetes egle (Drury 1773) June—July <0.01 8262 Ctenucha virginica (Esper 1794) June <0.01 8267 Cisseps fulvicollis (Hubner 1818) June <0.01 On Te VOLUME 106, NUMBER 1 Table |. Continued. Hodges # Taxa LYMANTRIIDAE 8316 Orgyia leucostigma (Smith 1797) 8318 Lymantria dispar (Linnaeus 1758) NOCTUIDAE 8322 Idia americalis (Guenee 1854) 8323 Idia aemula Hubner 1813 8334 Idia lubricalis (Geyer 1832) 8347 Zanclognatha obscuripennis (Grote 1872) 8348 Zanclognatha pedipilalis (Guenee 1854) 8355 Chytolita morbidalis (Guenee 1854) 8360 Macrochilo orciferalis Walker 1859 8362 Phalaenostola metonalis (Walker 1859) 8363 Phalaenostola eumelusalis (Walker 1859) 8364 Phalaenostola larentioides Grote 1873 8368 Tetanolita floridana (Smith 1895) 8370 Bleptina caradrinalis Guenee 1854 8386 Renia adspergillus (Bosc 1800) 8393 Lascoria ambigualis Walker 1866 8397 Palthis angulalis (Hubner 1796) 8398 Palthis asopialis (Guenee 1854) 8465 Plathypena scabra (Fabricius 1798) 8479 Spargaloma sexpunctata Grote 1873 8499 Metalectra discalis (Grote 1876) 8534 Plusiodonta compressipalpis Guenee 1852 8692 Zale galbanata (Morrison 1876) 8738 + Caenurgina sp. 8769 Spiloloma lunilinea Guenee 1873 8874 Catocola minuta Edwards 1864 8898 Allagrapha aerea (Hubner 1802—03) 8907 Megalographa biloba (Stephens 1830) 8908 Autographa precationis (Guenee 1852) 8924 Anagrapha falcifera (Kirby 1837) 8973 Baileya australis (Grote 1881) 9047 Lithacodia muscosula (Guenee 1852) 9053 Pseudeustrotia carneola (Guenee 1852) 9055.1 Maliattha synochitis (Grote and Robinson 1868) 9057 Homophoberia apicosa (Haworth 1809) 9065 Leuconycta diphteroides (Guenee 1852) 9095 Tarachidia erastrioides (Guenee 1852) S27) Spragueia leo (Guenee 1852) 9199 Acronicta rubricoma Guenee 1852 9582 Apamea vulgaris (Grote and Robinson 1866) 9367 Apamea dubitans (Walker 1856) 9402 Oligia chlorositigma (Harvey 1876) 9457 + Amphipoea americana (Speyer 1875) 9473 Papaipema impecuniosa (Grote 1881) 9484 Papaipema baptisiae (Bird 1902) 9626 Trachea delicata (Grote 1874) 9639 Amphipyra tragopoginis (Clerck 1759) 9647 Athetis miranda (Grote 1873) 9650 Anorthodes tarda (Guenee 1852) 9663 Balsa tristrigella (Walker 1866) 9666 Spodoptera frugiperda (Smith 1797) 9688 Galgula partita Guenee 1852 sss Month(s) Collected July July—August June June July—August August May—June June June, August June, August June, September June—August September June—July June August June—July August August-September June July May—June June May—September July July September April July—August June May, July, August June—July May, June, August June August June June, August July August June August June July September September June September May, July, September August June August-September April, June, July Proportion <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 0.02 <0.01 0.02 <0.01 <0.01 <0.01 <0.01 0.01 <0.01 <0.01 <0.01 <0.01 0.02 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 0.02 DPD PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Hodges # Taxa Month(s) Collected Proportion 9720 Ogdoconta cinereola (Guenee 1852) June <0.01 9893 Lithophane hemina Grote 1874 April <0.01 SI)5)]) Sunira bicolorago (Guenee 1852) September <0.01 10021 Copivaleria grotei (Morrison 1874) April <0.01 10300 Spiramater grandis (Guenee 1852) June <0.01 10368 Lacinipolia meditata (Grote 1873) September <0.01 10397 Lacinipolia renigera (Stephens 1829) June—September 0.03 10405 Lacinipolia lorea (Guenee 1852) June <0.01 10438 Pseudaletia unipuncta (Haworth 1809) April—September 0.02 10444 Leucania phragmitidicola Guenee 1852 June, September <0.01 10445 Leucania linda Franclemont 1952 June <0.01 10446 + Leucania sp. June, August <0.01 10461 + Leucania sp. June, August <0.01 10462 Leucania pseudargyria Guenee 1852 June <0.01 10488 Orthosia garmani (Grote 1879) April <0.01 10501 Crocigrapha normani (Grote 1874) May <0.01 10524 Nephelodes minians Guenee 1852 September <0.01 10585 Orthodes crenulata (Butler 1890) August <0.01 10587 Orthodes cynica Guenee 1852 May—June 0.04 10627 Tricholota signata (Walker 1860) July <0.01 10648 Agrotis gladiaria Morrison 1874 September <0.01 10651 Agrotis venerabilis Walker 1857 September <0.01 10663 Agrotis ipsilon (Hufnagel 1766) June, July, September <0.01 10674 + Feltia sp. August-September <0.01 10676 Feltia herilis (Grote 1873) August-September 0.05 10803 Euxoa velleripennis (Grote 1874) September <0.01 10891 Ochropleura plecta (Linnaeus 1761) July—August <0.01 10942 + Xestia sp. June, September 0.01 10942.1 Xestia dolosa Franclemont 1980 May—June <0.01 10950 + Xestia sp. September <0.01 10955 Xestia badinodis (Grote 1874) September <0.01 10994 Cerastis tenebrifera (Walker 1865) April <0.01 10998 Choephora fungorum Grote and Robinson 1868 September <0.01 11006 Protolampra brunneicollis (Grote 1865) September <0.01 11029 Abagrotis alternata (Grote 1864) September <0.01 11068 Helicoverpa zea (Boddie 1850) August-September <0.01 11149 Schinia trifascia Hubner 1818 August <0.01 MIL Noctua pronuba Linnaeus 1758 June—August <0.01 on the North Coleman Tract from 1999— 2001; the North Coleman Tract is posi- tioned slightly west of the center of the is- land. The trap was placed on the edge of a glacial groove that forms a small escarp- ment approximately | m above the valley floor (Fig. 1). The black light was turned on in the late afternoon before dusk, and specimens were recovered the next morn- ing. The immediate area around the trap was cleared to remove woody vegetation during early summer 1999 to provide a prairie remnant, leaving a forested environ- ment surounding the trap location. The list of moths we obtained is indexed according to family and Hodges’ number (Hodges et al. 1988). Specimens of ques- tionable identification were given a Hodg- es’ number for the species most closely ap- proximating the specimen, and are identi- fied on the list with a “‘plus” symbol (+) next to the Hodges’ number. Although a more recent classification is available (Poole 1996), we used Hodges et al. (1988) VOLUME 106, NUMBER 1 leva Il a. ? (24 Y all Kelleys Island Marblehead Sandusky anacaee Location of Kelleys Island in Lake Erie, and position of trap (filled square) on the North Coleman Tract. Contour lines show elevation in feet above sea level, and depict northward slope of trapping area. and their numbering system to better place those species of questionable identification. Information on host plant use was taken from Holland (1903), Rings et al. (1992), and Rings and Downer (2001). All speci- mens are housed at The Cleveland Museum of Natural History, Department of Inverte- brate Zoology. To obtain an estimate of the actual num- ber of moth species on Kelleys Island, data on species accumulation over ten years from the six papers prepared by Teraguchi Table 2. Number of moth specimens, number of species, cumulative number of species, and number of new species captured for three year survey of Kelleys Island. No. No. of Cumulative No. New Year Specimens spp- spp. spp 1999 7} 105 105 — 2000 408 103 163 58 2001 709 124 216 52 a and Lublin (1999a—f) were combined, and the average rate of species accumulation from the six northeastern Ohio sites they sampled was determined. A_ logarithmic curve exhibited a good fit with their data (y = 113.73 Ln(x) + 342.2, R? = 0.99, where y is the cumulative number of species and x is the year of collection). We applied a logarithmic, non-linear regression to the three years of species accumulation data from this study to give a preliminary esti- mate of the number of species anticipated after 10 years of sampling on Kelleys Is- land. The Kelleys Island flora was surveyed periodically during 1975—2002. Plants were identified or categorized using Fernald (1950), Voss (1996), and Rhoads and Block (2000). Representatives of all plant species taken were mounted on herbaria sheets, and are housed at The Cleveland Museum of Natural History, Department of Botany. 224 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. The flora of Kelleys Island, with species found at the site of light trap setup on the Coleman Tract property. Coleman Taxa Common Name Tract sp. Thelypteridaceae Thelypteris palustris Schott marsh fern Taxaceae Taxus canadensis Marsh American yew Cupressaceae Juniperus virginiana L. eastern red-cedar x Potamogetonaceae Potamogeton pectinatus L. sago false pondweed Potamogeton pusillus var. pusillus L. small pondweed Potamogeton richardsonii (Benn.) Rydb. red-headed pondweed Alismataceae Alisma subcordatum Raf. American water plantain xX Alisma triviale Pursh northern water plantain Xx Hydrocharitaceae Elodea canadensis L. C. Rich. common waterweed Vallisneria americana Michx. American eel-grass Poaceae Agrostis stolonifera var. palustris (Huds.) Farw. spreading vent Andropogon gerardii var. gerardii Vitman beardgrass xX Bromus erectus Hudson bromegrass Dactylis glomerata L. orchard-grass Elymus virginicus var. virginicus L. Virginia wild rye x Festuca elatior L. fescue Glyceria striata (Lam.) A. Hitche. fowl mannagrass Panicum flexile (Gattinger) Scribn. wiry panic grass Panicum gattingeri Nash Gattinger’s panic grass Panicum implicatum Scribn. panic grass xX Panicum philadelphicum Bernh. Philadelphia panic grass Phragmites australis (Cav.) Trin. Ex Steud. phragmites Poa compressa L. Canada bluegrass Poa pratensis L. Kentucky blue grass xX Sphenopholis intermedia Rydb. slender wedgescale x Sporobolus ozarkanus Fern. dropseed Triplasis purpurea (Walt.) Chapman purple sand grass Cyperaceae Carex aggregata Mackenzie glomerata sedge Carex amphibola var. turgida Fern. narrow-leaf sedge x Carex aurea Nutt. golden-fruit sedge Carex blanda Dewey eastern woodland sedge xX Carex cephalophora Willd. oval-leaf sedge x Carex comosa Boott bearded sedge Carex crawei Dewey Crawe’s sedge Carex davisti Schwein. & Torr. Davis’ sedge x Carex eburnea Boott bristle-leaf sedge Carex flava L. yellow-green sedge Carex garberi Fern. elk sedge Carex granularis Muhl. Ex Willd. limestone-meadow sedge x Carex lupulina Willd. hop-like sedge Carex meadii Dewey Mead’s sedge VOLUME 106, NUMBER 1 Table 3. Continued. oo EEEeEeEeEeSFSFSSMMMmmsmsmmHhFeFeseFeseFeFeFFFFfsFsfsfsfsfsfsMséS Taxa Carex molesta McKenzie Carex muskingumensis Schwein. & Torr. Carex oligocarpa Willd. Carex pellita Muhl. Carex radiata (Wahlen.) Small Carex rosea Schkuhr ex Willd. Carex shortiana Dewey Carex sparganioides Willd. Carex stipata Muhl. Ex Willd. Carex umbellata Schkuhr ex Willd. Carex viridula Michx. Carex vulpinoidea Michx. Cladium mariscoides (Muhl.) Torr. Cyperus bipartitus Torrey Cyperus engelmannii Steud. Cyperus odoratus L. Eleocharis caribaea (Rottb.) Blake Eleocharis compressa Sulliv. Eleocharis elliptica Kunth Eleocharis erythropoda Steud. Fimbristylis autumnalis (L.) Roemer & Schultes Scirpus atrovirens Willd. Scirpus lineatus Michx. Scirpus pendulus Muhl. Scleria verticillata Muhl. Ex Willd. Pontederiaceae Heteranthera dubia (Jacq.) MacM. Juncaceae Juncus alpinus Vill. Juncus articulatus L. Juncus stuckeyi Reinking Juncus tenuis var. uniflorus (Farw.) Farw. Juncus torreyi Coville Liliaceae Allium canadense L. Allium cernuum Roth Allium tricoccum Ait. Erythronium albidum Nutt. Iridaceae Sisyrinchium albidum Raft. Sisyrinchium angustifolium P. Mill. Sisyrinchium montanum Greene Sisyrinchium mucronatum Michx. Orchidaceae Liparis loeselii (L.) L. C. Rich. Spiranthes cernua (L.) L. C. Rich. Spiranthes magnicamporum Sheviak Salicaceae Populus deltoides Bartr. Ex Marsh Salix amygdaloides Anderss. Salix bebbiana Sarg. Coleman Common Name Tract sp. troublesome sedge X sedge x Richwood’s sedge xX woolly sedge eastern star sedge rosy sedge Short’s sedge burr-reed sedge stalk-grain sedge parasol sedge little green sedge common fox sedge twig-rush shining flat sedge Engelmann’s umbrella sedge umbrella sedge capitate spike rush flat-stem spike-rush elliptic spike-rush spike-rush slender fimbry dark-green bulrush bulrush rufous bulrush low nut-rush m~ m X grass-leaf mud plantain northern green rush joint-leaf rush Stuckey’s rush poverty rush Torrey’s rush xX meadow garlic x nodding onion x ramp small white fawn-lily white blue-eyed grass blue-eyed grass Xx strict blue-eyed grass needle tip blue-eyed grass yellow wide-lip orchid white nodding ladies’-tresses great plains ladies’-tresses eastern cottonwood xX peach-leaf willow gray willow N Table 3. Continued. 26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nl Taxa Coleman Common Name Tract sp. i Salix humilis Marsh Juglandaceae Carya ovata (P. Mill.) K. Koch Carya glabra (Miller) Sweet. Juglans nigra L. Betulaceae Betula pendula Roth Betula sandbergii Britt. Ostrya virginiana (P. Mill.) K. Koch Fagaceae Quercus macrocarpa Michx. Ulmaceae Celtis occidentalis L. Ulmus americana L. Ulmus rubra Muhl. Ulmus thomasii Sarg. Moraceae Morus alba L. Polygonaceae Polygonum lapathifolium L. Rumex crispus L. Chenopodiaceae Chenopodium standleyanum Aellen Molluginaceae Mollugo verticillata L. Portulacaceae Claytonia virginica L. Caryophyllaceae Cerastium arvense L. Stellaria media (L.) Vill. Ranunculaceae Aquilegia canadensis L. Clematis orientalis L. Ranunculus micranthus (Gray) Nutt. Ex Torr. & Gray Ranunculus recurvatus Poir. Fumariaceae Dicentra cucullaria (L.) Bernh. Brassicaceae Alliaria petiolata (Bieb.) Cavara & Grande Arabidopsis thaliana (L.) Heynh. Arabis glabra (L.) Bernh. Arabis hirsuta var. adpressipilis (M. Hopkins) Rollins Cardamine concatenata (Michx.) O. E. Schulz Cardamine douglassii (TYorr.) Britt. Cardamine hirsuta L. Diplotaxis muralis (L.) DC. Hesperis matronalis L. Rorippa palustris ssp. hispida (L.) Bess. (Desv.) Jonsell prairie willow shag-bark hickory x pignut hickory black walnut European birch hybrid (papyrifera * pumila) eastern hop-hornbeam burr oak common hackberry American elm slippery elm rock elm x * white mulberry x dock-leaf smartweed curlydock Standley’s goosefoot carpet weed Virginia spring beauty field mouse-ear chickweed common chickweed red columbine Oriental virgin’s bower rock buttercup Xx blisterwort Dutchman’s breeches garlic-mustard thalecress tower-mustard hairy rockcress cut-leaf toothwort limestone bittercress hairy bittercress wall or sand rocket dame’s rocket bog yellowcress VOLUME 106, NUMBER 1 Table 3. Continued. Sn ———————————————————————————————————————————— Taxa Coleman Common Name Tract sp. ee Saxifragaceae Heuchera americana L. Grossulariaceae Ribes americanum P. Mill. Rosaceae Amelanchier arborea (Michx. F.) Fern. Amelanchier spicata (Lam.) K. Koch Fragaria virginiana Duchesne. Geum canadense Jacq. Geum vernum (Raf.) Torr. & Gray Malus pumila Mill. Physocarpus opulifolius (L.) Maxim. Potentilla fruticosa L. Potentilla paradoxa Nutt. Ex T. & G. Potentilla simplex Michx. Prunus mahaleb L. Prunus virginiana L. Rosa blanda Ait. Rosa carolina L. Rosa multiflora Thunb. Rubus allegheniensis T.C. Porter Rubus pensilvanicus Poiret. Fabaceae Desmodium canescens (L.) DC. Gleditsia tricanthos L. Gymnocladus dioica (L.) K. Koch Medicago lupulina L. Oxalidaceae Oxalis stricta L. Geraniaceae Geranium carolinianum L. Geranium pusillum L. Rutaceae Ptelea trifoliata L. Zanthoxylum americanum P. Mill. Euphorbiaceae Acalypha rhomboidea Raf. Euphorbia maculata L. Euphorbia polygonifolia L. Anacardiaceae Rhus aromatica Ait. Rhus aromatica var. arenaria (Greene) Fern. Rhus glabra L. Rhus radicans L. Celastraceae Celastrus scandens L. Staphyleaceae Staphylea trifolia L. American alumroot wild black currant xX downy service-berry service-berry thick-leaved wild strawberry white avens spring avens x apple Atlantic ninebark cinquefoil bushy cinquefoil oldfield cinquefoil perfumed cherry choke cherry smooth rose Carolina rose Xx multiflora rose common blackberry Pennsylvania blackberry hoary tick-trefoil honey-locust Kentucky coffee tree black medick common yellow wood-sorrel Carolina crane’ s-bill xX small-flowered crane’s-bill common hoptree XxX toothache tree Xx common three-seed-mercury eyebane seaside spurge fragrant sumac Xx fragrant sumac smooth sumac poison ivy X American bittersweet American bladdernut 228 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Continued. Coleman Taxa Common Name Tract sp. Aceraceae Acer platanoides L. Norway maple Acer saccharum Marsh sugar maple Hippocastanaceae Aesculus glabra var. arguta (Buckl.) B.L. Robins. Ohio buckeye Vitaceae Parthenocissus inserta (Kern.) Fritsch Virginia creeper Vitis riparia Michx. river-bank grape Tiliaceae Tilia americana L. American basswood x Clusiaceae Hypericum perforatum L. common St. John’s-wort Violaceae Viola affinis LeConte sand violet Viola pubescens Ait. downy yellow violet Elaeagnaceae Elaeagnus umbellata Thunb. autumn olive Lythraceae Lythrum alatum Pursh wing-angle loosestrife Onagraceae Oenothera parviflora L. northern evening-primrose Haloragidaceae Proserpinaca palustris L. mermaid-weed Apiaceae Daucus carota L. queen anne’ s-lace Thaspium barbinode (Michx.) Nutt. hairy-jointed meadow parsnip Cornaceae Cornus amomum spp. amomum P. Mill. silky dogwood Cornus amomum ssp. obliqua (Raf.) J.S. Wilson silky dogwood Cornus drummondii C. A. Mey. rough-leaf dogwood x Primulaceae Lysimachia quadriflora Sims four-flower yellow-loosestrife Oleaceae Fraxinus americana L. white ash Xx Fraxinus pennsylvanica Marsh red ash x Fraxinus quadrangulata Michx. blue ash xX Apocynaceae Apocynum cannabinum L. Indian-hemp xX Vinca minor L. Asclepiadaceae Asclepias incarnata L. Asclepias syriaca L. Asclepias tuberosa L. Asclepias tuberosa ssp. tuberosa L. Asclepias verticillata L. Asclepias viridiflora Raf. lessor periwinkle swamp milkweed common milkweed x butterfly weed butterfly weed whorled milkweed green milkweed VOLUME 106, NUMBER 1 Table 3. Continued. ————————— EEE Taxa Polemoniaceae Phlox divaricata L. Hydrophyllaceae Phacelia purshii Buckl. Boraginaceae Lappula squarrosa (Retz.) Dumort. Verbenaceae Verbena simplex Lehm. Lamiaceae Clinopodium vulgare L. Lamium purpureum L. Lycopus americanus Muhl. Ex Bart. Lycopus europaeus L. Lycopus sherardii Steele Lycopus uniflorus Michx. Mentha piperita L. Nepeta cataria L. Scutellaria parvula Michx. Trichostema brachiatum L. Scrophulariaceae Chaenorrhinum minus (L.) Large Leucospora multifida (Michx.) Nutt. Penstemon hirsutus (L.) Willd. Scrophularia marilandica L. Bignoniaceae Campsis radicans (L.) Seem. Ex Bureau Plantaginaceae Plantago rugelii Decne. Rubiaceae Galium mollugo L. Hedyotis nigricans (Lam.) Fosberg Caprifoliaceae Lonicera morrowii A. Gray Campanulaceae Campanula americana L. Lobelia kalmii L. Asteraceae Aster ericoides L. Aster novae-angliae L. Aster pilosus var. demotus Blake Aster pilosus var. pilosus Willd. Aster pilosus var. pringlei (Gray) Blake Aster shortii Lindl. Aster simplex Willd. Bidens cernua L. Bidens bipinnata L. Cirsium arvense L. Scop. Cirsium discolor (Muhl. Ex Willd.) Spreng. Coleman Common Name Tract sp. wild blue phlox x Miami-mist bristly sheepburr narrow-leaf vervain wild basil red henbit cut-leaf water-horehound gypsywort water-horehound northern water-horehound water mint catnip small skullcap false penny royal dwart-snapdragon narrow-leaf paleseed hairy beard tongue carpenter's square trumpet-creeper American plantain false baby’s breath xX diamond flowers morrow’s honeysuckle American bellflower kalm’s lobelia many-flowered aster New England American aster heath aster heath aster Pringle’s American aster Short’s American aster panicled aster X nodding bar marigold Spanish-needles Canada thistle field thistle ee —————————— 230 Table 3. Continued. Taxa Chrysanthemum leucanthemum L. Eclipta prostrata (L.) L. Eupatorium altissimum L. Eupatorium perfoliatum L. Euthamia graminifolia (L.) Nutt. Ex Cass. Hieracium lactucella Wallr. Lactuca floridana (L.) Gaertn. Liatris spicata (L.) Willd. Polymnia canadensis L. Senecio pauperculus Michx. Solidago canadensis var. salebrosa (Piper) M. E. Jones Solidago gigantea Aiton. Solidago nemoralis Ait. Taraxacum officinale Weber RESULTS AND DISCUSSION We collected 1,690 moths representing 216 species from 20 families (Table 1). Noctuidae was the most species-rich family collected with 89 species taken, followed by Geometridae (33), Crambidae (24), Arc- tiidae (17), Notodontidae (17), and Tortri- cidae (14). Combined, those six families represent 89.8% of the total species taken. All other families were represented by 1—7 species. Twenty-two (10.2%) species each repre- sented =1% of the total catch. Crambus agitatellus (Crambidae) was the most nu- merous, representing 12.2% of the total catch (Table |). This species demonstrated a moderately long flight period from June to August on Kelleys Island, and the larvae are known to feed on grasses. Three species exhibited a long flight pe- riod spanning April through September: Udea_ rubigalis (Crambidae), Eupethecia sp. (Geometridae), and Pseudaletia uni- punctata (Noctuidae). Of these, only P. un- ipunctata was taken frequently, and repre- sented approximately 2% of all moth spec- imens obtained. All three species are known to be highly polyphagous on a variety of plants, including woody and herbaceous forms. Ten species were limited to an early flight period spanning only April and/or PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Coleman Common Name Tract sp. ox-eye daisy yerba-de-tajo tall thoroughwort boneset flat-top goldentop hawkweed woodland lettuce blazing-star white flower leafcup x ragwort x Canadian goldenrod smooth goldenrod gray goldenrod common dandelion May: Plutella xylostella (Plutellidae), Pal- pita magniferalis (Crambidae), Antepione thisoaria (Geometridae), Deidama inscripta (Sphingidae), Clostera inclusa (Notodonti- dae), and the noctuids Lithophane hemina, Copivaleria grotei, Orthosia garmani, Cro- cigrapha normani, and Cerastis tenebri- fera. All ten species are restricted to feeding on trees or cultivated plants as larvae. No early flying species were taken in any abun- dance. Conversely, 20 moth species were restricted to a late flight period (September only) on Kelleys Island: Crambus leachel- lus (Crambidae), Hypsopygia costalis (Crambidae), Prochoerodes transversata (Geometridae), Haematopsis grataria (Geometridae), Tolype velleda (Lasiocam- pidae), and the noctuids Tetanolita flori- dana, Allagrapha aerea, Papaipema impe- cuniosa, P. baptisiae, Amphipyra tragopo- ginis, Sunira bicolorago, Lacinipolia med- itata, Nephelodes minians, Agrotis gladiaria, A. venerabilis, Xestia sp., X. bad- inodis, Choephora fungorum, Protolampra brunneicollis, and Abagrotis alternata. No late flight species represented 21% of the total catch. Unlike the species that are re- stricted to an early flight period, the 20 late flight species are all highly polyphagous on woody and herbaceous vegetation as larvae. Yearly species accumulation data show VOLUME 106, NUMBER 1 that 111 species were collected during the second and third year of trapping that were not collected during the first year. The third year of collecting was the most species-rich with 124 species (Table 2). Applying a log- arithmic equation to the species accumula- tion data provided a good fit (R* = 0.99; y = 99.2 Ln(x) + 102.1), and gives an esti- mate that 330.5 moth species are anticipat- ed after ten years of collecting. This sug- gests that three years of effort have ob- tained 65% of the moth species on the is- land. Two-hundred-thirty-eight plant species from 65 families were found on the island, and are listed in Table 3. The Cyperaceae was the most species rich family with 39 species, and Carex spp. alone accounted for 26 species. Forty-nine species (23.1%) were collected from the North Coleman Tract. Comparing the calculated estimate of the number of moth species expected after ten years of collecting indicates that there are 1.39 moth species per plant species on Kel- leys Island. This implies that multiple insect herbivores put pressure on the Kelleys Is- land flora. Competitive interactions or niche packing may be limiting the number of moth species that the island can support. This is the only study we are aware of that can provide a predictive ratio (approximate- ly 1.4 lepidopteran herbivore species per plant species) for a herbivore community and its food sources. This study shows that, even when a rel- atively small area of terrestrial habitat is sampled (i.e., no confounding data obtained by moths flying in from areas outside of our sampling area), moths represent a species- rich community. Rings et al. (1987) at- tempted to associate the moth species they accumulated during a survey conducted in northeastern Ohio with their natural host plants. They were unsuccessful and deter- mined that this could not be done because the complete host range for most Lepidop- tera is unknown, and that moth species can be captured at great distances from their host plants using light traps. Although we i) eS) conducted no rearings, the concurrent pre- sentation of moth and plant species data from an insular environment may help other lepidopterists determine the host plant range of the moth species found during this study. Three years of data collection provided approximately two-thirds of the moth spe- cies anticipated after ten years of effort. Surveys of the entomofauna of a given area are hampered by rare taxa that may not be captured without long-term efforts. These results underscore the need for long-term survey work when dealing with Lepidop- tera, or any other speciose insect taxa. ACKNOWLEDGMENTS We thank the Ohio Biological Survey for funding this study. David J. Horn (Ohio State University) helped the senior author during the early phase of this study, and Bill Coleman kindly assisted with monthly field collections. Ruth Chase and Tom Puc- ci (CMNH) reviewed a draft of the paper. LITERATURE CITED Fernald, M. L. 1950. Gray’s Manual of Botany, 8th ed. The American Book Company, New York. Hodges, R. W., T. Dominick, D. R. Davis, D. C. Fer- guson, J. G. Franclemont, E. G. Munroe, and J. A. Powell, eds. 1988. Checklist of the Lepidoptera of America north of Mexico. E. W. Classey Ltd., London and Wedge Entomological Foundation, Washington D.C., 284 pp. Holland, W. J. 1903. The Moth Book. Reprinted 1968 with updating by A. E. Brower. Doubleday, Page, and Co., New York. Nault, B. A., R. W. Rings, and D. J. Horn. 1989. Lep- idoptera recorded from the islands of western Lake Erie, with a brief account of geology and flora. The Great Lakes Entomologist 22: 111—119. Poole, R. W. 1996. Nomina Insecta Nearctica, A Checklist of the Insects of North America, Vol. 3: Diptera, Lepidoptera, Siphonaptera. Entomologi- cal Information Services, Maryland, 1143 pp. Price, P. W. 1997. Insect Ecology. J. Wiley, New York, 874 pp. Rhoads, A. E and T. A. Block. 2000. The Plants of Pennsylvania: An Illustrated Guide. University of Pennsylvania Press, Philadelphia. Rings, R. W. and R. A. Downer. 2001. The Lepidop- tera of Wayne County, Ohio. Ohio State Univer- sity Research Bulletin, no. 1192, 164 pp. 232 Rings, R. W. and E. H. Metzler. 1989. A preliminary checklist of the Lepidoptera of Mohican State For- est and Mohican State Park, Ashland County, Ohio. Ohio Journal of Science 89: 78-88. Rings, R. W., E. H. Metzler, FE J. Arnold, and D. H. Harris. 1992. The owlet moths of Ohio. Bulletin of the Ohio Biological Survey, New Series 9: 1— 184. Rings, R. W., R. M. Ritter, and R. W. Hawes. 1987. A nine-year study of the Lepidoptera of The Wil- derness Center, Stark County, Ohio. Ohio Journal of Science 87: 55-61. Summerville, K. S., J. J. Jacquot, and R. E Stander. 1999. A preliminary checklist of the moths of Butler County, Ohio. Ohio Journal of Science 99: 66-76. Teraguchi, S. E. and K. J. Lublin. 1999a. Checklist of the moths of Pallister State Nature Preserve, Ash- tabula County, Ohio (1988-1992) with analyses of abundance. Kirtlandia 51: 3-18. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 1999b. Checklist of the moths of Lake Car- dinal Swamp, Ashtabula County, Ohio (1988— 1992) with analyses of abundance. Kirtlandia 51: 19-34. 1999c. Checklist of the moths of Groves Woods Preserve, Trumbull County, Ohio (1988— 1992) with analyses of abundance. Kirtlandia 51: 35-50. . 1999d. Checklist of the moths of Grand River Terraces Preserve, Ashtabula County, Ohio (1988-1992) with analyses of abundance. Kirtlan- dia 51: 51—66. . 1999e. Checklist of the moths of Resources Center, Hidden Valley Metropark, Lake County, Ohio (1988-1992) with analyses of abundance. Kirtlandia 51: 67—82. . 1999f. Checklist of the moths of Walden II Nature Preserve, Lake County, Ohio (1988-1992) with analyses of abundance. Kirtlandia 51: 83-98. Voss, E. G. 1996. Michigan Flora. University of Mich- igan Press, Ann Arbor. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 233-234 NOTE Ochlerotatus japonicus japonicus (Theobald) (Diptera: Culicidae), a New Invasive Mosquito for Georgia and South Carolina Exotic mosquitoes pose a threat to public health and the environment. Some recently introduced species such as Aedes albopictus (Skuse) are serious pests and transmit path- ogens to humans and domestic animals. Ochlerotatus japonicus japonicus (Theo- bald) is the most recently recognized spe- cies of exotic mosquito to become estab- lished in the continental United States (Pey- ton et al. 1999, Darsie 2002). Adult mos- quitoes were initially reported from New York and New Jersey (Peyton et al. 1999), Larvae of Oc. japonicus were later found in natural and artificial containers such as treeholes and used tires (Andreadis et al. 2001). Scott et al. (2001) reported that Oc. Japonicus was the most commonly collect- ed rockpool-dwelling mosquito in the Del- aware Water Gap National Recreation Area. Scott et al. (2001) and Andreadis et al. (2001) noted that this mosquito also inhab- its rockpools in Japan. Since the discovery of wild Oc. japonicus japonicus popula- tions in New Jersey and New York, it rap- idly expanded its range to Connecticut (An- dreadis et al. 2001), Pennsylvania, Ohio, Maryland (Fonseca et al. 2001), and Vir- ginia (Harrison et al. 2002). The public health implications of this exotic mosquito are unknown, but it can transmit Japanese encephalitis virus to mice in laboratory ex- periments (Peyton et al. 1999). We report the discovery of populations of Oc. japon- icus Japonicus in Georgia and South Caro- lina. We collected 13 mosquito larvae and 4 mosquito pupae from a leaf-lined rockpool adjacent to the Tallulah River, Southern Nantahalla Wilderness, Rabun County, Georgia, 34.9402°N, 83.5454°W, elevation 657 m, on 23 July 2003. In the laboratory, four adult mosquitoes emerged from the pu- pae within 24 hours and were identified as Oc. japonicus japonicus with the keys of Darsie (2002) and Darsie and Ward (1981). Ten of the larvae were Oc. japonicus ja- ponicus and three were a native rockpool mosquito Ochlerotatus atropalpus (Coquil- lett). We sampled similar habitats in South Carolina and collected 28 Oc. japonicus ja- ponicus and 2 Oc. atropalpus larvae from four rockpools along the Middle Saluda River, Jones Gap State Park, Greenville County, South~ Carolinas 35,1252 -N. 82.5733°W, elevation 420 m, on 26 and 30 July 2003. The larvae collected on 30 July were infected with a symbiotic trichomy- cete fungus Smittium sp. that develops in their hindguts. No other symbiotes were noted. Our collections suggest that Oc. ja- ponicus japonicus has invaded the southern Appalachians since Darsie (2002) reported it from the Northeast and central Atlantic States. Our collection in Georgia was less than 3 km from the North Carolina border, indicating that this mosquito probably oc- curs in North Carolina. There are no major urban centers near either collection site and these mosquitoes probably colonized both sites without human-assisted transportation. Our collections are significant for mosquito control programs because Oc. japonicus will bite humans. This mosquito could be involved in the transmission of economi- cally important pathogens such as West Nile encephalitis virus or Dirofilaria im- mitis (Leidy). Voucher specimens of larval and adult Oc. japonicus japonicus including larvae preserved in 95% ethanol for DNA analysis were deposited in the Clemson University Arthropod Collection. We thank P. H. Adler for reviewing a draft of this manuscript. 234 LITERATURE CITED Andreadis, T. G., J. E Anderson, L. E. Munstermann, R. J. Wolfe, and D. A. Florin. 2001. Discovery, distribution, and abundance of the newly intro- duced mosquito Ochlerotatus japonicus (Diptera: Culicidae) in Connecticut, USA. Journal of Med- ical Entomology 38: 774-779. Darsie, R. F, Jr. 2002. Revision of Darsie and Ward (1981) to include Ochlerotatus japonicus Theo- bald and a checklist of species referred to the ge- nus Ochlerotatus in the Nearctic region. Journal of the American Mosquito Control Association 18: 237-240. Darsie, R. FE, Jr. and R. A. Ward. 1981. Identification and geographical distribution of the mosquitoes of North America, north of Mexico. Mosquito Sys- tematics (Supplement) 1: 1—313. Fonseca, D. M., S. Campbell, W. J. Crans, M. Mogi, I. Miyagi, T. Toma, M. Bullians, T. G. Andreadis, R. L. Berry, B. Pagac, M. R. Sardelis, and R. C. Wilkerson. 2001. Aedes (Finlaya) japonicus (Dip- tera: Culicidae), a newly recognized mosquito in the United States: analyses of genetic variation in the United States and putative source populations. Journal of Medical Entomology 38: 135-146. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Harrison, B. A., P. B. Whitt, S. E. Cope, G. R. Payne, S. E. Rankin, L. J. Bohn, E M. Stell, and C. J. Neely. 2002. Mosquitoes (Diptera: Culicidae) col- lected near the Great Dismal Swamp: new state records, notes on certain species, and a revised checklist for Virginia. Proceedings of the Ento- mological Society of Washington 104: 655—662. Peyton, E. L., S. R. Campbell, T; M. Candeletti, M. Romanowski, and W. J. Crans. 1999. Aedes (Fin- laya) japonicus japonicus (Theobald), a new in- troduction into the United States. Journal of the American Mosquito Control Association 15: 238— 241. Scott, J. J.. E L. Carle, and W. J. Crans. 2001. Och- lerotatus japonicus collected from natural rock- pools in New Jersey. Journal of the American Mosquito Control Association 17: 91—92. W. K. Reeves and J. A. Korecki, De- partment of Entomology, Soils, and Plant Sciences, 114 Long Hall, Clemson Univer- sity, Clemson, SC 29634, U.S.A. (e-mail: wreeves @clemson.edu). PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 235-238 NOTE Overwintering Cave Mosquitoes (Diptera: Culicidae) of the Arkansas and Missouri Ozarks Some 60 species of mosquitoes occur in Arkansas and Missouri (Darsie and Ward 1981, Crockett 2002). We have long known that mosquitoes occur in Ozark caves, but few species records have been available in the literature. As a result of extensive col- lecting effort in Missouri caves by the Mis- souri Department of Conservation and in Arkansas caves by the Subterranean Bio- diversity Project at the University of Ar- kansas we now know of four species that occur in Ozark caves. The specimens recorded here are all fe- males found in caves in late fall, winter, and early spring. Few mosquitoes were ob- served in caves during the summer months despite an equal degree of collecting effort during those months. Mated females appar- ently enter caves to pass the winter. Male mosquitoes are rarely collected in caves. Ives (1938) found only 8 males among 3102 specimens collected in, Tennessee caves. They were all found in the early part of the hibernation period and may not have lived far into the winter. Anopheles punctipennis (Say).—AR- KANSAS: 1 2, Newton County, Tom Wat- son’s Bear Cave, 26 January 2002. MIS- SOURI: | 2, Boone County, Hunters Cave, 1 November 1998. 2 2, Crawford County, Campsite Cave, 18 December 1998. 5 9, Maries County, Indian Ford Cave, 4 Janu- ary 2002. 2 2, Maries County, Indian Ford Crevice Cave, 4 January 2003. 2 2 Mc- Donald County, MoDot Cave, 21 Novem- ber 2002. Anopheles punctipennis adults are noc- turnal, resting during the daytime in hollow trees, under rock overhangs, and in similar dark, moist shelters (Irby and Apperson 1992). Females take blood meals from mammals and birds. They rarely enter homes to feed. Females winter in buildings, cellars, hollow trees, and other well-pro- tected shelters (Carpenter and LaCasse 1955, Horsfall 1955). In caves, they tend to congregate in the twilight zone. Movement into and out of caves is apparently governed by outdoor temperature (Ives 1938, Hess and Crowell 1949). Adult females emerge from overwintering quarters as early as February, take blood meals, and lay eggs (Breeland et al. 1961). This species was previously recorded from Canadian caves by Peck (1988); Massachusetts subterra- nean habitats by Berg and Lang (1948): a New York cave by Lawlor (1935); Tennes- see caves by Ives (1938); Kentucky caves by Barr (1967); Illinois caves by Peck and Lewis (1977); and Bat Cave, Crawford County, Missouri, by Craig (1977). Culex erraticus (Dyar and Knab).—AR- KANSAS: 2 2, Marion County, Brown Cave, 16 November 2001. 1 2, Marion County, Reed Cave, 9 March 2002. 1 2, Marion County, Reed Cave, 15 November 2001. 1 2, Newton County, Big Hole, 10 November 2001. 4 @, Newton County, Saltpeter Cave, 17 March 2002. 2 2, New- ton County, Tweet’s Cave, 26 October 2001. 4 2, Stone County, Gustafson Cave, 6 October 2002. 2 2, Stone County, Hidden Spring Cave, 5 October 2002. MISSOURI: 4 2, Benton County, Estes Cave, 8 Novem- ber 2002. 2 2, Boone County, Hunters Cave, 1 November 1998. 1 2, Boone Coun- ty, Hunters Cave, 16 October 2002. 2 9, Boone County, Rocheport Cave, 6 Septem- ber 2002. 2 2, Camden County, Adkins Cave, 8 November 2002. 3 2, Camden County, Moles Cave, 27 January 2003. 1 2, McDonald County, McDot Cave, 21 No- vember 2002. 4 2°, Shannon County, Bank- er Cave, 13 December 1998. 3° 2, Shannon 236 County, Forester Cave, 17 October 1998. 2 ?, Shannon County, Knee Suck Cave, 20 January 1998. 4 2, Wright County, Little Smittle Cave, 14 November 2002. 2 9, Wright County, Lowell Cave, 13 November 2002. At dusk, C. erraticus females move from sheltered marsh and swamp forest sites to nearby grasslands (Snow 1955), where they take blood meals opportunistically from mammals, birds, reptiles and amphibians (Robertson et al. 1993, Williams and Meisch 1981). During the day, they rest on vegetation and in natural and man-made sheltered areas (Irby and Apperson 1992). This is the most common species found wintering in Ozark caves during the course of this study. It was previously reported from Missouri caves by Sutton (1993). Culex pipiens Linnaeus or C. quinque- fasciatus Say— ARKANSAS: 1 2, Benton County, Marshall Caves, 18 December 1999. MISSOURI: 2 2, Camden County, Adkins Cave, 8 November 2002. 2 2, Mar- ies County, Indian Ford Cave, 4 January 2002. 1 2, Polk County, Mcdermic Cave, 15 March 2001. 5 2, Wright County, Little Smittle Cave, 15 December 1999. Only females of this species were seen. It is not possible to reliably determine if the specimens are C. pipiens or C. quinquefas- ciatus. The northern and southern house mosquitoes are commonly found in homes, and they bite humans at night. Adults rest during the daytime on vegetation and in natural and man-made sheltered areas (Irby and Apperson 1992). Culex pipiens com- prised 89% of the 1562 mosquitoes found wintering in a small natural cave in the city of Montpellier, southern France, between October 1997 and the following March. Only four of the specimens were males (Gazave et al. 2001). Culex pipiens was previously recorded from Massachusetts subterranean habitats by Berg and Lang (1948), a New York cave by Lawlor (1935), Tennessee caves by Ives (1938), and Illinois caves by Peck and Lewis (1977). It was found in large numbers in a Minnesota PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mushroom cave in early March (Owen 1937): Uranotaenia sapphirina (Osten Sack- en).—MISSOURI: 1 @, Boone County, Hunters Cave, | November 1998. 1 2, Camden County, Moles Cave, 27 January 2003. 5 ¢, Oregon County, Long Point Cave, 1 April 1998. 1 2, Wright County, Little Smittle Cave, 14 November 2002. Uranotaenia sapphirina adults are most active in early evening. They rest during the day in dark cavities and other protected shelters (Irby and Apperson 1992). Peterson and Smith (1945) reported this species overwintering in considerable numbers in hollow trees with Anopheles quadrimacu- latus Say and A. punctipennis in Mississip- pi. It was collected from a cave in eastern New York State in February, along with A. punctipennis and C. pipiens (Lawlor 1935). The cave-inhabiting mosquito species are not restricted to overwintering in caves. They will choose any convenient dark, moist, sheltered area that provides protec- tion from freezing temperatures, such as hollow trees, mammal burrows, rock piles, wells, mine shafts, culverts, cellars, sheds, stables, and similar structures (Shemanchuk 1965, Zukel 1949). Most of the Ozark mosquitoes that have not been found in caves winter as eggs or larvae. Aside from the four species record- ed here, local species that are thought to overwinter as fertilized adult females in- clude A. guadrimaculatus, Culex peccator Dyar and Knab, Culex restuans Theobald, Culex salinarius Coquillett, Culex tarsalis Coquillett, Culex territans Walker, and Cu- liseta inornata (Williston). We might expect to find these overwintering in Ozark caves. Among them, A. quadrimaculatus (Hess and Crowell 1949, Ives 1938, Peck 1988) and C. restuans (Peck 1988) have been found overwintering in caves elsewhere. Culex territans (misidentified as C. apical- is) has been found overwintering in subter- ranean basement structures (Berg and Lang 1948). Adult female Culex tarsalis have been found wintering in abandoned mine VOLUME 106, NUMBER | tunnels in Colorado (Mitchell 1979, Black- more and Dow 1962) and Nevada (Chap- man 1961) and in rodent burrows in Cali- fornia (Mortenson 1953). Culex tarsalis and C. inornata have been found wintering in subterranean burrows of large mammals in Canada (Shemanchuk 1965). Our Aedes, Ochlerotatus and Psorophora species are thought to overwinter in the egg stage. Adult Aedes canadensis (Theobald), A. ci- nereus Meigen, A. vexans (Meigen), and Ochlerotatus sticticus (Meigen) have been found in Canadian caves in late spring and summer, but not in winter (Peck 1988). The Missouri specimens recorded here are the property of the Missouri Department of Conservation, Jefferson City. They will be deposited in the Enns Entomology Mu- seum, University of Missouri, Columbia. The Arkansas specimens are deposited in the University of Arkansas Arthropod Mu- seum, Fayetteville. LITERATURE CITED Barr, T. C. 1967. Ecological studies in the mammoth Cave system of Kentucky. I. The biota. Interna- tional Journal of Speleology 3: 147-204. Berg, M. and S. Lang. 1948. Observations of hiber- nating mosquitoes in Massachusetts. Mosquito News 8: 70-71. Blackmore, J. S. and R. P. Dow. 1962. Nulliparity in summer and fall populations of Culex tarsalis Coq. Mosquito News 22: 291-294. Breeland, S. G., W. E. Snow, and E. Pickard. 1961. Mosquitoes of the Tennessee Valley. Journal of the Tennessee Academy of Science 36(4): 249— ZIG) Carpenter, S. J. and W. J. LaCasse. 1955. Mosquitoes of North America (north of Mexico). University of California Press, Berkeley, 360 pp. Chapman, H. C. 1961. Abandoned mines as overwin- tering sites for mosquitoes, especially Culex tar- salis Coq. in Nevada. Mosquito News 21: 324— 3271. Craig, J. L. 1977. Invertebrate faunas of caves to be inundated by the Meramec Park Lake in eastern Missouri. NSS (National Speleological Society) Bulletin 39(3): 80-89. Crockett, R. J. 2002. Arbovirus surveillance and con- trol of Arkansas mosquitoes. Master’s thesis, Uni- versity of Arkansas (Fayetteville). Darsie, R. F, Jr. and R. A. Ward. 1981. Identification and geographical distribution of the mosquitoes of North America, north of Mexico. American Mos- 235 quito Control Association, Fresno, California, 313 Pp. Gazave, E., C. Chevillon, T. Lenormand, M. Maruine, and M. Raymond. 2001. Dissecting the cost of insecticide resistance genes during the overwin- tering period of the mosquito Culex pipiens. He- redity 87: 441—448. Hess, A. D. and R. L. Crowell. 1949. Seasonal history of Anopheles quadrimaculatus in the Tennessee Valley. Journal of the National Malaria Society 8: 159-170. Horsfall, W. R. 1955. Mosquitoes: their bionomics and relation to disease. The Ronald Press Company, New York, 723 pp. Irby, W. S. and C. S. Apperson. 1992. Spatial and tem- poral distribution of resting female mosquitoes (Diptera: Culicidae) in the coastal plain of North Carolina. Journal of Medical Entomology 29(2): 150-159. Ives, J. D. 1938. Cave hibernation of mosquitoes. Jour- nal of the Tennessee Academy of Science 13(1): 15-20. Lawlor, W. K. 1935. Hibernation of Uranotaenia sap- phirina (Osten Sacken) (Diptera: Culicidae). Bul- letin of the Brooklyn Entomological Society 30(1): 14. Mitchell, C. J. 1979. Winter survival of Culex tarsalis (Diptera: Culicidae) hibernating in mine tunnels in Boulder County, Colorado, USA. Journal of Medical Entomology 16(6): 482—487. Mortenson, E. W. 1953. Observations on the overwin- tering habits of Culex tarsalis Coquillett in nature. Proceedings and Papers of the Annual Conference of the California Mosquito Control Association 21: 59-60. Owen, W. B. 1937. The mosquitoes of Minnesota, with special reference to their biologies. University of Minnesota Agricultural Experiment Station Tech- nical Bulletin 126: 75 pp. Peck, S. B. 1988. A review of the cave fauna of Can- ada, and the composition and ecology of the in- vertebrate fauna of caves and mines in Ontario. Canadian Journal of Zoology 66: 1197-1213. Peck, S. B. and J. J. Lewis. 1977. Zoogeography and evolution of the subterranean invertebrate faunas of Illinois and southeastern Missouri. NSS (Na- tional Speleological Society) Bulletin 40(2): 39— 63. Peterson, A. G. and W. W. Smith. and distribution of mosquitoes in Mississippi. Journal of Economic Entomology 38(3): 378-383. Robertson, L. C., S. Prior, C. S. Apperson, and W. S. Irby. 1993. Bionomics of Anopheles quadrima- 1945. Occurrence culatus and Culex erraticus (Diptera: Culicidae) in the Falls Lake Basin, North Carolina: seasonal changes in abundance and gonotrophic status, and host-feeding patterns. Journal of-Medical Ento- mology 30(4): 689-698. 238 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Shemanchuk, J. A. 1965. On the hibernation of Culex tarsalis Coquillet, Culiseta inornata Williston, and Anopheles earlei Vargas, (Diptera: Culicidae) in Alberta. Mosquito News 25(4): 456-462. Snow, W. E. 1955. Feeding activities of some blood- sucking Diptera with reference to vertical distri- bution in bottomland forest. Annals of the Ento- mological Society of America 48: 512-521. Sutton, M. R. 1993. Caves and cave wildlife in a min- eral prospecting area, Oregon and Shannon Coun- ties, Missouri. Missouri Speleology 33(1—4): 138 pp. Williams, D. C. and M. V. Meisch. 1981. A blood host study of riceland mosquitoes in Arkansas County, Arkansas. Mosquito News 41(4): 656-660. Zukel, J. W. 1949. A winter study of Anopheles mos- quitoes in southwestern Georgia, with notes on some culicine species. Journal of the National Ma- laria Society 8: 224—233. Jeffrey K. Barnes, The Arthropod Mu- seum, Department of Entomology, Univer- sity of Arkansas, Fayetteville, Arkansas 72701, U.S.A. (e-mail: jbarnes @ uark.edu). PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 239-242 NOTE First Report of Pryeria sinica Moore (Lepidoptera: Zygaenidae) in North America Pryeria Moore is a monotypic genus ac- commodating P. sinica Moore, 1877. It is eastern Palaearctic, recorded from Russia (Far East), China, Korea, Japan, and Taiwan (Yen and Horie 1997). The holotype of P. sinica, deposited in The Natural History Museum, London, is from China; the ho- lotype of Neopryeria jezoensis Matsumura, 1927, the only synonym of P. sinica, de- posited in the Entomological Institute, Hok- kaido University, Sapporo, Japan, is from Japan. The species has been recorded only recently in Taiwan, where it occurs primar- ily in the subalpine zone (Yen and Horie M97): Pryeria sinica is moderately divergent from other zygaenids in several superficial and morphological features. Although for- merly associated with Phauda Walker (Phaudinae) (Inoue 1982), its assignment to Zygaeninae was convincingly demonstrated by Alberti (1954) and Naumann (1987, 1988). According to Naumann (1987), the genus represents the most primitive lineage of the subfamily. Food plants of P. sinica appear to be re- stricted to Celastraceae, which contain acyanogenic compounds (Epstein et al. 1998). Two genera have been reported as host plants—Celastrus and Euonymus. Yen and Horie (1997) list specimens reared from Celastrus punctatus Thunb., Euony- mus sieboldianus Blume, and Euonymus ja- ponica Thunb. They indicate that the geo- graphical range of Celastrus punctatus, the documented host plant in Taiwan, extends to south China and Japan, which agrees well with the distribution of P. sinica. Sato (1969) reported rearing larvae on Euony- mus sieboldianus, Euonymus alatus (Thunb.) f. ciliatidentatus (Fr. et Sar.) and Celastrus orbicularis Thunb. Yen and Horie (1997) were unsuccessful in transferring larvae from Celastrus to Euonymus in the laboratory, suggesting the possibility of a degree of geographic host specialization. Although Cave and Redlin (1996) iden- tified P. sinica as an important plant pest of Buxus in Korea and China, this is likely an error. According to Yen (personal commu- nication), the common name of Pryeria sin- ica in China is translated as the “buxus zygaenid moth’; however, the common name for Buxus and Euonymus is exactly the same. As a result, hosts for this species are common confused in the entomological literature of China. According to Yen, Eu- onymus and Buxus frequently occur sym- patrically in the mountains of Japan and Taiwan, and he has never observed larvae of P. sinica on Buxus. In April and May of 2001, 2002, and 2003 large infestations of larvae of P. sin- ica were reported on ornamental Euonymus in the city of Fairfax, Fairfax County, Vir- ginia, U.S.A., causing severe damage to the plants. In March 2003 adult specimens were submitted to the USDA Systematic Ento- mology Laboratory at the National Museum of Natural History where they were identi- fied as P. sinica. Subsequently, specimens were reported from Glen Burnie, Anne Arundel County, Maryland, in May 2003. The species previously was unreported from North America. The origins of the Fairfax and Glen Burnie populations are unknown, but introduction via nursery stock from the Far East seems the most likely pathway. Based on the success of P. sinica in northern Virginia on ornamental Euonymus—surviving local winter condi- tions and producing considerable proge- ny—and the degree of damage at the site of the infestation, P. sinica has the potential to become an important pest in situations where Euonymus is used in ornamental landscaping. 240 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. The life cycle of P. sinica has been stud- ied in detail by several workers (e.g., Ishii et al. 1983, Shiotsu and Arakawa 1982, Shiotsu and Tsubaki 1986, Tamura 1981, Tamura and Ouchi 1977, Tsubaki 1981, Tsubaki and Shiotsu 1982, Wipking and Naumann 1992). Eggs are laid in clusters on the stems of the host plant in November and December. They are ovoid, flattened, and covered with scales from the female ab- Pryeria sinica. 1, Adult. 2, Larvae. 3, Damage caused by larvae on Euonymus. dominal hair tufts. At 10—18°C, the egg stage lasts about 80 days. Larvae hatch in March and April and exhibit a group-feed- ing behavior (Tsubaki 1981, Tsubaki and Shiotsu 1982). They may be found on the upper or under surface of the leaves of the food plant, feeding on the leaf edges (Fig. 3). They readily drop on a line of silk when disturbed. Pupation occurs in late May, with an obligate pupal diapause that lasts until VOLUME 106, NUMBER 1 November, when adults emerge. Adults are diurnal, with a slow, fluttering flight gen- erally 1-5 m above the ground. Adults are active from about 0900-1500 hours. They typically mate during the daytime and re- main coupled for about 20 hours. Oviposi- tion takes place during the day, usually within a few hours after copulation. The adult of P. sinica is a medium-sized wasp-mimic (Fig. 1), with a forewing length of 10—13 mm in the male and 12— 14 mm in the female. The wings are trans- parent with a few scattered black scales and a small, diffuse patch of yellow-orange scales in the basal portion of the forewing. Antennae are bipectinate in the male, non- pectinate but clubbed in the female. The distinct forewing coloration, highly reduced proboscis, and tufts of orange scales at the end of the abdomen distinguish this species from all other Zygaenidae. Additional illus- trations of the adult can be found in Yen and Horie (1997) and Epstein et al. (1998). The mature larva (Fig. 2) is 15—22 mm in length. The color is assumed to be apo- sematic (Johki and Hidaka 1979): a white ground color with dorsal, subdorsal, medial, and submedial black longitudinal stripes (comprised of adjacent spots), and a yellow venter. Chaetotaxy is described in detail by Yen and Horie (1997); additional illustra- tions of the larva can be found in Nakajima (1987) and Yen and Horie (1997). The pupa is 8-10 mm in length, 4.0—4.5 mm in width, stout, and dorsoventrally compressed, enclosed in a tough, relatively smooth, beige, flattened, semispherical co- coon. The cocoon typically is spun within a folded leaf or between adjacent leaves on the live plant. Coloration of the pupa is sim- ilar to that of the larva with dorsal, sub- dorsal, submedial, and medial black longi- tudinal stripes (see Yen and Horie 1997 for illustrations). The species is attacked by at least two parasitoids—A grothereutes minousubae Nakanishi (Hymenoptera: Ichneumonidae) (Shiotsu and Arakawa 1982, Shiotsu and Tsubaki 1986) and Bessa parallela (Mei- 241 gen) (= B. selecta fugax Rondani or B. fu- gax Rondani) (Diptera: Tachinidae) (Shima 1973). Based on laboratory studies, larvae of P. sinica produce an (E)-phytol that in- duces predation by the generalist predatory stinkbug, Eocanthecona furcellata (Wolff) (Hemiptera: Pentatomidae) (Yasuda 1998). Given the widespread use of Euonymus as ornamentals, especially in the northeast- ern U.S., the potential seems high for the persistence and expansion of the present in- festation in the U.S. We thank the following for comments on the manuscript that improved its quality and clarity: David Smith, USDA, Systematic Entomology Laboratory, National Museum of Natural History, Washington, D.C., U.S.A.; Shen-Horn Yen, Department of Bi- ological Sciences, Imperial College, Lon- don, United Kingdom; and Clas Naumann, Alexander Koenig Research Institute and Museum of Zoology, Bonn, Germany. LITERATURE CITED Alberti, B. 1954. Uber die stammesgeschichtliche Gliederung der Zygaenidae nebst Revision einiger Gruppen (Insecta, Lepidoptera). Mitteilungen Zoologischen Museum, Berlin 30: 115—480. Cave, G. L. and S. C. Redlin. 1996. Importation of Chinese penjing into the United States, with par- ticular reference to Buxus sinica. Unpublished re- port prepared by USDA, APHIS, PPQ. Epstein, M. E., H. Geertsema, C. M. Naumann, and G. M. Tarmann. 1998. 10. The Zygaenoidea, pp. 159-180. Jn Kristensen, N., ed. Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Sys- tematics, and Biogeography. Handbook of Zool- ogy 4 (35), Arthropoda: Insecta. Walter de Gruy- ter, Berlin and New York. Inoue, H. 1982. Zygaenidae. /n Inoue, H., S. Sugi, H. Kuroko, S. Moriuti, and A. Kawabe, eds. Moths of Japan 1: 191-196, 2: 217-219. Ishii, M., Y. Johki, and T. Hidaka. 1983. Studies on summer diapauses in zygaenid moths. |. Factors affecting the pupal duration in Pryeria sinica. Kontya@ 51: 122-127. Johki, Y. and T. Hidaka. 1979. Function of the “‘warn- ing coloration” in larvae of a diurnal moth, Pry- eria sinica Moore (Lepidoptera: Zygaenidae). Ap- plied Entomology and Zoology 14: 164-172. Nakajima, H. 1987. Pryeria sinica Moore, p. 13. In Sugi, S., ed. Larvae of Larger Moths in Japan. Naumann, C. M. 1987. On the phylogenetic signifi- cance of two Miocene zygaentd moths (Insecta, Lepidoptera). Palaeontological Zoology 61: 299— 308. . 1988. The internal female genitalia of some Zygaenidae (Insecta, Lepidoptera): Their mor- phology and remarks on their phylogenetic sig- nificance. Systematic Entomology 13: 85—99. Sato, R. 1969. The larvae of Pryeria sinica feeding on Euonymus sieboldianus Blume, Euonymus alatus (Thunb.) Sieb. f. ciliatidentatus (Fr. et Sar.) Hi- yama and Celastrus orbicularis Thunb. Yugato (36): 32. Shima, H. 1973. New host records of Japanese Tach- inidae (Diptera: Calyptrata). Sieboldia Acta Biol- ogia (Fukuoka) 4: 153-160. Shiotsu, Y. and R. Arakawa. 1982. One-host one-par- asitoid system: Seasonal life cycles of Pryeria sin- ica (Lepidoptera) and Agrothereutes minousubae (Hymenoptera). Research on Population Ecology, Kyoto University 24: 43-57. Shiotsu, Y. and Y. Tsubaki. 1986. One-host one-para- sitoid system: Population dynamics of a zygaenid moth Pryeria sinica Moore in an undisturbed hab- itat. Research on Population Ecology, Kyoto Uni- versity 28: 333-346. Tamura, M. 1981. Influence of temperature on the ter- mination of diapause in the egg of Pryeria sinica Moore (Lepidoptera: Zygaenidae). Zoen-zasshi 44: 220-224. Tamura, M. and M. Ouchi. 1977. Studies on larval development of Pryeria sinica Moore at constant temperature. Scientific Reports of the Faculty of the Agricultural Ibaraki University 25: 1—4. Tsubaki, Y. 1981. Some beneficial effects of aggrega- tion in young larvae of Pryeria sinica Moore (Lepidoptera: Zygaenidae). Research on Popula- tion Ecology, Kyoto University 23: 156—167. 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Tsubaki, Y. and Y. Shiotsu. 1982. Group feeding as a strategy for exploiting food resource in the burnet moth Pryeria sinica. Oecologia 55: 12-20. Wipking, W. and C. M. Naumann. 1992. Diapause and related phenomena in Zygaenidae moths, 107— 128. In Dutreix, C., C. M. Naumann and W. G. Tremewan, eds. Recent advances in burnet moth research (Lepidoptera: Zygaenidae). Proceedings of the 4th Symposium on Zygaenidae, 11—13 Sept. 1987. Yasuda, T. 1998. Effects of (E)-phytol of several lep- idopteran species in prey-locating behavior of a generalist predatory stinkbug, Eocanthecona fur- cellata (Heteroptera: Pentatomidae). Entomologi- cal Science 1(2): 139-164. Yen, S.-H. and K. Horie. 1997. Pryeria sinica Moore (Lepidoptera, Zygaenidae), a newly discovered relic in Taiwan. Transactions of the Lepidoptero- logical Society of Japan 48(1): 39-48. John W. Brown, Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC-168, Washington, DC 20013-7012, U.S.A. (e-mail: —jbrown@sel.barc.usda. gov); Marc E. Epstein, Department of Sys- tematic Biology, Smithsonian Institution, National Museum of Natural History, Washington, DC 20013-7012, U.S.A.; and Eric R. Day, Insect Identification Lab., De- partment of Entomology, Virginia Tech University, Blacksburg, VA 24061, U.S.A. PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 243-244 OBITUARY Alexander Douglas Campbell Ferguson 1926—2002 (a | Stay ay Ot ears = > mee, = wy Douglas C. Ferguson, 1926—2002 [1979, National Museum of Natural History Staff Directory] Dr. Douglas C. Ferguson (Doug to ev- eryone who knew him) died on 4 Novem- ber 2002 following surgery on 16 October. Doug was born in Halifax, Nova Scotia, on 17 February 1926, attended local schools, and received a B.S. from Dalhousie Uni- versity in 1950. His M.S. (1956) and Ph.D. (1967) were awarded by Cornell University. He was a field assistant to J. H. Mc- Dunnough in 1946; Curatorial Assistant, Curator of Entomology, and Chief Curator (Science Division) at the Nova Scotia Mu- seum (1949-1963); Research Associate in Entomology (Peabody Museum of Natural History) then Research Staff Biologist and Lecturer (Department of Biology) and Cu- ratorial Associate in Entomology (Peabody Museum of Natural History), Yale Univer- sity (1963-1969); and Research Entomol- ogist, Systematic Entomology Laboratory U.S.D.A. at the National Museum of Nat- ural History (1969-1996). Upon retirement he continued as a Collaborator of the U.S. Department of Agriculture and Research Associate of the Smithsonian Institution. Doug’s interest in natural history began in childhood when he seriously watched birds and discovered the nests of most local species. After reading W. J. Holland’s ac- count of sugaring for moths in The Moth Book in 1941, he tried it on the trees around his home and was thrilled to catch five spe- cies of underwing moths (Catocala) the first night. Halifax was a-small city with 244 many collecting sites within walking or cy- cling distance, and it had a museum with a collection of local Lepidoptera, a library, and a helpful director. Doug’s initial in- volvement with the Lepidoptera increased exponentially and resulted in The Lepidop- tera of Nova Scotia, part 1, Macrolepidop- tera in 1954. He was deeply influenced by the prominent lepidopterists James Mc- Dunnough, W. T. M. Forbes, Charles Rem- ington, and John Franclemont during his formative years. Throughout his career Doug was an avid, knowledgeable collector. Field trips were directed to learn and document the fauna of particular areas. He collected in the south- ern parts of the Provinces and all States but Hawaii and several times in many. He spread and labelled an estimated 200,000+ specimens during his career. These speci- mens have augmented significantly the holdings of the National Museum of Natu- ral History, the Peabody Museum of Nat- ural History, and the Nova Scotia Museum. He was extremely interested in learning the life history of species and reared to the adult stage more than 600 species, docu- menting many of them with 35 mm slides of the larvae and adults. A chance meeting in 1967 with Richard B. Dominick, a Yale alumnus and Lepidop- teran enthusiast, at the Peabody Museum led to several collecting trips at The Wedge, Dick’s estate near McClellanville, South Carolina. Here began the Moths of North America project and subsequently the es- tablishment of the Wedge Entomological Research Foundation, which funds and pub- lishes the series. Doug enlisted the partici- pation of John Franclemont, Eugene Mun- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON roe, and me for the project, originally pro- jected to be a synoptic update of Holland’s moth book. Studied consideration led to the project’s present scope of an anticipated 130° fascicles to treat the estimated 16,000* species in the area. Doug contributed fas- cicles on the Saturniidae and Geometrinae and had the text and line drawings com- pleted for a major revision of the geometrid tribes Cassymini and Macariini before his death. Doug was an excellent field biologist who interacted and collaborated with many Lepidopterists. As well, he aided many col- lectors by identifying specimens and occa- sionally describing species whose identity was needed for economic or biologic pur- poses. Doug had two students: Roger Heitz- man (Ennominae) and Alma Solis (Pyralo- idea). He was very generous with his knowledge and would drop what he was doing to answer their questions. Doug was a quiet, thoughtful, well-read person who had many interests, history, gardening, and classical music among them. In group meet- ings he usually was reserved; however, he often brought a carefully reasoned and val- ued view to discussions. Doug is survived by Charlotte, his wife of 49 years, daughters Stephanie and Car- oline, and six grandchildren. He is held in high esteem and is sorely missed by many friends and colleagues. Ronald W. Hodges, retired, Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Ag- riculture. Current address: 85253 Ridgetop Drive, Eugene, OR 97405, U.S.A. (e-mail: rwhodges @ earthlink.net). NOTICE OF NEW PUBLICATION A Catalog of the Cecidomytidae (Diptera) of the World Entomological Society of Washington Memoir No. 25 This catalog, the first complete catalog of the family since 1913, lists the 5,451 species and 598 genera of living and fossil Cecidomyiidae or gall midges of the world. It provides information on species distribution, hosts, and types, and original and subsequent helpful references. Within subfamilies, genera are listed in alphabetical order but are each cross referenced in an appendix where they are arranged in an annotated classification. One new species is named, many new names, new synonyms, and new combinations are proposed, and several type species are designated. A single index lists all generic and specific names of Cecidomyiidae with their authors, as well as hosts, host family for plants, host order and family for arthropods, and order for fungi. ENTOMOLOGICAL SOCIETY OF WASHINGTON http://entomology.si.edu/ES W/ES W Menus. lasso Information and Officers Bylaws History First 100 Years Past Presidents History of the ESW Seal Information for Authors Available Publications Meetings Members Database Membership Application Subscription THE ENTOMOLOGICAL SOCIETY OF WASHINGTON PRELIMINARY SCHEDULE OF THE REGULAR MEETINGS FOR 2003-2004 October 2: Chris Desjardins (University of Maryland & Maryland Center for Systematic Entomology). Diparine Wasp Diversity and Collecting in South Africa and Aus- tralia November 6: Stefan Cover (Museum of Comparative Zoology, Harvard University). Ant Systematics in the 21st Century: North America, a Case Study December 4: John Strazanac (University of West Virginia). Diversity of Central Ap- palachian Forest Canopy Macrolepidoptera, Symphyta, and their Tachinid Para- sitoids January 8: Jonathan Mawdsley (National Fish & Wildlife Foundation). Pollination by Beetles (Insecta: Coleoptera): An Overview February 5: Jeremy Miller (Smithsonian Institution, Entomology). The Unbearable Lightness of Being Monophyletic: Clade Stability and the Addition of Data in Erigonine Spiders March 4: Frank Carle (Rutgers University). Evolution of Dragonflies April 1: Caroline Chaboo (Cornell University & American Museum of Natural History). Untangling Fecal Shield Architecture in Tortoise Beetles: Behavior, Ecology, Morphology and Phylogeny (Coleoptera: Chrysomelidae: Cassidinae) May 6: Leeanne Alonso (Conservation International). Contributions of Rapid Assess- ment of Insects to Biodiversity Conservation Meetings are held at: 7:00 pm Cathy Kerby Seminar Room (CE-340) National Museum of Natural History 10th & Constitution Ave., Washington, D.C. Meetings are open to the public Please bring new literature, specimen demonstrations, ento-T-shirts, etc. Pre-meeting dinner: R. Reagan Building Food Court, 5:30 p.m. Refreshments served following the meetings Jonathan Mawdsley, President David G. Furth & John Brown, Co-Program Chairs PROC. ENTOMOL. SOC. WASH. 106(1), 2004, pp. 247-248 INSTRUCTIONS FOR AUTHORS General Policy.—Articles for publica- tion are ordinarily accepted only from members of the Society and must be in En- glish. A summary in French, German, Spanish, or Russian is acceptable. Such a summary should be placed immediately af- ter the English abstract and may be a trans- lation of that abstract. Manuscripts should not exceed 15 printed pages (about two and one-half double-spaced typewritten pages equal one page). Manuscripts are peer-re- viewed before acceptance. Final acceptance is the responsibility of the Editor. Articles are published in the order received, not the date of acceptance. Immediate publication is available with payment of full page charges. Notes, book reviews, and obituar- ies are published as space is available. 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See recent issues for styles for journals, books, book chapters, etc. When referring to references in text, use the following; Henry (1990), (Henry 1990), (Henry 1987, 1990), (Smith 1990, Henry 248 1992), (Smith 1990>"Henry F990; 1991); (Miller 1990a, 1990b) where two articles are published by the same author in the same year, or Henry et al. 1990 where more than two authors are involved (do not ital- icize ‘‘et al.’’). Citations “‘in press’’ should be cited as “‘in press,’ not with year of expected publication. Names of persons providing unpublished information should include initials and be referenced in the text as: W. Mathis (per- sonal communication) [information ob- tained orally] or W. Mathis (in litt.) [infor- mation obtained in a writing]. Names of organisms.—The first men- tion of a plant or animal should include the full scientific name including the authority. Use only common names approved in Com- mon Names of Insects and Related Organ- isms published by the Entomological Soci- ety of America. Spell out the entire scien- tific name the first time it is mentioned, thereafter abbreviate generic names; how- ever, do not abbreviate a genus name at the beginning of a paragraph or sentence, or if two or more genera being discussed have the same first letter. Within sentences, al- ways use the genus name or initial preced- ing a species name. Taxonomic papers.—Taxonomic papers must conform to requirements of the Inter- national Code of Zoological Nomenclature. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON In addition, type specimens must be desig- nated for new species described, type de- positories must be clearly indicated, and new taxa must be clearly differentiated from existing taxa by means of keys or dif- ferential diagnoses. In short, these papers must conform to good taxonomic practices. 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Acceptance of papers is based only on scientific merit without regard to the author’s financial sup- port. PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael E. Schauff. 85 pp. 1990 A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. Eric Grissell and Michael E. Schauff. 87 pp. 1997 Revision of the Oriental Species of Aphthona Chevrolat (Coleoptera: Chrysomelidae), by Alexander S. Konstantinov and Steven W. Lingafelter. 349 pp. 2002 Revision of the Genus Anoplophora (Coleoptera: Cerambycidae), by Steven W. Lingafelter and E. Richard Hoebeke. 236 pp. 2002 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 No. 5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 __. No.6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi VEE: Ser sy PS go s/h REDS MR ee le ees ERA a SW od OLB EH Sia d Bh A AE AORTA IR VLE No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato- pogoenidae), by W.:L. Grogan; Jr. and W: W.2 Wirth: 125 pp.1979 _ te 9S eee No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. 07/10) OFC he SE See Wee RO ENR ee TM Oe Ie LIS Se me Rie NeT oF ott No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 POP Sy wae a erg IS ad AN, eed ah eek A Se A ee Beaton Paes, Medea) REE WA Eth es a el a No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent andi Wee KGrogan: Irv OS pp OO Sek uses te cud lu EAU eT ae ee No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 ___ No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited bye Be Nordenand: AY SiMenke 21 6rpp. 1990: <8 See ee eee eee No. 18. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and Wilhtamyl: Grogan yn DOI pp yhoo Te Se ie ee se A oe 0S PT he Ee Saye A este tee tp 5 ere No. 19. Monograph of the Stilt Bugs. or Berytidae (Heteroptera), of the Western Hemisphere, by Wihomasw henry: LAO py lO OG hee Sa jek Cy Le a a SE eee en See ee No. 20. The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- Se Ved Leer QU IMP oy oR MEL ep ee Renee OS ip ea Me lh ue ncaa ee PARE ROA a Le NA PN alte TN Mee No. 21. New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. Bertins LOW shOO Ss 3 SSL Sere ANY a dee bs) Mie Ne Bee oe 2 eR ple ete oe ee See ae No. 22. Systematics of the North American Species of 7richogramma Westwood (Hymenoptera: iimaichorrammatidae): (by. John Dy Pintos,287 ppt LOGS nae Sei Bee ee eee ee No. 23. Revision of the Net-Winged Midges of the Genus Blepharicera Macquart (Diptera: Blepha- riceridae) of Eastern North America, by Gregory W. Courtney. 99 pp. 2000) __------------ No. 24 Holcocerini of Costa Rica (Lepidoptera: Gelechioidea: Coleophoridae: Blastobasinae), by David f'sWakct an st a'Vyil &2 Wf 0) 0494 0] OPA ph SeASAMMRIRRUE aeae EM nage Sarat eben, MFe dtr lguiNy op Aree MNES sk Ae paieek tPA SUS A Ad No. 25 A Catalog of the Cecidomyiidae (Diptera) of the World, by Raymond J. Gagné. 408 pp. 2004 $10.00 15.00 40.00 30.00 $15.00 15.00 15.00 15.00 12.00 5.00 25.00 25.00 25.00 25.00 25.00 18.00 12.00 12.00 28.00 14.00 18.00 50.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume to non-members and $25.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra on all orders. Dealers are allowed a discount of 10 percent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Treasurer, Entomological Society of Washington, % Department of Entomology, MRC 168, Smithsonian Institution, Washington, D.C. 20560-0168. CONTENTS (Continued from front cover) MACKAY, WILLIAM P.—A new species of the ant genus Acanthostichus Mayr (Hymenoptera: Formicidae) from Paraguay, and a description of the gyne of A. brevicornis Emery ............... 97 MAWDSLEY, JONATHAN R.—Pollen transport by North American TJrichodes Herbst (Coleopteran\Glemndachana. toro: hoa = se cic eee gee CRIN LTE chat emia ial’ asi tictoe nie cities 199 MENDES, LUIS F. and GEORGE O. POINAR, JR.—A new fossil Nicoletiidae (Zygentoma, SApLeny gota) TaD omni Canam Dei ie ccrmaee eens Oommen cnc. eeicls ita) clare niet ateiatealcte & dlatael sere GES 102 MORSE, JOHN C. and LIANFANG YANG—The world subgenera of Glossosoma Curtis (Trichoptera: Glossosomatidae), with a revision of the Chinese species of Glossosoma subgenera Synafophora Martynov and Protoglossa ROSS .............0 cee eect e eet e ees 52 NISHIDA, KENJI and DAVID ADAMSKI—Two new gall-inducing Saphenista Walsingham (GepidopteraPortricidacs Cochyliny) trom! CostaiRica yt) Seabees) eo aeeaiietan sane eke 133 SONG, HOJUN—Post-adult emergence development of genitalic structures in Schistocerca Stal and Locusta L. (Orthoptera: Acrididae) .................. nh Ripe tea Oe Ce cs Mam ake 18] SUEYOSHI, MASAHIRO and WAYNE N. MATHIS—A new species of Cyamops Melander 1913 (Diptera: Periscelididae) from Japan and a review of Japanese Periscelididae ................ 74 WAGNER, DAVID L., DAVID ADAMSKI, and RICHARD L. BROWN—A new species of — Mompha Hiibner (Lepidoptera: Coleophoridae: Momphinae) from buttonbush (Cephalanthus occidentalis \..) with, descriptions) of the; carly stages ¥ion.i... stan yee ioe cee on ee tees ieee ee 1 = WHEELER, A. G., JR.—Largidea davisi Knight, a rarely collected plant bug (Hemiptera: Miridae: Deraeocorinae) associated with pitch pine in the northeastern United States ........ 85 WHEELER, A. G. JR. and E.. RICHARD HOEBEKE—New North American records of the European broom psyllid Arytaina genistae (Latreille) (Sternorrhyncha: Psyllidae) .......... oL/G NOTES BARNES, JEFFREY K.—Overwintering cave mosquitoes (Diptera: Culicidae) of the Arkansas AMI SSOULINOAATKS otal Soe Se. coe oe cick. 2 obill. Sac rade eee) Se MS AiR OS ne a Sars eal RO 235 BROWN, JOHN W., MARC E. EPSTEIN, and ERIC R. DAY—First report of nae sinica Moore Gzepidopteras Ay caemdae).inviNorth Amesica, 2 'o:k.scawek oo sk oe ote as 2are eae see ae 239 REEVES, W. K. and J. A. KORECKI—Ochilerotatus japonicus japonicus (Theobald) (Diptera: Culicidae), a new invasive mosquito for Georgia and South Carolina ........................ 233 OBITUARY HODGES, RONALD W.—Alexander Douglas Campbell Ferguson 1926-2002 ............. ee oe 243 MISCELLANEOUS Notice of New Publication: A Catalog of the Cecidomytidae (Diptera) of the World .............. 245 Preliminary Schedule of the Regular Meetings for 2003-2004 ................ ccc cece cece eee e es 246 TAStructloms OPA TUTMOTS Wyo. iss. is eee cece eecdeee ear a cel eee sg Re Te eee Gra 247 VOL. 106 APRIL 2004 NO. 2 OL ; (ISSN 0013-8797) L{ wl eof PROCEEDINGS E of the ENTOMOLOGICAL SOCIETY = st : jy Ny ¥ . CONTENTS BRAILOVSKY, HARRY and ERNESTO BARRERA—Two new genera and four new species of Colpurini (Heteroptera: Coreidae: Coreinae) fron New Guinea ................0. 00 cece e eee 424 BROWN, JOHN W. and RICHARD L. BROWN—A new species of Cryptaspasma Walsingham (Lepidoptera: Tortricidae: Olethreutinae) from Central America, the Caribbean, and south- eastern United States, with a catalog of the world fauna of Microcorsini .................... 288 COSTA, LUIZ A. A. and THOMAS J. HENRY—Fulvius chaguenus Carvalho and Costa (Heteroptera: Miridae: Cylapinae: Fulviini): Redescription and recognition of type specimens 417 ETNIER, DAVID A., CHARLES R. PARKER, and IAN C. STOCKS—A new species of Rhyacophila Pictet (Trichoptera: Rhyacophilidae) from Great Smoky Mountains National Park, with illustrations of females of R. appalachia Morse and Ross and R. mycta Ross .... 396 GAGNE, RAYMOND J., ALEJANDRO SOSA, and HUGO CORDO—A new Neotropical species of Clinodiplosis (Diptera: Cecidomyiidae) injurious to alligatorweed, A/ternanthera Pam R erties CNIMATAM HACE AE) Rasen Mnep eee ake ss Ticats Ui diel Us dio dcie ale ches oes sacle sleis-¥ Sleia n oidteleiets 305 HARRINGTON, RICHARD C. and JOHN C. MORSE—A new species of Brachycentrus Curtis (Trichoptera: Brachycentridae) from the southern Appalachian Mountains and variation in the egtalsi tay EL IG aL r mare OIE) Aces A Oh Ud eH SAD oe a a ee ee 453 ISHIKAWA, TADASHI and SHUJI OKAJIMA—A new species of the saicine assassin bug genus Carayonia Villiers (Heteroptera: Reduviidae) from Indochina ......................0.0.000005 319 KONSTANTINOV, ALEXANDER S. and BORIS A. KOROTYAEV—Sexual dimorphism and size of aedeagi in apionid weevils (Coleoptera: Apionidae) and flea beetles (Coleoptera: Chrysomelidae): Why some masculine males have small aedeagi ..................--....0++- 324 MATHIS, WAYNE N. and TADEUSZ ZATWARNICKI—A revision of the shore-fly genus Cressonomyia Arnaud (Diptera: Ephydridae), with comments on species that have been mea pee eee riaein mamas eek ptr Lenten PNM ATE Wee Eyl RLS cls. Al isieile-a acts wiles spate See's «oe 249 NICKLE, DAVID A.—Commonly intercepted thrips (Thysanoptera) from Europe, the Mediterranean, and Africa at U.S. ports-of-entry. Part Il. Frankliniella Karny and Iridothrips ia eevatere (CAM ain eae eet SP Uh Coes eI a ee 438 OCAMPO, FEDERICO C. and MIGUEL ANGEL MORON—Description of the third instar larva of Hemiphileurus dispar Kolbe (Coleoptera: Scarabaeidae: Dynastinae: Phileurini) .. 412 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2004 E. E. GrissEL_L, President Jon A. Lewis, Custodian JASON P. W. Hatt, President-Elect MIcHAEL G. PoGuE, Treasurer Stuart H. McKamey, Recording Secretary | JOHN W. BROWN AND Davip G. FurtH, Program Chairs Ho iis B. WiLLiAMs, Membership Secretary JONATHAN R. Mawps-ey, Past President Davip R. SmitH, Editor Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY Wayne N. MartuHis Honorary President DonALD M. ANDERSON Honorary Members WILLIAM E. BICKLEY KARL V. KROMBEIN RONALD W. HopGEsS PAUL J. SPANGLER Manya B. STOETZEL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168. MEETINGS.—Regular meetings of the Society are held in the Natural History Building, Smithsonian [nstitu- tion, on the first Thursday of each month from October to June, inclusive, at 7:00 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $25.00 (U.S. currency). PROCEEDINGS.—The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub- lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send address changes to the Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomo- logical Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE PP. 247-248 OF THE JANUARY 2004 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 30 March 2004 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 249-279 A REVISION OF THE SHORE-FLY GENUS CRESSONOMYIA ARNAUD (DIPTERA: EPHYDRIDAE), WITH COMMENTS ON SPECIES THAT HAVE BEEN EXCLUDED WAYNE N. MATHIS AND TADEUSZ ZATWARNICKI (WNM) Department of Entomology, NHB 169, P.O. Box 37012, Smithsonian Institu- tion, Washington, D.C. 20013-7012, U.S.A. (e-mail: mathis.wayne@nmnh.si.edu); (TZ) Museum and Institute of Zoology, Polish Academy of Sciences, ul. Wilcza 64, 00-679 Warsaw, Poland, and Department of Biosystematics, University of Opole, ul. Oleska 22, 45-052 Opole, Poland (e-mail: zatwar@uni.opole.pl) Abstract.—Five species of the shore-fly genus Cressonomyia Arnaud are revised, in- cluding C. bolivia, n. sp. (Bolivia. La Paz: Tajlihui). The genus is known only from the New World tropics and subtropics and is the sister group to Peltopsilopa Cresson. Two species groups are recognized, the aciculata group (two species) and the skinneri group (three species). Psilopa aeneonigra (Loew), which had been included in Cressonomyia, is returned to Psilopa. A neotype is designated for Psilopa aciculata Loew, the senior synonym for Plagiops nitidifrons Cresson (new synonym). All species are illustrated and distribution maps are also provided. Key Words: This revision treats species of Cresson- omyia Arnaud, which are known only from the New World. The species of Cressono- myia have never been treated comprehen- sively, although Cresson (1942, 1946) pro- vided a review, including keys, in his treat- ments of the Nearctic and Neotropical shore-fly faunas. The primary objectives of this paper are to revise the species and to present a phylogenetic context for the genus Cressonomyia within the tribe Psilopini Cresson, subfamily Discomyzinae Acloque. We recharacterize Cressonomyia and de- scribe and illustrate several characters of the male terminalia that have not been re- ported or analyzed previously. These struc- tures are described and illustrated for all known species and are included in the phy- logenetic analysis. Although the generic concept of Cres- sonomyia has remained relatively stable review, Diptera, Ephydridae, shore flies, Cressonomyia since its description, the genus-group no- menclature was confused for decades be- cause of preoccupied names. The generic status was first recognized in 1918 as the genus Plagiops Cresson, a preoccupied ge- nus-group name (Townsend 1911). Cresson recognized his nomenclatural lapse and pro- posed the replacement name of Plagiopsis Cresson, which, unfortunately, was also preoccupied (Brauer and Bergenstamm 1889). Both Plagiops and Plagiopsis were first proposed for tachinid flies, and Arnaud (1958), a specialist on Tachinidae, ultimate- ly corrected the nomenclatural oversights by suggesting Cressonomyia as the valid, replacement name. As E. T. Cresson, Jr. de- scribed most of the species included in the genus, it is appropriate that the generic name be a patronym to honor and recognize him. The type species for Plagiops is P. nitidifrons Cresson, which automatically 250 became the type species for the replacement names. Thus far eight names are available for species of Cressonomyia (Mathis and Za- twarnicki 1995). Loew (1862) described the first species, Psilopa aciculata, from spec- imens collected in Cuba, and the second species, Ephygrobia metallica Schiner (1868), was later found to be conspecific with C. aciculata (Cresson 1925). Loew (1878) also described the third species, Psi- lopa aeneonigra, from specimens collected in Texas, and like the first species he named, it too became a senior synonym when the fourth species, Psilopa fulvipennis Hine (1904) from Louisiana, was discov- ered to be conspecific with C. aeneonigra (Cresson 1942). The correct generic status of Psilopa aeneonigra and P. fulvipennis 1s reported in this paper. Cresson named the other four species, although none was de- scribed in Cressonomyia. The four species in their original generic combination are: Plagiops hinei (Cresson 1922); Psilopa meridionalis (Cresson 1918); Plagiops ni- tidifrons (Cresson 1918); and Psilopa skin- neri (Cresson 1922). Aside from catalog en- tries (Wirth 1965, 1968; Mathis and Za- twarnicki 1995) and Cresson’s synopses (1942, 1946), there are no other substantive papers on Cressonomyia. Cressonomyia has always been associat- ed with the genus Psilopa and related gen- era, which were usually recognized as the tribe Psilopini (Cresson 1942, 1946; Wirth 1965, 1968; Mathis and Zatwarnicki 1995). We continue with that precedent, as all mor- phological evidence (see key and diagnosis below) substantiates that tribal placement. We also adhere to Zatwarnicki’s (1992) characterization of Psilopini, which ex- cludes genera that are more closely related to Discomyza Meigen as a separate tribe, Discomyzini Acloque. Both Discomyzini and Psilopini are tribes in the subfamily Discomyzinae (Mathis and Zatwarnicki 1895): METHODS AND MATERIALS The descriptive terminology, with the ex- ceptions noted in Mathis (1986) and Mathis PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and Zatwarnicki (1990a), follows that pub- lished in the Manual of Nearctic Diptera (McAlpine 1981). Because specimens are small, usually less than 3.5 mm in length, study and illustration of the male terminalia required use of a compound microscope. We have followed the terminology for most structures of the male terminalia that other workers in Ephydridae have used (see ref- erences in Mathis 1986, and Mathis and Za- twarnicki 1990a, b), such as pre- and post- surstylus. Zatwarnicki (1996) has suggested that the pre- and postsurstylus correspond with the pre- and postgonostylus and that the subepandrial plate is the same as the medandrium. The terminology for struc- tures of the male terminalia is provided di- rectly on Figs. 2-15. The species descrip- tions are composite and not based solely on the holotypes. One head and two venational ratios that are used in the descriptions are defined below (all ratios are based on three specimens: the largest, smallest, and one other). Gena-to-eye ratio is the genal height measured at the maximum eye height di- vided by the eye height. Costal vein ratio: the straight line distance between the apices of R,,, and R,, 23 INBIO); Chomogo area (10°18'N, 84°47'W; 16205m)s 13 sum 1973, Te Erwint (Ge E Hevel (1 d, 1 2; USNM). Cartago: Peralta (9°58/N; 83°37 W; 332 m); 7 Aus 1909.2 P. Calvert (1 6; ANSP). Guanacaste: Ha- cha, Finca el Oro (5 km S de Hacienda Ale- mania; 11°01.4'N, 85°27.5’W; 400 m), 14— 19 Apr 2002, D. Bricefio (8 6, 11 2; IN- BIO); Pitilla (9 km S Santa Cecilia; 10359:54N; 85 25.8 W; 700 m), Feb—7 Apr 1989, 1993, P Rios (6 6, 8 2; INBIO). Heredia: Finca Naranja Valenciana (2 km S Pueblo Nuevo, Sarapiqui; 10°28'N, 84°07'W; 90 m), 31 Jan 1993, M. Ortiz (2 2; INBIO); La Virgen de Sarapiqui (10°23.7'N, 84°08.5'W), 9-24 Apr 1993, M. Ortiz (1 2; INBIO); Santo Domingo, INBio Parque (9°58.4'N, 84°5.6’W), 18 Feb 2003, W. N. Mathis (1 do, 1 2; USNM). Limon: Batan, 16 Jun 1951, L. L. Cart- wright (1 6; USNM); Guacimo (10°13’N, 83°41'W; 110 m), 6 Jun 1909, P. P. Calvert (1 6; paratype; ANSP). Puntarenas: Esta- ci6n Agujas (8°32.2'N, 83°25.5'W; 300 m), 19—24 Mar 1997, A. Azofeifa (1 2; IN- BIO); Estacié6n Cabuya (9°35.3'N, 85°05.9'W; 50 m), 22 Mar 1997, E Alva- rado (1 ¢; INBIO); Estaci6n Carara (9°46.5'N, 84°31.6'W; 200 m), Feb 1990, R. Zuniga (1 6, 2 2; INBIO); Estacién Quebrada Bonita (9°46'N, 84°36.5'W; 50 m), 21 Mar-21 Apr, 1989, R. Zuitiga (1 6. 1 2: INBIO); Estacion Sirena, Corcovado (8°28.8'N, 83°35.5'W), 21 Mar—21 Apr 1992, Z. Fuentes (1 2; INBIO); Montev- erde, 26 Mar 1987, W. E. Steiner (13 ¢, 21 2; USNM); Rancho Quemado (8°40.8'N, 83°34'W; 200 m), Apr 1992, K. Flores (1 2; INBIO). San José: La Caja (8 km W Sanmeliosé)) 28° Apr 1B. iSchmidti(2Qeue: USNM); Pedregoso (9°22'N, 83°43'W), D. L. Rounds (1 9; ANSP). CUBA. Camagiiay: Camagiiay, 3 Dec 1917, J. V. McGuire (1 6; USNM). Cien- fuegos: Soledad, Jardin Botanico (22°7.5'N, 80°19.2’W), Jan—13 Dec 1927, 1994, C. T. Brues, W. N. Mathis, G. Salt G 6, 6 2; MCZ, USNM). Granma: Cayamas, 26 Jan, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON E. A. Schwarz (1 46; USNM). Guantdna- mo: Baracoa (30 km SE), 26 Feb 1992, M. von Tschirnhaus (1 6; USNM); Guantana- mo, 10 Nov 1914, H. Skinner (1 6, 4 9; ANSP); Pt. Barrios, 3-14 Mar 1905 (1 @; ANSP); San Carlos Est., 4-8 Oct 1913 (1 36; ANSP). Havana: Puerto Escondido, 26 Apr 1983, W. N. Mathis (1 2; USNM). Holguin: Holguin, north, Feb 1992, M. von Tschirnhaus (3 6, 3 2; USNM); Santa Lu- cia (3 km N), 22 Feb 1992, M. von Tschirn- haus (1 2; USNM). Isla de Pinos: Mar, C. W. Metz (1 6, 2 2; MCZ, USNM). Matan- zas: Palpite (1 km NE), 2 May 1983, W.N. Mathis (1 2; USNM). Oriente: Herradura (20°40'N, 76°48°W), ‘CC. W. Metz (ise; MCZ); Santigo de Cuba (20°45'N, 76°2'W), 23 Feb 1992, M. von Tschirnhaus 2 6,2 82? USNM): Pinar del Rios Soroa (2 km NW; 22°48.6’N, 83°1.0’W), 4—5 Dec 1994, W. N. Mathis (1 2; USNM). Santa Clara: Soledad, 1—8 Jun 1939, C. T. Par- sons (1 6, 1 2; MCZ, USNM). DOMINICA. Layou River mouth, 9 Jan 1965, W. W. Wirth (1 6; USNM). Spring- field Plantation, 18—22 Jul 1978, G. C. Steyskal (2 ¢6, 5 2; USNM). DOMINICAN REPUBLIC. Distrito Na- cional: Ciudad Trujillo (6 km W; = Santo Domingo), 16—20 Dec 1955, J. Maldonado Capriles (1 2; USNM). La Vega: El Rio (9.5 km E; 19°0.9'N, 70°33.5'W; 980 m), 6 May 1995 (1 2; USNM); Rio Camu (3.5 km NW La Vega; 19°13.7'N; 70°35.2'’W; 100 m), 10 May 1995, W. N. Mathis (2 d, 10 2: USNM); Valle del Rio, 28 Dec 1955, J. Maldonado Capriles (2 2; USNM). Ped- ernales: Cabo Rojo (23.5 km N; 18°06’N, 71°38'W; 540 m), 13-19 Jul 1990, L. Mas- ner, J. Rawlins, C. Young (1 2; CMP). ECUADOR. Los Rios: Quevedo (40 km S), 11 Jul 1975, J. Cohen, Peterson, Thorn- dal (1 6; USNM). EL SALVADOR. San Salvador, 14 Jun 1958, L. J. Bottimer (1 2; USNM). GRENADA. St. Andrew: La Force Bridges (12°07.6'N, 61°39.8'W),, 19° Sep 1996, W. N. Mathis (2 2; USNM). GUYANA. Georgetown (6°48.6'N, 58°08.6'W), 20-29 Aug 1997, W. N. Math- VOLUME 106, NUMBER 2 is (1 6; USNM):; Georgetown, Atkinson Airport Road, 2 Jun 1965, S. M. Gaud, L. E Martorell (1 2; USNM). Karanambo, Ru- pununi River (ox bow; 3°45.1'N, 59°18.6’W), 2 Apr 1994, W. N. Mathis (9 36, 8 2; USNM). HAITI. (1 6, 1 9; USNM). HONDURAS. Lancetilla (15°42'N, 87°28'’W), Aug, Stadelmann (1 2; MCZ). JAMAICA. Clarendon: Barnswell Beach (17°45.’N, 77°08.5’W), 13 May 1996, D. and W. N. Mathis, H. Williams (4 6, 4 2°; USNM); Farquhars Beach (17°50.9'N, 77°22.8'W), 9 May 1996, D. and W. N. Mathis, H. Williams (3 ¢, 4 2; USNM); Jackson Bay (17°44.7'N, 77°12.6'W), 13 May 1996, D. and W. N. Mathis, H. Wil- liams (3 6,5 2; USNM); Portland Cottage (17°45.4'N, 77°11'W), 13 May 1996, D. and W. N. Mathis, H. Williams (1 6,3 &; USNM); Portland Cottage (1 km S; 17°45.8'N, 77°12.6'W), 13 May 1996, D. and W. N. Mathis, H. Williams (1 ¢, 1 @; USNM); Salt River (4 km N; 17°52.1'N, 77°09.5'’W), 13 May 1996, D. and W. N. Mathis, H. Williams (1 2; USNM); Toll Gaten(7 a kui S5el7°58/ N... 77°22:3) W)..9 May 1996, D. and W. N. Mathis, H. Wil- liams (1 6, 1 2; USNM). Manchester: Ba- tersea (17°13'N; :77°29' W),. Feb: 1910, R: Thaxter (1 6; ANSP); Mandeville, Feb 1910, R. Thaxter (1 2; ANSP); near Man- deville (18°03.5'N, 77°31.9'W), 15 Apr—7 May 1996, 2000, D. and W. N. Mathis, H. Williams (12 6, 24 9; USNM); near War- wick (17°54.1'’N, 77°25.5'W), 7 May 1996, D. and W. N. Mathis, H. Williams (1 &; USNM). St. Andrew: Irishtown (5 mi SW; via road), 7 Dec 1975, G. E Hevel (1 6; USNM); Ferry River, 13 May 1941, E. Chapin (1 2; USNM); Hardwar Gap, 10 Marni1970s; a. ABareW.) W.\Wirthy Gord; USNM). St. Catherine: Fresh River at Fer- Py 22 Jul (1962; TeFarr,:OnS Punt (oie; USNM). St. Elizabeth: Balaclava (18°10'N, 77°39'W), R. Thaxter (1 2; MCZ); Black River (18°01.4’N, 77°51.1’W), 11 May 1996, D. and W. N. Mathis, H. Williams (3 Cn 2 162s SUSNND ss Blinin: (ISPO7k Ne 77°40.5'W), 10 Apr 2000, W. N. Mathis (1 261 oiyiv2, 1 2s LUSNM): Mageotty) Falls (18°08.2'N, 77°45.1’W), 18 Apr 2000, W. N. Mathis (1 2; USNM). MEXICO. Chiapas: Boca de Ciela (17 km S Puerto Arista), 18 May 1985, A. Freidberg, W. N. Mathis (1 2; USNM); Cascadas de Agua Azul (62 km S Pal- enque), 7 May 1985, W. N. Mathis (1 6, 1 2: USNM): Finca Prusia (33 km S Jalten- ango; 1,000 m), 10-12 May 1985, W. N. Mathis (1 2; USNM); Union Juaréz (9 km S), 23 Apr 1983 (1 6, 2 2; USNM). Jal- isco: Puerto Vallarta, 5 Oct 1984, G. E. Bo- hart (2 6; BYU). Tabasco: Teapa (8 km SW), 6 May 1985, W. N. Mathis (1 2; USNM). Veracruz: Ciudad Aleman, 3 May 1985, W. N. Mathis (2 6, 2 2; USNM); Fortin de las Flores, 2 May 1985, W. N. Mathis (1 ¢; USNM); Ocotal Chico (600 m), 4-5 May 1985, W. N. Mathis (1 <4; USNM). MONTSERRAT. Trinidad, 29 Jun 1905, A. Busck (1 2; USNM). PANAMA. Canal Zone: Coraza, 1—6 Mar 1911, A. Busck (1 6, 1 2: ANSP): Paraiso, 3—7 Feb 1911, A. Busck (1 6, 2 2; ANSP); Plantation “‘Borracho,”’ 10 Jul 1918, H. E Dietz, J. Zetek (1 2; USNM). PARAGUAY. Asuncion, May 1905, Ve- zéenyi (1 (dsn21 23 AINSP)=) Villaricay Jul 1937;:B. Schade (ud 337925 UUSNM): PERU. Madre de Dios: Manu, Rio Manu, near Romero, 8 Sep 1988, W. N. Mathis (1 6, 3 2; USNM). PUERTO RICO. Aguadilla, Jan 1899, A. Busck (2 2; USNM). Arecibo (beach; 18°28.7'N, 66°42'W), 23 Sep 1995, D. and W. N. Mathis (2 2; USNM). Arroyo, Feb 1899, A. Busck (1 2; USNM). Guanica (18°N, 66°54'W; Santa Rita), 12 May 1965, S. M. Gaud (5 2; USNM). Henry Bks, 19 Nov 1952, FE S. Blanton (2 2; USNM). Ma- yaguez, 21-23 Jun 1915 (1 2; ANSP). Pla- ya de Guayanilla (18°0.4'N, 66°46.1’W), 19 Sep 1995, D. and W. N. Mathis (2 &; USNM). Utuado, Jan 1899, A. Busck (1 @; USNM). Yabucoa (18°03'N, 65°52'W), 14 May 1965, S. M. Gaud (1 2°; USNM). ST. VINCENT. St. Andrew: Buccament Bay (near beach; 13°11'N, 61°16’W), 8 Jun 262 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1991, D. and W. N. Mathis (1 2; USNM). St. Patrick: Cumberland Bay (near beach; 13°16'N, 61°16’W), 28 Mar—10 Jun 1989, 1991, D. and W. N. Mathis (2 6, 2 &; USNM). TOBAGO. St. John: Charlotteville (beach; 11°19.5’N, 60°32.9'W), 16-18 Apr 1994, D. and W. N. Mathis (1 ¢6; USNM). Charlotteville (5 km S; 11°19'N, 60°34'W), Hermitage River and beach, 22 Apr—10 Jun 1993, 1994, W. N. Mathis (2 6, 5 Q; USNM); Parlatuvier (creek; 11°17.9'N, 60°35’W), 20 Apr 1994, W. N. Mathis (2 3,4 2; USNM); Speyside (1 km NW; Doc- tor River; 11°18'’N, 60°32'W), 12-13 Jun 1993, W. N. Mathis (3 ¢, 4 2; USNM). Sz. Paul: Argyle Falls (11°15'N, 60°35’W), 21 Apr 1994, W. N. Mathis (5 6, 4 2; USNM); Delaford, Kings Bay (11°16'N, 60°32.8’W), 13 Jun 1993, W. N. Mathis (2 3,4 2; USNM); Roxborough (6 km NNW; 11°16’N, 60°35.4'W), 20 Apr 1994, W. N. Mathis (7 2; USNM). TRINIDAD. St. Andrew: Valencia (1 km W; 10°39’N, 61°13'W), Aripo River, 20 Jun 1993, W. N. Mathis (6 6, 3 2; USNM). VENEZUELA. Aragua: Maracay (NW at km 16), 14 Mar 1982, G. FE and J. EK Hevel (6 6,9 2; USNM). Bolivar: El Miamo (10 km SW), 20 Mar 1982, G. E and J. E Hevel (1 2; USNM); Guasipati (7.6 km SE), 22 Mar 1982.) G) Buand) J. Bevel "@''\c; USNM); Upata (19 km SE), 20 Mar 1982, G. E and J. EF Hevel (2 2; USNM). La Pied- rita, 16 Nov 1911, S. Brown (1 2; ANSP). Distribution (Fig. 16).—Nearctic: USA (AL?, TX). Neotropical: Bahamas, Belize, Brazil (Distrito Federal, Rio de Janeiro), Colombia (Magdalena), Costa Rica (Ala- juela, Cartago, Guanacaste, Limon, Puntar- enas, San José), El Salvador, Guyana, Hon- duras, Mexico (Chiapas, Jalisco, Tabasco, Veracruz), Panama, Paraguay, Peru, Trini- dad and Tobago, Venezuela, West Indies (Cuba, Dominica, Dominican Republic, Grenada, Haiti, Jamaica, Montserrat, Puerto Rico, St. Vincent). Remarks.—We found considerable intra- specific variation in the degree of micro- sculpturing of the mesonotum and in the convexity of the scutellum of this species. Cresson (1946: 159-162) used these char- acters to distinguish between C. aciculata and his C. nitidifrons. Our attempts to use these characters proved futile, given the variation, and when we examined structures of the male terminalia, we quickly discov- ered that these two species are essentially identical. Thus, our observations reveal that there is intraspecific variation in the mi- crosculpturing and scutellar convexity but that structures of the male terminalia are consistent. Cressonomyia aciculata and C. nitidifrons are conspecific, and their names are synonyms with the former being senior. Although primarily occurring in the Neo- tropical Region, we have examined speci- mens of this species from southern Texas. Cresson (1942) reported a specimen from Alabama, which we have not confirmed and thus did not include on the distribution map of this species (Fig. 16). Cressonomyia bolivia Mathis and Zatwarnicki, new species (Figs. 17—30) Description.—This species is distin- guished from congeners, especially of the aciculata group, by the following combina- tion of characters: Small to moderately small shore flies, body length 1.50—1.90 mm. Head: Fronto-orbital setae minute. Aris- ta bearing 8 dorsal rays. Face and frons in lateral view only slightly arched, almost straight; face bare, shiny. Thorax: Mesonotum microsculptured, subopaque, granulose, or aciculate. Wing with costal vein ratio 0.97—1.05; M vein ra- tio 0.62—0.67. Abdomen: Male terminalia (Figs. 17— 30): Epandrium broadly U-shaped in pos- terior view (Fig. 17), much wider ventro- laterally than dorsally; cerci almost rodlike (Fig. 17), more rounded laterally, medial margin nearly straight; presurstylus in pos- terior view (Fig. 17) incised medially, form- ing subbasal notch, 3 setulae in notch along basomedial portion, thereafter ventrally swollen medially to slightly less than width at base then tapered and recurved to pointed VOLUME 106, NUMBER 2 263 Figs. 17-20. Structures of the male terminalia of Cressonomyia bolivia (Bolivia. La Paz: Tajlihui, 15°40.8'S, 67°41.7'W; 590 m). 17, Epandrium, cerci, and presurstylus, posterior view. 18, Same, lateral view. 19, Aedeagus (shaded), phallapodeme, postsurstylus, pregonite, and hypandrium, ventral view. 20, Same, lateral view. 264 apex, in lateral view (Fig. 18) wider basally and slightly recurved posteriorly, very gradually tapered to blunt ventral apex; postsurstylus in lateral view (Figs. 20, 30) deeply bilobed, posterior lobe longer and wider than anterior lobe, bearing numerous setulae, slightly angulate, apex bluntly rounded; pregonite (Figs. 19—20, 27, 29) moderately long, slightly tapered, broadly rounded apically, bearing 3 well-developed setulae apically; aedeagus in ventral view (Fig. 19) with slight bulge on basal two- thirds, lateral margin of swollen portion ir- regularly wavy, thereafter apically parallel sided, apex broadly rounded, in lateral view (Figs. 20, 22) with medial margin deeply sinuous, apex irregularly truncate; phalla- podeme with keel moderately wide basally, extended portion trapezoidal in lateral view; subepandrial in lateral view (Fig. 26) ta- pered to curved, point; hypandrium in lat- eral view moderately deeply pocketlike, in ventral view subquadrate anteriorly, wider than long, anterior margin nearly flat, with wide posterolateral projections posteriorly, lacking anterior projections. Type material.—The holotype male is la- beled ““BOLIVIA. Depto. La Paz[,] Tajli- hui, 590 m[,] 15°40.8'S 67°41.7’W[,] 12-i1i- 2001 [12 Mar 2001], S.D. Gaimari/HO- LOTYPE Cressonomyia bolivia ¢ W.N. Mathis USNM & Zatwarnicki [red; species name and “‘& Zatwarnicki’’ handwritten].”’ The holotype is double mounted (minuten in a block of plastic), is in excellent con- dition, and is deposited in the MNBL. Twenty-six paratypes (14 ¢d, 12 2; MNBL, USNM) bear the same locality label data as the holotype. Other specimens examined.—Neotropi- cal: BOLIVIA. La Paz: Guanay (15°29.8’'S, 67°52.7'W; 460 m), 13 Mar 2001, W. N. Mathis (1 6; USNM); Guanay (3 km E; 15°30.2'S, 67°52.3'W; 500 m), 14 Mar 2001, W. N. Mathis (2 2; USNM). Distribution (Fig. 16).—Neotropical: Bo- livia (La Paz). Etymology.—The species epithet, boli- via, alludes to the country where the type series was collected. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Remarks.—Although this species is now only known from Bolivia, we would expect that it will be found elsewhere at mid-ele- vational sites. THE HINEI GROUP Remarks.—The hinei group, which com- prises C. hinei and C. skinneri, is distin- guished from the aciculata group or meri- dionalis groups by the following combina- tion of characters (synapomorphies that characterize the meridionalis group and es- tablish its monophyly are indicated by an *): (1) fronto-orbital setae well developed; (2*) face shorter, height 1.5 that of frons; vestiture of face and frons variable; (3*) face microsculptured or with vertical micro- tomentose stripe; (4) darkened base of wing less extensive, at most occupying basal 4 of cell cua,; and (5*) postsurstylus with a posterior lobe that is elongate, narrow, spur- like (Figs. 43, 57). Cressonomyia hinei (Cresson) (Figs. 31—45) Plagiops hinei Cresson 1922: 135. Plagiopsis hinei: Cresson 1942: 127 [ge- neric combination]; 1944: 162 [correc- tion of lapsus calami in 1942: 127]; 1946: 159 [review, Neotropical fauna]. Psilopa hinei: Cresson 1942: 127 [generic combination; lapsus calami]. Cressonomyia hinei: Wirth 1965: 742 [Ne- arctic catalog; generic combination]. Description.—This species is distin- guished from congeners, especially of the skinneri group, by the following combina- tion of characters: Small to moderately small shore flies, body length, 1.75—2.35 mm. Head: Fronto-orbital setae well devel- oped. Arista bearing 7—8 dorsal rays. Face and frons moderately arched in lateral view; face short, height 1.5 that of frons, mod- erately microsculptured with some irides- cent greenish blue coloration. Thorax: Mesonotum moderately micro- scuptured; darkened base of wing at most occupying basal 4 of cell cua,. Wing with VOLUME 106, NUMBER 2 29 265 22 re Figs. 21-30. Structures of the male terminalia of Cressonomyia bolivia (Bolivia. La Paz: Tajlihui, 15°40.8’S, 67°41.7'W; 590 m). 21, Aedeagus and phallapodeme, ventral view. 22, Same, lateral view. 23, Hypandrium, ventral view. 24, Same, lateral view. 25, Subepandrial plate, ventral view. 26, Same, lateral view. 27, Pregonite, ventral view. 28, Postsurstylus, ventral view. 29, Pregonite, lateral view. 30, Postsurstylus, lateral view. costal vein ratio 0.96—1.10; M vein ratio 0.62-0.63. Abdomen: Male terminalia (Figs. 31— 44): Epandrium broadly U-shaped in pos- terior view (Fig. 31), rounded, moderately wider ventrolaterally than dorsally; cerci (Fig. 31) almost rodlike and parallel sided; presurstylus in posterior view (Fig. 31) in- cised medially, forming a large subbasal notch, 3 setulae in dorsal portion of notch, thereafter ventrally greatly swollen medi- ally to slightly more than width at base then 266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 31-34. Structures of the male terminalia of Cressonomyia hinei (Mexico. Tabasco: Teapa (8 km SW)). 31, Epandrium, cerci, and presurstylus, posterior view. 32, Same, lateral view. 33, Aedeagus (shaded), phalla- podeme, postsurstylus, pregonite, and hypandrium, ventral view. 34, Same, lateral view. VOLUME 106, NUMBER 2 tapered and recurved to short, pronglike apex, in lateral view (Fig. 32) wider basally, thereafter almost straight, very gradually ta- pered to blunt ventral apex; postsurstylus in lateral view (Figs. 34, 43) trilobed, medial lobe much longer and wider, spatulate on apical half, bearing numerous setulae, pos- terior margin of spatulate portion with 4—5 setulae from dentate bases, posterior lobe 4% length of medial lobe, slender, spurlike, bearing 1—2 setulae, anterior lobe very short, broadly triangular; pregonite (Figs. 33-34, 39, 42) moderately long, slightly narrowed medially, thereafter widened to broadly rounded to truncate apex, bearing 3 well-developed setulae apically; aedeagus in ventral view (Figs. 33, 35) with slight bulge on basal third, thereafter apically very slightly flared to mostly parallel sided, apex broadly truncate, in lateral view al- most parallel sided, medial margin very shallowly and widely incised at middle third; phallapodeme (Figs. 34, 35-36) with narrowly projected, parallel sided keel; sub- epandrial plate in ventral view (Fig. 40) broadly inverted V-shaped, in lateral view (Fig. 44) shallowly bilobed, longer lobe slightly tapered, pointed apically; hypan- drium in lateral view (Fig. 38) more deeply pocketlike, in ventral view (Figs. 33-37) with anterior, moderately wide, posteriorly curved and pointed processes, posterior processes oriented posteriorly. Type material.—The holotype female of Plagiops hinei Cresson is_ labeled **Pluer]t[o].Barrios. Guatemala Mch.3-14’ O05 [3-14 Mar 1905]/Holo-TYPE Plagiops HINEI E. T. Cresson Jr [maroon; species name handwritten].’’ The holotype is dou- ble mounted (glued to a narrow, paper tri- angle), is in excellent condition, and is de- posited in OHSU. Other specimens examined.—Nearctic: UNITED STATES. Florida. Alachua: 10 Mar 955, H:. V. Weems; In (1 6, 4° 9°; USNM); Gainesville, 23 Feb 1918, J. M. Aldrich (5 2°; ANSP, USNM). Broward: Fort Lauderdale, 26 Jan 1933, A. L. Melan- der (2 6,5 2; ANSP USNM): Hammond- ville (26°14’N, 80°12’W), 2 Jun 1953, M. 267 R. Wheeler (2 2; USNM). Dade: Royal Palm Park, 22 Apr 1930, A. L. Melander Cl 2>ANSP):. Hendry 77 Mec. 1955. Re A. Morse (2 2; USNM). Henry: Clewiston, 20 Jan 1938, A. L. Melander (1 6; USNM). Highlands: Highlands Hammock State Park, 31 Mar 1956, H. V. Weems, Jr. (1 6; USNM); Sebring, 25 Nov 1954, H. V. Weems, Jr. (1 36; USNM). Miami-Dade: Paradise Key, 21—28 Feb 1919, H. S. Bar- ber, E. A. Schwarz (i 6, 3° 2; ANSP. USNM). Okeechobee: Okeechobee, 25 Jun 1953 (1 2; USNM). Orange: Orlando, 18 Feb—18 Apr 1918, 1956, J. M. Aldrich, W. W. Wirth (6 2; ANSP, USNM). Osceola: Kissimmee, | Feb 1932, A. L. Melander (2 2: ANSP USNM). Palm Beach: Lake Worth, A. T. Slosson (1 2; USNM). Sara- sota: Sarasota, 2 Mar 1937 (1 2; USNM). Louisiana. Orleans: New Orleans (near), 1 Jan—23 Feb 1923, 1932, T. EK Hubbell, A. L. Melander (2 2; ANSP, USNM). Mississippi. Hancock: Bay St. Louis, 3 Sep-1958 dC). 2; USNM). South Carolina. Clarendon: Manning, 29-30 May 1914, W. Stone (1 2; ANSP). Texas. Bexar: San Antonio, 3 Apr 1942, A. L. Melander (1 2; USNM). Brazoria: Angleton, 3 Jan 1946, R. H. Beamer (1 &; USNM). Neotropical: BELIZE. Stann Creek: Pla- centia Lagoon, Rum Point, 4—5 Nov 1987, D. and W. N. Mathis (2 2; USNM). COSTA RICA. Alajuela: Cano Negro (10°53.6'N, 84°47.4’W; 20 m), 14-27 Apr 1994, K. F Flores (2 6, 3 2>,INBIO). EL SALVADOR. Santa Tecla (12 km NW), Feb 1954, W. B. Heed (1 2; USNM). MEXICO. Tabasco: Teapa (8 km SW), 6 May 1985,0W. N. Mathis (I 6, 20 &; USNM). Veracruz: Ciudad Aleman, 3 May 1985, W. N. Mathis (2 ¢6, 2 2; USNM); Tampico, 29 Dec 1908 (1 2; ANSP). PANAMA. Canal Zone: Coraza, 1 Mar 1911, A. Busck (1 2; USNM). Panama: Tocumen, 13 Feb—25 May 1952, 1953, E S. Blanton (4 2; USNM). Distribution (Fig. 45).—Nearctic: USA (FL, LA, MS, SC, TX). Neotropical: Be- lize, Costa Rica (Alajuela), El Salvador, 268 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 35—44. Structures of the male terminalia of Cressonomyia hinei (Mexico. Tabasco: Teapa (8 km SW)). 35, Aedeagus and phallapodeme, ventral view. 36, Same, lateral view. 37, Hypandrium, ventral view. 38, Same, lateral view. 39, Pregonite, ventral view. 40, Subepandrial plate, ventral view. 41, Postsurstylus, ventral view. 42, Pregonite, lateral view. 43, Postsurstylus, lateral view. 44, Subepandrial plate, lateral view. Guatemala, Mexico (Tabasco, Veracruz), tion there is primarily confined to the south- Panama. eastern United States. Remarks.—In North America, this is the The moderately microsculptured face most widespread species, but its distribu- with some iridescent greenish blue colora- VOLUME 106, NUMBER 2 100° 90° 100° 90° Fig. 45. Distribution map for Cressonomyia hinei. tion distinguishes this species most easily from congeners. Cressonomyia skinneri (Cresson) (Figs. 46-58) Psilopa skinneri Cresson 1922: 136.—Wol- cott 1936: 383 [list, Puerto Rico]. Plagiops skinneri: Cresson 1946: 159 [ge- neric combination]. Plagiopsis skinneri: Cresson 1942: 126 [ge- neric combination]; 1946: 158-159 [re- view, Neotropical fauna, Brazil, Cuba, Panama, Paraguay, Puerto Rico]. Cressonomyia skinneri: Wirth 1965: 742 [Nearctic catalog; generic combination]; 1968: 11 [Neotropical catalog].—Mathis and Zatwarnicki 1995: 35 [world cata- log]. Description.—This species is distin- guished from congeners, especially of the skinneri group, by the following combina- tion of characters: Small to moderately 269 small shore flies, body length 1.65—2.35 mm. Head: Fronto-orbital setae well devel- oped. Arista bearing 8 dorsal rays. Face and frons moderately arched in lateral view; face short, height 1.5 that of frons, with vertical, microtomentose stripe medially; facial setae in minute pits. Thorax: Mesonotum smooth, shiny; darkened base of wing at most occupying basal % of cell cua,. Wing with costal vein ratio 0.70—0.78; M vein ratio 0.64—0.7. Tib- ial apices yellowish. Abdomen: Male terminalia (Figs. 46— 57): Epandrium broadly U-shaped in pos- terior view (Fig. 46), rounded, moderately wider ventrolaterally than dorsally; cerci (Fig. 46) almost rodlike, wider ventrally, lateral margins rounded, medial margin al- most straight; presurstylus in posterior view (Fig. 46) incised medially, forming an an- gulate, moderately deep subbasal notch, 2 270 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ia skinneri (Jamaica. Clarendon: Grantham Figs. 46-49. Structures of the male terminalia of Cressonomyia : (18°09.3'N, 77°23.8'W; 340 m)). 46, Epandrium, cerci, and presurstylus, posterior view. 47, Same, lateral view. 48, Aedeagus (shaded), phallapodeme, postsurstylus, pregonite, and hypandrium, ventral view. 49, Same, lateral view. VOLUME 106, NUMBER 2 setulae in dorsal portion of notch, thereafter ventrally shallowly swollen medially to slightly less than width at base then tapered and recurved to short, blunt apex, in lateral view (Fig. 47) much wider basally, there- after almost straight, very slightly tapered to blunt apex; postsurstylus in lateral view (Figs. 49, 57) somewhat trilobed, medial lobe much longer and wider, slightly spat- ulate on apical half, bearing numerous se- tulae, posterior lobe moderately long, nar- row, spurlike, bearing 2 setulae, anterior lobe broadly triangular; pregonite (Figs. 48—49, 52) moderately long, in lateral view (Fig. 55) subquadrate, apex truncate, bear- ing 3 well-developed setulae apically; ae- deagus in ventral view (Figs. 48, 550) with basal third slightly swollen, lateral margins of swollen portion almost parallel sided, subquadrate, thereafter almost parallel sided to apex, apex broadly truncate, in lateral view (Figs. 49, 51) with medial margin widely and moderately deeply and irregu- larly incised, external margin more evenly arched; phallapodeme (Figs. 48-51) with keel wide basally, irregularly and asym- metrically projected, projection longest to- ward hypandrium; subepandrial plate in ventral view (Fig. 53) shallowly and broad- ly V-shaped, each arm wider than narrow medial portion, in lateral view (Fig. 56) subquadrate with apical corners slightly produced; hypandrium in lateral view (Fig. 49) more moderately deeply pocketlike, slightly angulate, in ventral view (Figs. 48, 50) with anterior margin symmetrically and shallowly sinuous, moderately wide anteri- orly, thereafter posteriorly more or less par- allel sided, posterior, subanteriorly with very slender, pointed posterolateral projec- tions. Type material.—The holotype female of Psilopa skinneri Cresson is labeled ‘*“Guan- tanamo Cuba/H. Skinner 10 11 714 [10 Nov 1914; “10” handwritten]/d [sic, specimen is a female]/Holo-TY PE Psilopa skinneri E. T. Cresson Jr [red; species name handwrit- ten]... The holotype is double mounted (minuten in a thin, rectangular piece of cardboard), is in fair condition (head miss- 271 ing), and is deposited in the ANSP (6346). The type locality is in the province of Guantanamo. Other specimens examined.—Neotropi- cal: ARGENTINA. Corrientes: Corrientes, 23 Feb 1927, R. C. Shannon (1 2; USNM). BRAZIL. Sdo Paulo: Mongagua (24°06'S, 46°37'W), 24 Aug 1961, N. L. H. Krauss 2) 6, 172: USNMD. CUBA. Clenfuegos: Aguada de Pasajero (22°23Nz 80°51) WW). Reb ies Give ot 2: USNM); Jardin Botdnico (Soledad; 22°7.5'N, 80°19.2'W), Jan—Feb 1927, C. T. and B. B. Brues (2 36, 6 2; USNM). Gran- ma: Cayamas, Baker (1 2; USNM). Guan- tanamo: Guantanamo, 10 Nov 1914, H. Skinner (1 @; paratype; ANSP). Havana: Havana, Baker (1 ¢; ANSP); Ojo de Agua (22°54.6’N, 82°29.1'W), 8 Dec 1994, W_N. Mathis (1 2; USNM). Pinar del Rio: Soroa (22°47.7'N, 83°W), 4-6 Dec 1994, W.N. Mathis (1 6, 1 2; USNM). DOMINICAN REPUBLIC. /ndependen- cias, Los Bolos G38 37-8°N, 71.39.27 W: 1370 m), 24 Mar 1999, W. N. Mathis (4 6, 17 2; USNM). La Vega: El Rio (9.5 km E; 19°0.9'N, 70°33.5’W; 980 m), 6-24 May 1995. 19985 W. IN. Mathis (@tvyd, 4) 9: USNM); Jarabacoa (1—2 km S; 19°06.9'N, 70°37'W; 520 m), 8-21 May 1995, 1998, W. N. Mathis (5 6, 6 2; USNM); Rio Camu (3.5 km NW La Vega; 19°13.7'N, 70°35.2'W; 100 m), 10 May 1995, W. N. Mathis (1 2; USNM). Puerto Plata: Rio Camu (14 km E Puerto Plata; 19°11.9'N, 70°37.4'W), 17 May 1995, W. N. Mathis (3 61 23)USNM): JAMAICA. Clarendon: Grantham (18°09.3'N, 77°23.8’W; 340 m), 16 Apr 2000, W. N. Mathis (4 6, 13 2; USNM). Manchester: near Clandon (18°09'N, 77°28.3'W), 8 May 1996, D. and W. N. Mathis, H. Williams (1 2°; USNM). Port- land: Berridale (18°06.5'N, 76°20’W), Rio Grande River, 25 Apr 2000, W. N. Mathis (4 ¢6, 4 2; USNM). St. Andrew: Kingston (Fresh River), 24 Feb 1969, W. W. Wirth (1 2: USNM); Mavis Bank (1.7 km E; 18°02):4! N37 72 39:5 OW: 15752m),. Yallahs River, 21-22 Apr—1 May 2000, W. N. i) | Nw PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 50-57. Structures of the male terminalia of Cressonomyia skinneri (Jamaica. Clarendon: Grantham (18°09.3'N, 77°23.8'W; 340 m)). 50, Aedeagus, phallapodeme, and hypandrium, ventral view. 51, Same, lateral view. 52, Pregonite, ventral view. 53, Subepandrial plate, ventral view. 54, Postsurstylus, ventral view. 55, Pregonite, lateral view. 56, Subepandrial plate, lateral view. 57, Postsurstylus, lateral view. Mathis (13 6, 11 2; USNM); Mavis Bank (4.3 km SE; 18°01.4'N, 76°38.1'W; 480 m), Yallahs River, 22—23 Apr 2000, W. N. Mathis (11 6, 8 2; USNM); Silver Hill Gap, (18°05 WN, 76°41 LW; ° 9209). 26 Apr 2000.) WinNi Mathis @ad ei 2; USNM); Wag Water River, 25 Feb 1969, W. W. Wirth (1 2; USNM). St. Mary: Annotto Bay (marsh), 25 Feb 1969, W. W. Wirth (3 2; USNM). MEXICO. Chiapas: Cacahoatan (7 km N), 22 Apr 1985, W. N. Mathis (1 6, 1 9; USNM); Union Juaréz, 23 Apr 1983, W.N. Mathis (1 2; USNM). Veracruz: Tampico, 25 Feb 1972, D. Miller, E Parker (1 6; USNM). VOLUME 106, NUMBER 2 110° 100° 90° 80° Fig. 58. PANAMA. Canal Zone: Balboa, Oct 1946, N. L. H. Krauss (1 2: USNM). Ta- boga Island, 26 Feb 1912, A. Busck (1 6; USNM). PUERTO RICO. Barranquitas, 5 Jun 1961, J. Maldonado (1 2; USNM). Baya- mon, Jan’ 1899, Ay Buseck Giods\7); USNM). Dorado, 22 Nov 1964, S. M. Gaud, E. Medina (1 2; USNM). El Yunque (road 191:-1;380 ft), 28: Apr-1965; S< M: Gaud (1 2; USNM). Fajardo, Feb 1899, A. Busck (1 2; USNM). Jayuya (2 km E; Rio Saliente; 18°12.8’N, 66°33.9'W), 22 Sep 1995. (D.rand W.N. «Mathis:.Qe6; dor; USNM). Maricao (4 km WNW; 18°10.7'N, 66°59.6’W), 21 Sep 1995, D. and W. N. Mathis (1 ¢6; USNM). Mayaquez, Jan 1899, A. Busck (1 2°; USNM). Rio Hoconuco (18°7.6’N, 67°2.6’W), 20 Sep 1995, D. and W. N. Mathis (3 6; USNM). Rio Piedras (18°20'N, 66°3’W), 25 Apr—2 May 1965, S. M. Gaud, E. Medina (10 3, 24 2; USNM). Distribution map for Cressonomyia skinneri. 70° 60° 50° 40° Rio Piedras, San Gerardo, 24 Jan 1965, E. Medina, P. Medina (1 2; USNM). Rio Aba- jo Forest (road 621; 18°18'N, 66°41'W), 29 Feb 1965. S. M:Gaud @ 3, 3.2; USNM). San German, 23 Dec 1962, P. and P. Span- gler (1) 2; USNM). Toa Baja, S5-Apr 1915, G. Garb (2 2; ANSP). Utuado, Jan 1899, A. Busck (3 6, 7 2; USNM). Vieques Is- land, Feb 1899, A. Busck (1 2; USNM). TRINIDAD. Caroni: Caroni (3 km W), 22 Mar 1985, G. E and J. E Hevel (3 d, 1 2; USNM); Chaguanas (near), 22 Mar 1985,0G), Byand (J.-E: Hevel (3 d. 13 2; USNM). St. Andrew: Valencia (1 km W; 10°39'N, 61°13’W), Aripo River, 20 Jun 1993, W. N. Mathis (3 ¢, 3 2; USNM). VENEZUELA. Santa Rosa: Barinas, Feb 1943, E Anduze (2 2; USNM). Distribution (Fig. 58).—WNearctic: USA (FL). Neotropical: Argentina (Corrientes), Brazil (Sao Paulo), Mexico (Chiapas, Ve- racruz), Panama, Trinidad, Venezuela, West 274 Indies (Cuba, Dominican Republic, Jamai- ca, Puerto Rico). Remarks.—The vertical, microtomen- tose, facial stripe is consistently evident but varies in its dimensions, sometimes extend- ing dorsally to the ptilinal suture and some- times becoming gradually wider basally, to- ward the oral margin. Although we have not examined any specimens of this species from the Nearctic Region, Cresson (1942) reported a speci- men from Florida. This record needs con- firmation and is not included on our distri- bution map (Fig. 58). The Brazilian (Sao Paulo. Mongagua: 24°06'S, 46°37'W) and Argentinian (Cor- rientes: 27°28’S, 58°50’W) localities are disjunct and appear to be outliers (Fig. 58), which prompted us to re-examine the spec- imens, especially the male from Brazil. We dissected this male and confirmed its iden- tification, and here suggest that the known distribution of this species represents a sam- pling artifact. Thus, we predict that with better sampling, the known distribution will include sites between southeastern Brazil and northern Argentina and northern South America. The specimen from Paraguay that Cresson (1946) listed was not located, but if accurately identified, it probably reflects the same pattern just noted. THE MERIDIONALIS GROUP Remarks.—The meridionalis group com- prises a single species, C. meridionalis, which is somewhat similar externally to species of the hinei group. Structures of the male terminalia of this group are the most derived and divergent within Cressonomyia and are the primary basis for recognizing the group. The group is distinguished from the aciculata or the hinei groups by the fol- lowing combination of characters (synapo- morphies that characterize the meridionalis group and establish its monophyly are in- dicated by an *): (1) fronto-orbital setae well developed; (2*) arista with 4—5 dorsal rays; (3*) face shorter, height 1.5 that of frons; (4*) face polished; (5) darkened base of wing less extensive, at most occupying PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON basal % of cell cua,; (6*) presurstylus elon- gate, narrow, almost parallel sided, shallow- ly curved; (7*) phallapodeme with portion that attaches to hypandrlium projected; (8*) gonite narrow; and (9*) hypandrium flat- tened in lateral view. Cressonomyia meridionalis (Cresson) (Figs. 59-73) Psilopa meridionalis Cresson 1918: 52. Plagiopsis meridionalis: Cresson 1946: 158 [generic combination, review, Neotropi- cal fauna]. Cressonomyia meridionalis: Wirth 1968: 11 [generic combination, Neotropical cata- log].—Mathis and Zatwarnicki 1995: 35 [world catalog]. Description.—This species is distin- guished from congeners, especially of the skinneri group, by the following combina- tion of characters: Small to moderately small shore flies, body length, 1.70—2.25 mm. Head: Fronto-orbital setae well devel- oped. Arista bearing 4—5 dorsal rays. Face and frons moderately arched in lateral view; face short, height 1.5 that of frons, pol- ished, shiny. Thorax: Mesonotum smooth, shiny; darkened base of wing at most occupying basal % of cell cua,. Wing with costal vein ratio 0.98—1.0; M vein ratio 0.60—0.62. Tib- iae mostly black. Abdomen: Male terminalia (Figs. 59— 72): Epandrium moderately broadly U- shaped in posterior view (Fig. 59), much wider ventrolaterally than dorsally; cerci (Fig. 59) somewhat rodlike, more rounded laterally, medial margin shallowly curved, pointed mediodorsally; presurstylus in pos- terior view sicklelike, gently curved, lack- ing medial notch and swelling, very slightly and gradually tapered to pointed apex, in lateral view as a long, narrow rod, only very slightly wider basally, apex slightly curved, almost truncate; postsurstylus in lateral view (Figs. 60, 72) longer than wide, angulate, bearing numerous setulae, at cor- ner of angle slightly wider from small, ex- VOLUME 106, NUMBER 2 275 pp ta 7 at I hes . © Figs. 59-62. Structures of the male terminalia of Cressonomyia meridionalis (Costa Rica. San José: Rio Savegre, San Gerardo de Dota (9°39.5'N, 83°51'W; 2180 m)). 59, Epandrium, cerci, and presurstylus, posterior view. 60, Same, lateral view. 61, Aedeagus (shaded), phallapodeme, postsurstylus, pregonite, and hypandrium, ventral view. 62, Same, lateral view. 276 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 65 rw 72. ee aay A 69 } Ss Figs. 63-72. Structures of the male terminalia of Cressonomyia meridionalis (Costa Rica. San José: Rio Savegre, San Gerardo de Dota (9°39.5'N, 83°51'W; 2180 m)). 63, Aedeagus and phallapodeme, ventral view. 64, Same, lateral view. 65, Hypandrium, ventral view. 66, Same, lateral view. 67, Pregonite, ventral view. 68, Subepandrial plate, ventral view. 69, Postsurstylus, ventral view. 70, Pregonite, lateral view. 71, Subepandrial plate, lateral view. 72, Postsurstylus, lateral view. > ternal lobe, apex with 2 small dentate pro- lae apically; aedeagus in ventral view (Figs. jections at each corner; pregonite (Figs.61— 61, 63) almost triangular, base much nar- 62, 67, 70) moderately long, narrow, apical rower than long, apical extension, like dor- half rodlike, parallel sided, narrowly round- sal view of a bicycle seat, apex narrowly ed apically, bearing 3 well-developed setu- truncate, in lateral view (Figs. 62, 64) with VOLUME 106, NUMBER 2 100° 90° Bios: swelling on basal third, swollen portion ex- tended medially, rounded, apical half slight- ly tapered, apex shallowly concave; phal- lapodeme (Figs. 62—64) comparatively large with wide, asymmetrically extended keel, longest extension toward hypandrium; subepandrial plate in ventral view (Fig. 68) much wider than long, each lateral portion slightly curved posteriorly, bluntly rounded, in lateral view (Fig. 71) slightly tapered to rounded apex; hypandrium in lateral view (Fig. 62) very shallowly pocketlike, some- what flattened, in ventral view (Fig. 61) suboval, lacking anterior and posterior pro- jections. Type material—The holotype male of Psilopa meridionalis Cresson is labeled "@artaco! M17 V"09 '* [7 “May * 1909] C[osta].Rica PPCalvert/3/TYPE 6126 [red; number handwritten]/TYPE Psilopa meri- dionalis ET. GCRESSON_ IR: [ted:. “LL YPE” written along left margin of label].’’ The Pia 80° Distribution map for Cressonomyia meridionalis. holotype is double mounted (minuten in a thin, rectangular piece of cardboard), is in good condition (slightly dusty), and is de- posited in the ANSP (6126). The type lo- cality, Cartago, is in the province of Car- tago. Other specimens examined.—Neotropi- cal: COSTA RICA. Cartago: Cartago, 17 May 1909, P. P. Calvert (1 6, 1 @; para- types; ANSP); El Alto, Laguna de Ochom- ogo! (9°53N, ©83°57' W3 1510-am)5 47 Tul 1909, P. P. Calvert (1 2; ANSP); Near Ti- erra Blanca (Rio Toyogres; 9°50'N, 83°55’W), 6 Apr 1910, P. P. Calvert (2 6, 2 2; ANSP); Trinidad (9°41.3'N, 83°54’W; 2530 m), 29 Jun 2001, A. Freidberg (1 d; USNM). Heredia: Santo Domingo (INBIO Parque; 9°58.4'N, 84°05.6'W), 23 Jun 2001, W. N. Mathis (2 ¢6; USNM). San José: La Caja (8 km W San José), 1930, H. Schmidt (5 6, 1 2; USNM); Rio Savegre, Cabinas de Quetzal (9°33.9'N, 83°48'W; 2,270 m), 278 7-8 Aug 2001, D. and W. N. Mathis (8 4, 2 2: USNM); Rio Savegre, San Gerardo de Dota (9°39.5'N, 83°51'W; 2,180 m), 7-8 Aug 2001, D. and W. N. Mathis (22 6, 8 2; USNM). MEXICO. San Luis de Potosi: Ciudad del Maiz (40-50 mi NW), 20 Nov 1948, E. S. Ross (1 2; USNM). Distribution (Fig. 73).—Neotropical: Costa Rica (Cartago, Heredia, San José), Mexico (San Luis de Potosi). Remarks.—The structures of the male terminalia, the presurstylus in particular, are quite remarkable and represent a major de- parture from other species in Cressono- myia. The narrow, almost parallel-sided, sickle-shaped presurstylus is quite unique, and readily identifies this species. Other- wise, however, we did not find this species to differ greatly from congeners. ACKNOWLEDGMENTS We gratefully acknowledge the assistance and cooperation of many organizations and individuals who contributed to the field work and production of this paper. To Da- vid A. Grimaldi (AMNH), Jon K. Gelhaus and Jason D. Weintraub (ANSP), Richard W. Baumann (BYU), Chen Young (CPM), Manuel Zumbado (INBio), Philip D. Per- kins (MCZ), Ruth Contreras-Lichtenberg (NMW), Peter W. Kovarik (OHSU), and their institutions, who loaned specimens, we express our sincere thanks. Hollis B. Williams provided Galatea support and produced the maps. For re- viewing a draft of this paper, we thank Ste- phen D. Gaimari and Allen L. Norrbom. We also express thanks to EK C. Thompson who advised us on nomenclatural issues and to Manuel Zumbado who converted Lambert coordinates for the Costan Rican grid sys- tem into standard latitude and longitude. In 1995, 1996, and 1998, field work on the West Indies was funded in large measure by grants from the Biodiversity Program (Bi- otic Surveys and Inventories, BSI), Nation- al Museum of Natural History, Smithsonian Institution (Lynne R. Parenti, former chair, George R. Zug, chair). Field work in Bo- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON livia was also supported by BSI. Field work on the West Indies and/or in Bolivia was greatly expedited through the able and pleasant assistance of N. Dianne Mathis, Amnon Freidberg, Stephen D. Gaimari, Hollis B. Williams, Kelvin Guerrero, Dan- iel E. Pérez-Gelabert, and Oliver S. Flint, Jr. Field work in Guyana was supported by the Smithsonian Institution’s Biological Di- versity of the Guianas Program (publication number 72; Vicki A. Funk, Director; Carol L. Kelloff, Coordinator). This is contribution number 653 from the Caribbean Coral Reef Ecosystems (CCRE), Smithsonian Institution. LITERATURE CITED Arnaud, P. H., Jr. 1958. Cressonomyia new name for Plagiopsis Cresson Preoccupied (Diptera: Ephy- dridae). Entomological News 69(1): 24. Brauer, E and J. E. von Bergenstamm. 1889. Die Zwei- fliigler des Kaiserlichen Museums zu Wien. IV. Vorarbeiten zu einer Monographie der Muscaria Schizometopa (exclusive Anthomyidae). Pars I. Denkschriften der Kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaf- tliche Classe. Wien 56(1): 69-180. Coquillett, D. W. 1900. Report on a collection of dip- terous insects from Puerto Rico. Proceedings of the United States National Museum 22(1198): 249-270. Cresson, E. T., Jr. 1918. Costa Rican Diptera collected by Philip P. Calvert, Ph.D., 1909-1910. Paper 3. A report on the Ephydridae. Transactions of the American Entomological Society 44(1): 39-68. 1922. Descriptions of new genera and species of the dipterous family Ephydridae——vV. Ento- mological News 33(5): 135-137. . 1925. 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Van Duzee, Scientific Survey of Porto Rico and Virgin Islands. Annals of the New York Academy of Science 11(1): 1-118. Gowdey, C. C. 1926. Catalogus Insectorum Jamaicen- sis. Department of Agriculture. Jamaica. Ento- mological Bulletin 4(1—2): xiv+114 pp. Hendel, E 1930. Die Ausbeute der deutschen Chaco- Expedition 1925/26. Diptera. XIX. Ephydridae. Konowia 9(2): 127-155. Hine, J. S. 1904. On Diptera of the family Ephydridae. The Ohio Naturalist 4(3): 63—65. Johnson, C. W. 1913. Insects of Florida. I. Diptera. Bulletin of the American Museum of Natural His- tory 32(3): 37-90. . 1919. A revised list of the Diptera of Jamaica. Bulletin of the American Museum of Natural His- tory 41(8): 421-449. Loew, H. 1862. Monographs of the Diptera of North America. Part 1. Smithsonian Institution, Smith- sonian Miscellaneous Collections 6(141): 1—221. . 1878. Neue nordamerikanische Ephydrinen. Zeitschrift fiir die Gesammten Naturwissenschaf- ten 3(51): 192-203. Mathis, W. N. 1986. Studies of Psilopinae (Diptera: Ephydridae), I: A revision of the shore fly genus Placopsidella Kertész. Smithsonian Contributions to Zoology 430: iv+30 pp. Mathis, W. N. and T. Zatwarnicki. 1990a. A revision of the Western Palearctic species of Athyroglossa (Diptera: Ephydridae). Transactions of the Amer- ican Entomological Society 116(1): 103-133. . 1990b. Taxonomic notes on Ephydridae (Dip- tera). Proceedings of the Biological Society of Washington 103(4): 891—906. . 1995. A world catalog of the shore flies (Dip- tera: Ephydridae). Memoirs on Entomology, In- ternational, 4: vit+423 pp. McAlpine, J. E 1981. Morphology and terminology— Adults. In McAlpine, J. FE et al., eds. Manual of Nearctic Diptera, 1: 9-63. Ottawa. [Volume 1 is Monograph 27 of Research Branch Agriculture Canada. | Osten Sacken, C. R. 1903. Record of my life work in ATS) entomology. University Press, Cambridge, Mas- sachusetts, ix+204 pp. Ragués, P. V. 1908. Dipteros Cubanos. Anales de la Academia de Ciencias de la Habana 45: 302-319. Schiner, I. R. 1868. Diptera. (Article 1). 7n (Wiiller- storf-Urbair, B. von, in charge), Reise der Oster- reichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859 unter den Befehlen des Commodore B. von Wiillerstorf-Urbair. Zoolo- gischer Theil. Karl Gerold’s Sohn, Wien. 2(1B), 388 pp. Townsend, C. H. T. 1911. Announcement of further results secured in the study of muscoid flies. An- nals of the Entomological Society of America 4(2): 127-152. Williston, S. W. 1896. On the Diptera of St. Vincent (West Indies). The Transactions of the Entomo- logical Society of London 1896: 253-446. . 1897. Diptera Brasiliana. Part IV. Kansas Uni- versity Quarterly Series A 6: 1-12. . 1908. Manual of North American Diptera. Third edition. New Haven, Connecticut, 405 pp. Wirth, W. W. 1965. Family Ephydridae, pp. 734—759. In Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson, eds. A Catalog of the Diptera of America North of Mexico. United States Department of Agriculture, Agriculture Handbook No. 276: iv+1696 pp. . 1968. 77. Family Ephydridae, pp. 1—43. Jn Papavero, N., ed. A Catalogue of the Diptera of the Americas South of the United States. Depar- tamento de Zoologia, Secretaria da Agricultura. Sao Paulo. Wolcott, G. N. 1924 [1923]. “‘Insectae Portoricensis.” A preliminary annotated check-list of the insects of Porto Rico, with descriptions of some news [sic] species. The Journal of the Department of Agriculture of Porto Rico 7(1): 1-313. . 1936. Insectae Borinquenses. A revised an- notated check-list of the insects of Puerto Rico. The Journal of Agriculture of the University of Puerto Rico 20(1): 1—600. Zatwarnicki, T. 1992. A new classification of Ephy- dridae based on phylogenetic reconstruction (Dip- tera: Cyclorrhapha). Genus 3(2): 65-119. . 1996. A new reconstruction of the origin of eremoneuran hypopygium and its implications for classification (Insecta: Diptera). Genus 7(1): 103— Wis! PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 280-287 KEY TO THE GENERA OF NEW WORLD ALYDIDAE (HEMIPTERA: HETEROPTERA) CARL W. SCHAEFER Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs CT 06269-3043, U.S.A. (e-mail: schaefer@uconnvm.uconn.edu) Abstract.—There are two subfamilies in Alydidae, Alydinae and Micrelytrinae, the latter with two tribes, Micrelytrini and Leptocorisini. The Alydinae contain two tribes, Daclerini (Old World genus Daclera Signoret only) and Alydini (the remainder of the subfamily). The Alydidae contain 53 genera, of which 19 occur in the New World exclu- sively, and two others in the Holarctic; a third is pantropical. The numbers break down as follows: Alydinae: Alydinae, 25 genera, 8 in the New World; Micrelytrinae: Micrely- trini, 22 genera, 11 in the New World; Micrelytrinae: Leptocorisini: Leptocorisidi, 4 gen- era, | in the New World; Noliphidi, 2 genera, 1 in the New World. Keys to subfamilies, tribes, genera, and subgenera of New World Alydidae are given. Key Words: keys Alydidae are a family of somewhat to greatly elongate heteropterans included with Stenocephalidae, Hyocephalidae, Rho- palidae, and Coreidae in the Coreoidea (Schaefer 1964, Henry 1997). Although Al- ydidae were once included as a subfamily of Coreidae, they are now treated as a sep- arate family (Schaefer 1964). Three of the family’s higher groups (based on Alydus Fa- bricius, Micrelytra Laporte, and Leptocori- sa Latreille) have been variously treated as subfamilies and/or as tribes within several subfamilies; the three groups have always been considered distinct, but their taxonom- ic affinities and levels have been ques- tioned. The head and thorax of Alydidae are somewhat flattened, with no bumps or ridg- es (except the thoracic spines of some Mi- crelytrinae), and usually without troughs. The family’s common sobriquet “broad- headed bugs”’ is best restricted to Alydinae, whose heads are indeed broader between Hemiptera, Heteroptera, Alydidae, Micrelytrinae, Leptocorisini, New World, the eyes than are those of other coreoids. The heads of Alydinae and of many Micre- lytrini are quite antlike, as would be those of other Micrelytrinae if their paraclypei were shorter than their clypet. A recent analysis (Schaefer 1999) con- cludes that there are two subfamilies, Aly- dinae and Micrelytrinae, the latter with two tribes, Micrelytrini and Leptocorisini. Aly- dinae contains two tribes, Daclerini (Old World genus Daclera Signoret only) and Alydini (the remainder of the subfamily). Leptocorisini is divided into two subtribes, Leptocorisidi and Noliphidi; both occur in the New World (one genus each). Characterizing the Alydinae is a feeding preference for legumes (Schaefer 1980, Schaefer and Mitchell 1983), a linear ar- rangement of the fifth-sternal trichobothria, spines on the hind femora, and the nymphs’ excellent mimicry of ants, among other fea- tures. Micrelytrinae may have spines on the humeral angles of the pronotum as well VOLUME 106, NUMBER 2 (sometimes) as on the scutellum’s apex; these spines are occasionally broken off, but their bases remain. The fifth-sternal tri- chobothria of Micrelytrinae are grouped in a triangle. More genera of Micrelytrini than of Leptocorisini have humeral and scutellar spines, and members of one group of Mi- crelytrini mimic ants both as adults and as nymphs. (Note: The humeral angles of some Alydinae are acuminate and may be extended as spines; this occurs for example in many Hyalymenus Amyot and Serville; these spines are horizontal, not vertical like those of Micrelytrinae.) Leptocorisini are quite elongate grass-feeding bugs (some are serious pests of rice in the Old World) whose more advanced members have elon- gated paraclypei (juga) which pass anteri- orly to the clypeus (tylus) and often meet in front of it. Micrelytrini in some ways seem to be in- termediate between Alydinae and Leptocor- isini (I do not mean to suggest they are in- termediate phylogenetically). The adults and nymphs of one group (the New World Trachelium Herrich-Schaeffer and some Cydamus Stal; the Old World Dulichius Stal) mimic ants, as do immature alydines; and some members of another group are nearly as elongate as Leptocorisini and probably also (Protenor Stal certainly) feed on Gramineae; at least one species of the antlike Cydamus also feeds on grasses (For- ero and Schaefer, unpublished), as does Es- peranza texana Barber (Wheeler and Henry 1984). In some nonantlike Micrelytrini the paraclypei surpass the clypeus, as they do in most Leptocorisini. And some Leptocor- isini have thoracic spines, as do some Mi- crelytrini. These similarities among the three family groups occur both in the New World and in the Old World Alydidae. I dis- cuss these differences and similarities, and their systematic significance, elsewhere (Schaefer 1972, 1999; and see 2003a). It should be noted that Bliven (1973) cre- ated a new subfamily in Alydidae. His Ar- aphinae Bliven 1973 was necessary, Bliven argued, because “‘along the one-way evo- 281 lutionary road, there was a successful com- mingling of genes between a Tollius-like form and a largid,”’ the result of which were the genera Araphe Herrich Schaeffer 1850, Japetus Distant 1883 (now Pararhaphe Henry [Henry 1988]), and Jarhaphetus Bli- ven 1956. (Note: The last two genera were synonymized with Araphe by Brailovsky [1981] with brief comment. More detailed reasons for the synonymy will be given by Schaefer and Ahmad [in preparation], in a revision of the New World Largidae.) These three genera (Araphe, Pararhaphe, and Jar- haphetus) had been included in the Largi- nae, of the pyrrhocoroid family Largidae, until the genera, as a new subfamily, Ara- phinae, were moved by Bliven (1973) to Alydidae. As Froeschner (1988: 5) points out, Bliven’s “‘changes, made with a mini- mum of discussion of critical and exact characters, have not been followed by other authors.”” The most important character, in Bliven’s view, uniting these three genera with Alydidae (in Bliven’s restricted defi- nition: see below), is the possession in both of a forewing stridulitrum and correspond- ing hind-femoral plectrum. However, these occur in only a few genera of Alydinae (and not at all in Micrelytrinae), and also occur in other groups only very distantly related to Araphinae and Alydidae (Schaefer and Pupedis 1981). The three largid genera re- moved by Bliven (1973) to Alydidae are now part of the tribe Araphini (Largidae: Larginae) (see Schaefer 2000). Bliven (1973) also removed to the Cor- eidae, without explanation, the alydid sub- families (now tribes of a single subfamily, Micrelytrinae) Leptocorisinae and Micre- lytrinae (s.s.). This action left Bliven’s Al- ydidae with two subfamilies, Alydinae and Araphinae. The Alydidae today contain 53 genera of which 19 occur in the New World exclu- sively; two others are Holarctic, and a third is pantropical: @ Alydinae: Alydini—25 genera, 8 in New World. 282 @ Micrelytrinae: Micrelytrini—22 gen- era, 11 in New World. @ Micrelytrinae: Leptocorisini: Lepto- corisidi—4 genera, | in New World. @ Micrelytrinae: Leptocorisini: Noliphi- di—2 genera, 1 in New World. The Alydinae comprise primarily Old World genera: One tribe (Daclerini, with just Daclera) occurs there only, six genera are exclusively New World, and two others (Alydus and Megalotomus Fieber) are Hol- arctic (Schaffner 1965). The New World genera seem to be mostly Neotropical: sev- eral genera are exclusively so, and a few others, although primarily Nearctic, have ranges extending into Mexico and (in one instance, Stachyocnemus, also into Cuba [Schaefer and Schaffner 1997]). The world genera, and some of the species, of Alydi- nae have been ably revised by Schaffner (1965), but the revision remains unpub- lished. The Micrelytrinae: Micrelytrini badly need revision. My scant analysis comes from several published sources, but mostly from unpublished data of Imtiaz Ahmad. Of the 22 genera (two unpublished), 11 occur only in the New World; all 11 are Neotrop- ical; of these 11, three occur also in the southwestern United States, and one (Pro- tenor) occurs north from Central America throughout the United States and into cen- tral and eastern Canada (Brailovsky and Flores 1979, Froeschner 1988, Maw et al. 2000). The Old World Acestra may represent a separate micrelytrine tribe (Ahmad, unpub- lished; Li and Zheng 1993). Of the six gen- era of Leptocorisini, one (the monotypic Lyrnessus Stal) is exclusively neotropical, and another (Stenocoris Burmeister) has 22 tropical species, only six of them in the New World (Ahmad 1965). Thus, of the 53 alydid genera, only 19 occur exclusively in the New World, and only three (Alydus, Megalotomus, Stenocor- is) have species in both the Old and the New Worlds (different species in most cas- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON es). A thorough revision of both the New World and the Old World Micrelytrini will probably change these numbers somewhat (I suspect several genera should be synon- ymized). But the fact that Alydidae—as well as its subfamilies—are basically Old World, will remain evident. The north-south division will also remain clear: most gen- era, including those with the most species, are tropical, in both the New and Old Worlds, and have little extension north. In this paper I am concerned only with New World alydids, especially with provid- ing keys to encourage more work. Interest in and collections of the family have increased recently, especially in the New World. There are separate surveys un- derway of the Alydidae (and other heter- opterans) of French Guiana, Colombia, Nic- aragua; and doubtless there are, or will be, others. The revisions of Schaffner (1965, Alydinae), Ahmad (1965, Leptocorisini), and to a more restricted degree that of Kor- milev (1953, Micrelytrini of Argentina) provide a good basis for further work, al- though only parts of the first have been published. Because of this interest, and with this basis, I provide the following keys to the genera and subgenera of New World Al- ydidae. Some of these keys are major modifica- tions of existing ones and others are new. Some existing keys are too vague or too inaccurate or too regional to be useful; I have not used these, and I have checked those keys I did use against specimens (with indicated exceptions). I hope these keys stimulate further work on these fasci- nating bugs, and I hope that work includes even better keys. KEY TO SUBFAMILIES AND TRIBES OF NEW WORLD ALYDIDAE I have used several keys to these groups, and that by J. C. Schaffner (1965) is the best. Because that key remains unpublished, I pre- sent it here, slightly modified, with gratitude to its author (see also Schaefer 1999). A character in some published keys is the pres- VOLUME 106, NUMBER 2 ence (Leptocorisini) or absence (Micre- lytrini) of a rostrum-receiving groove on the thoracic sterna. However, at least in New World Micrelytrinae, genera of both tribes have the groove, although not always on all three sterna (Schaefer, unpublished). 1. Hind femur bearing spines (except in the Asian Euthetus); trichobothria of abdominal sternum five (= visible sternum three) arranged in a row lateral or anterior to spiracle (Fig. 1) MEN Bek Send eee fe oer eet poh ag Ai Oke ous Alydinae — Hind femur without spines; trichobothria of ab- dominal sternum five arranged in a triangle pos- terior to spiracle. (Figs. 2, 3) Micrelytrinae ... 2 . Second rostral segment shorter than third and fourth together; third rostral segment more than half as long as fourth; evaporative area of meta- thoracic scent gland smooth Leptocorisini — Second rostral segment longer than third and fourth together; third rostral segment less than half as long as fourth; evaporative area ridged (may be very lightly ridged, as in Calamocoris) Micrelytrini i) KEY TO GENERA AND SUBGENERA OF NEW WORLD ALYDINAE This key is based on those of Fracker (1918) and Schaffner (1965), and examina- tion of specimens of Al/ydus (3 species), Ap- idaurus (2), Burtinus (2), Hyalymenus (4—5, both subgenera), Megalotomus (2), Neome- galotomus (2), Stachyocnemus (1), and Tol- lius (1); and on descriptions by Van Duzee (1906, 1921), Torre Bueno (1939, 1941), Brailovsky and Flores (1979), inter al. 1. Metathoracic scent gland opening reduced or notypenizeme- absent or reduced! =). 3. - A) — Metathoracic scent gland apparatus (peritreme) well developed Metathoracic scent gland peritreme absent; hind tibiae with two rows of prominent spines (Ne- arctic into Mexico and Cuba) N FS: Richy ac eeaca ee cee Stachyocnemus Stal — Metathoracic scent gland opening and peritreme reduced; without tibial spines 3. With stridulatory device (stridulitrum on ante- rior edge of corium [Fig. 4], plectrum near base of hind femur); clypeus (tylus) not extended be- yond paraclypei (juga) (Fig. 5) (western United States into Mexico Tollius Stal — Without stridulatory device; clypeus extended (Giese 6)x(Neotropical) Rms eee Apidaurus Stal 4. Hind tibia curved, with apical spine (Hyalymen- us Stal) Hind tibia straight, without apical spine 5. First antennal segment slightly longer than head; parameres of male directed towards one another, not projecting above ventral rim of genital cap- suley(Neotropical) aan aoe Hyalymenus (Hyalymenus Stal) — First antennal segment equal to or (more often) shorter than head; parameres directed dorsally and rising above capsule’s ventral rim (Nearctic and Neotropical) .... Hyalymenus (Tivarbus Stal) 6. Without stridulatory device (stridulitrum on an- terior edge of corium, plectrum near base of hind femur); posterolateral (humeral) angle of pronotum with small spine (Neotropical) ees eee Neomegalotomus Schaffner and Schaefer — With stridulatory device (Fig. 4); without hu- meral spine 7. Distance between bases of ocellar tubercles equal to or greater than distance from base of ocellar tubercle to eye (Neotropical) ..... A ee ee SS Saks Gy Sie ees Burtinus Stal — Distance between ocellar bases less than dis- tance fromibase to)eye (Aolarctic)= eae 8 8. First antennal segment longer than second (Hol- arctic) Megalotomus Fieber — First antennal segment shorter than second (Hol- ALCH GE). cee eas, nese ween Bee Alydus Fabricius Note.—Darmistidus Uhler (Uhler 1893), described from the island of St. Vincent in the Caribbean, is not included in this key. Uhler writes that the single included species, D. maculatus Uhler, is “*[c]losely related to Stachyocnemus.”’ However, Torre Bueno (1941) synonymized Darmistidus with Xen- ogenus, without remarking that the latter is in a different family (Rhopalidae) and ap- parently without seeing specimens; this syn- onymy has since been confirmed by GolIner- Scheiding (1980, 1983), and discussed by me (Schaefer 2003b). Because Darmistidus is now in the Rhopalidae (as Xenogenus), I do not include it in this key. KEY TO GENERA OF NEW WORLD MICRELYTRINAE: MICRELYTRINI This key is based in part on those of Stal (1873), Fracker (1918), Torre-Bueno (1941), Kormilev (1953), and Brailovsky and Flores (1979), as well as on descriptions and dis- cussions in Breddin (1901, 1903), Brailov- sky (1991), and Schaefer (1996). Specimens of the following genera were studied: Bac- 284 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON @) — 1 2 3 4 OTN \ nf as ant. 5 6 SNe thee Figs. 1-9. 1-3, Trichobothrial patterns of fifth abdominal sternum (lateral), diagrammatic; 0 = spiracle, x = trichobothrium. 1, Alydinae, Burtinus notatipennis (Stal). 2, Micrelytrinae: Micrelytrini Darmistus subvittatus Stal. 3, Micrelytrinae: Leptocorisini, Stenocoris (Stenocoris) sp. 4, Stridulitrum on costal margin of forewing of Tollius sp. (modified from Schaefer and Pupedis 1981), best seen at high dissecting scope magnification. 5—8, Dorsal views of anterior of heads; ant. = antenna (cut off), clyp. = clypeus (tylus), p.clyp. = paraclypeus (juga). 5, Tollius sp. 6, Apidaurus conspersus Stal. 7, Bactrodosoma parallelum Stal. 8, Darmistus subvittatus. 9, Lateral view of head (anterior to antennae) of Protenor belfragei Haglund. trodosoma (1 species), Bactrophya (1), Cal- (1953), respectively; and see discussion of amocoris (2), Cydamus (5), Darmistus (3), Eudarmistus in Schaefer (2003a). Esperanza (1), Protenor (2), and Trachelium (2). I have not seen specimens of Eudarmis- d id Ig ate : der”) angles of pronotum and on tip of scutel- tus, Bactrophyamixia, or Bactrocoris, and 1 : PAL atyat pki bes 5 s = wig i um (some species ant-mimetic) .......... 2 have relied upon the descriptions by Breddin _ with spines either on humeral angles or on scu- (1903), Brailovsky (1991), and Kormilev tellami@ot both). ae eee ee ee 3 1. With spines on posterolateral (humeral, “‘shoul- VOLUME 106, NUMBER 2 — Without spines on either humeral angles or scu- tellum 2. Metathoracic scent gland apparatus with auricle; head not elongated before eyes (distance from antennifers to apex of clypeus less than distance between eyes); head behind eyes tapering gently Neotropical) he fk pare can sh Swe uereres ote Cydamus Stal — Metathoracic scent gland apparatus without au- ricle; head elongated before eyes (distance from antennifers to apex of clypeus equal to distance between eyes); head behind eyes tapering sharp- ly, narrowed (Neotropical) eee erica stems cil & Trachelium Herrich-Schaeffer . Spine on scutellum only (Note: Calamocoris will key here in Froeschner’s [1981] key, which errs in placing a spine on the scutellum.) (south- Chom ES ss WIeExXICGO) ee |e ses - Esperanza Barber — Spine on each humeral angle only (Note: Cy- damus adspersipes Stal [Brazil] will key out here. However, the first antennal segment of Calamocoris is longer than the head, whereas that of Cydamus [all species] is shorter.) (north- ern South America) ...... Calamocoris Breddin 4. Tip of clypeus (tylus) not sharply deflected (Fig. OM eee eh Nore TRON Ore AG. Tee A es 5 — Tip of clypeus sharply deflected below tips of paraclypei (juga) (Fig. 8) 5. Small spur just medial to antennifers (Neotrop- ical) Bactrodosoma Stal — Without small spur (Neotropical) ........ Ra SaOe ie dey ced. RAISES anh. Bactrophya Breddin 6. Paraclypei in side view split (bifid) (Fig. 9) (Ne- arreine, INjojimoyteell)) 5p ee ubcu eau Protenor Stal Paraclypei not split 7. Longer than 16 mm; paraclypei acuminate dis- tally ((Miex1GO)) = ey. = Bactrophyamixia Brailovsky — Shorter than 14 mm; paraclypei rounded distally o>) 8. Body greatly elongate (width/length x 100 = 10-13) (Argentina) Bactrocoris Kormilev — Body not greatly elongate (width/length larvae: Comicntes. Prov. -Cor rientes, 19-XI-2002, A.J. Sosa & J. Dorado, 2 6; reared from tip galls on Alternanthera aquatica, Chaco Prov., Rt. 16 and road to Isla del Cerrito, 20-XI-2002, A.J. Sosa & J. Dorado, 3 6, 10 2, 8 pupae; Chaco Prov., near Resistencia, 19-IV-1961, G. Vogt, 3 2; Tucuman Prov., Ingenio San Pablo, 21-V- 2002, A.J. Sosa, 4 6, 3 2; Tucuman Prov., Rt. 380: ules. 21-V-2002. AJ. Sosa, 2°. URUGUAY: Montevideo, II-10-1962, G. Vogt, 2 2, 2 pupal exuviae, 4 larvae. PARAGUAY: Asuncion, 24-III-1960, G. Vogt, 2 pupae (d genitalia visible in one). BRAZIL: Rio Grande do Sul, Porto Al- 309 legre, G. Vogt, 6; Bahia, Salvador, 1-II- 1962, G. Vogt, 8 larvae; Santa Catarina, Ita- jai, 14-II-1962, G. Vogt, 2 larvae. Etymology.—The specific name, alter- nantherae, means “of Alternanthera.” Remarks.—Clinodiplosis alternantherae is one of five Neotropical species of the ge- nus known to have tarsal claws that are curved beyond midlength (Figs. 7, 13) in- stead of at the basal third. The other four are C. americana (Felt), C. cattleyae (Mol- liard), C. coffeae (Felt), and C. eupatorii (Felt). Of these five species only the new species and C. americana, a species reared from a dead wild fig branch found in Pan- ama (Felt 1911), show a general similarity in the male genitalia. They both have a deeply lobed hypoproct, with long and nar- row lobes, and a long, narrow aedeagus, but in C. alternantherae (Figs. 8—9) the cerci are wider and each bears several setae in- stead of only two, the hypoproct is not so deeply lobed, and the gonocoxite not so narrow as in C. americana (Fig. 12, illus- trated for the first time). Unlike in the new species, the circumfilar loops of C. ameri- cana are very irregular, with some loops of the first and third circumfila nearly surpass- ing the next distal flagellomere node in length. In addition, the claws of C. ameri- cana are bent at nearly right angles (Fig. 13) and are much more flared beyond the bend than in C. alternantherae (Fig. 7). ACKNOWLEDGMENTS We thank Jimena Dorado, Ana and Julia Faltlhauser, and M. Cristina Hernandez for their invaluable field assistance and ac- knowledge the initial discovery and rear- ings of the new gall midge in the early 1960s by the late George B. Vogt, then in the Systematic Entomology Laboratory (SEL), USDA, Washington, DC. We thank P. Malikul for making the slide prepara- tions, Lucrecia Rodriquez for computer as- sistance in making the plates, and, for their comments on drafts of the manuscript: K. M. Harris, formerly of the CAB Interna- tional Institute of Entomology (CIE), Lon- 310 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Vv vw aa vjms a v ve Me Ge ae ye v ~ ls: 1 ee Uo vai wEivyy Viv iv _ ViPv tio WY v s “ v Vv v SY v ROE u - . vJ v Aa ERENT Ys , v vv vay. do vantavy a 19 (Oe “ Figs. 14-20. Clinodiplosis alternantherae. 14, Female abdomen, seventh segment to end (dorsolateral). 15, Female tenth segment and cerci (dorsolateral). 16, Pupal head and prothoracic spiracles (ventral). 17, Pupal abdomen, sixth segment to end (dorsolateral). 18, Pupal abdomen, detail of dorsum of sixth segment (dorsolat- eral) with further detail showing pair of dorsal papillae. 19, Larval spatula and associated papillae. 20, Larval eighth and terminal segments (dorsal). VOLUME 106, NUMBER 2 don, UK, E. E. Grissell and A. L. Norrbom, Systematic Entomology laboratory, and V. C. Maia, Museu Nacional, Rio de Janeiro, Brazil. The field portion of this study was conducted under an agreement between USDA-ARS and CSIRO (Australia). LITERATURE CITED Felt, E. P 1911. Three new gall midges (Dipt.). Journal of the New York Entomological Society 19: 190— 193% 311 Gagné, R. J. 1989. The Plant-Feeding Gall Midges of North America. Cornell University Press, Ithaca, New York. xiii & 355 pp. & 4 pls. . 1994. The Gall Midges of the Neotropical Re- gion. Cornell University Press, Ithaca, New York, xv & 352 pp. McAlpine, J. E, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds. 1981. Manual of Nearctic Diptera. Vol. 1. Mono- graph 27, vi & 674 pp. Research Branch, Agri- culture Canada, PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 312-318 A NEW SPECIES OF BRACHYCERCUS CURTIS (EPHEMEROPTERA: CAENIDAE) FROM CHINA CHANG-FA ZHOU, LU SUN, AND W. P. MCCAFFERTY (CFZ) Institute of Genetic Resources, College of Life Sciences, Nanjing Normal Uni- versity, Nanjing 210097, China (e-mail: cfzhou@eyou.com); (LS, WPM) Department of Entomology, Purdue University, West Lafayette, IN 47907, U.S.A. Abstract.—Brachycercus capnicus, new species, is described from larvae and reared male adults recently collected from the southern provinces of China. The new species represents the first true record of the genus Brachycercus Curtis and subfamliy Brachy- cercinae from China, and the eighth such from the Eastern Hemisphere. Brachycercus capnicus is morphologically similar to B. gilliesi Soldan and Landa from southern Asia and B. japonicus Gose from eastern Asia; however, the larvae of B. capnicus are distin- guished by color pattern, antennal setae, and shape of the ocellar tubercles. The adults of B. capnicus are distinguished from other known Asian congeners by body size, pedicel length compared to that of the scape, color pattern, and development of vestigial abdom- inal processes. Larval habitat of the new species is typical of the genus. Key Words: The genus Brachycercus Curtis of the subfamily Brachycercinae (McCafferty and Wang 2000) of the cosmopolitan mayfly family Caenidae is distributed in the Hol- arctic and Oriental regions. Larvae of the genus are typified by their possession of ocellar tubercles and two-segmented labial palpi. Seven species have been known pre- viously from the Eastern Hemisphere (Tshernova 1952, Soldan 1986, Kluge 1991, Soldan and Landa 1991, Tojo 2001), but only one Brachycercus species has been recorded from China. It was taken from the northeastern region of that country and in- formally referred to as B. YUa by Quan et al. (2002). In the vicinity of China, B. har- risella Curtis has been recorded from Mon- golia and the Amur River Basin, and B. tub- ulatus Tshernova has been recorded from Korea and the Amur River Basin (Tsher- nova 1952, Soldan 1986, Kluge 1991, Hwang and Bae 1999). These records sug- mayflies, China, Brachycercus, new species gest that the two latter species eventually will be found in China. The Chinese species originally described as Brachycercus par- viforcipis Zhou, Gui, and Su (2000) is ac- tually a member of the genus Caenis Ste- phens (subfamily Caeninae) and was re- combined as such by Zhou and Zheng (in press). The description of a new Chinese species of Brachycercus is presented below. Brachycercus capnicus Zhou, Sun, and McCafferty, new species (Figs. 1-18) Mature larva.—Body (Fig. 1) length 3.4— 5.0 mm. Caudual filaments length 1.8—2.3 mm. General coloration pale yellow brown. Head: Coloration pale yellow; occiput and frons stained with brown centrally, without dark brown patches or black stripes poste- rior to lateral ocellar tubercles. Head cap- sule with anterolateral transverse row of long setae on either side between base of VOLUME 106, NUMBER 2 Figs. 1-5. 5, Hypopharynx. antenna and base of mandible. Lateral ocel- lar tubercles triangulate in lateral view (Fig. 2), with length slightly greater than width at base, and subequal to width of eye, with anterior margin slightly concave and with apex round and lacking long setae. Middle ocellar tubercle triangulate in dorsal view (Fig. 1) and straight in lateral view, with length ca. two-thirds that of lateral ocellar tubercles and ca. 1.5 basal width; apex round to bluntly pointed. Eyes not elevated above level of vertex (Fig. 2). Antenna (Fig. 3) pale, with pedicel 2.0 scape length; scape and pedicel without maculae; pedicel with ca. 10 setae subequal to, or longer than, one-half length of pedicel. YY. .Idy | ) A \ f.-J 7 SS is yy / —— , 4 . : = Nh) Brachycercus capnicus, larva. 1, Habitus. 2, Ocellar tubercles (lateral). 3, Antenna. 4, Labrum. Clypeus with transverse row of relatively long setae anteriorly. Labrum (Fig. 4) near- ly trapezoidal, with lateral margins slightly protruding, with medioapical margin slight- ly concave, and with relatively dense, stout setae along anterior margin. Lingua of hy- popharynx (Fig. 5) with distal margin straight; superlinguae with lateral margins moderately and roundly produced. Maxilla (Fig. 6) with galealacinia length 2.9 basal width; palp segment 2 length 1.8 that of segment 1; segment | width 1.6 that of segment 2; palp segment 2 with few long, stout setae at apex extending somewhat ba- sally along inner margin, and with only sparse, fine setae near base. Labial palp 314 (Fig. 7) with dense, long setae in lateral as- pect of segment 2. Thorax: Notal coloration yellow brown, without distinct black mac- ulae; sternal coloration pale yellow with diffuse black stain. Pronotum (Fig. 8) trap- ezoidal, darker medially, with lateral mar- gin slightly produced. Prothoracic pleura and lateral mesonotum with sparse marginal setae. Prosternum with median transverse ridge roundly curving, but not produced ventrally. Mesosternum with anterior mar- gin straight, without long, fine setae. Me- sosternum and metasternum without protru- sions. Legs pale throughout. Ratios of length of body: foreleg : midleg : hindleg, 3.1:1.0:1.5:1.5. Ratios of length of forefe- mur : tibia: tarsus : claw, 3.3:1.6:1.7:1.0. Ra- tios of length of hindfemur: tibia: tarsus: claw, 3.9:2.5:2.7:1.0. Ratio of hindfemur length to width, 4.3:1.0. Forefemur (Fig. 9) with row of very short setae in addition to relatively long setae along dorsal margin, and with row of stout setae in addition to some short setae along ventral margin; fore- tibia with row of four or five short setae along ventral margin; foretarsus with row of ca. 10 short setae along ventral margin; foretibia and tarsus also with scattered, sparse, short, fine setae on both anterior and posterior surfaces. Hindfemur (Fig. 10) with row of short setae interspersed with long setae along dorsal margin and short setae along ventral margin; hindtibia and tarsus with row of sparse, long setae along dorsal margins and row of short, stout setae along ventral margin; hindclaw (Fig. 10) adenticulate and moderatly curved, with length 4.0 basal width. Abdomen: Terga pale yellow, without distinct maculae; terga 1, 2, 7-9 with medial area somewhat dark- er; tergum 10 darkened anteriomedially. Tergum 1 with fine setae along posterior margin; tergum 7 with row of relatively dense, long setae along posterior margin; tergum 8 with much fewer setae along pos- terior margin. Segment 2 without short fin- gerlike process near base of operculate gill. Lateral processes on segments 2-8 (Fig. 11); process 2 triangulate and blunt; pro- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cesses 3—6 strongly curved dorsally, with apices bluntly pointed; processes 5 and 6 subequal in length; processes 7 and 8 short; process 9 small, triangulate, with apex acutely pointed. Tergum 9 with posterior margin straight. Sterna flat, pale yellow, with pale diffuse black stain in some indi- viduals, without noticable setae; sternum 9 (Fig. 12) with posterior margin convex. Operculate gill (Fig. 13) subquadrate, with length 1.3 width, without protruding edge at outer-posterior corner; dorsal surface yel- low brown except pale along outer and pos- terior margins, with few long setae in outer region; Y-ridge somewhat darker than ad- jacent areas, with outer branch stained with black, branching in anterior third of gill, and with few long setae; ventral surface lacking dense, short setae; inner margin with row of relatively sparse, short setae; inner-posterior corner with row of relatively dense, long setae; outer-posterior corner with row of long setae (shorter than those of inner-posterior corner), intermixed with several very short setae; outer margin with row of long setae shorter than those of pos- terior margin. Caudal filaments pale yellow, with segments in apical third with long se- tae. Male adult (in alcohol).—Body (Fig. 14) length 3.5—3.7 mm. Caudal filaments length 11.0-12.0 mm. Wing length 3.5 mm. Head: Occiput pale yellow, stained with diffuse pale brown. Stem of epicranial suture bor- dered with diffuse pale brown stain. Frons diffuse brown centrally above middle ocel- lus. Scape diffuse pale brown; pedicel pale brown, 2.3 length of scape; flagellum pale. Thorax: Pronotum pale yellow, with extensive black shading. Mesonotum and metanotum yellow brown. Metanotum with posteromedial protrusion broadly triangu- late, with apex bluntly pointed. Thoracic sterna yellow, with diffuse black stain. Prosternum with blackened median trans- verse ridge straight, not prominent ventral- ly; mesosternum with sternacostal suture not blackened; mesosternum and metaster- num without vestige of central protrusions. VOLUME 106, NUMBER 2 \ Q\\ oe Figs. 6—12. Brachycercus capnicus, larva. 6, Maxilla. 7, Labial palp. 8, Pronotum (left half, dorsolateral view). 9, Foreleg (anterior). 10, Hindleg (anterior). 11, Left lateral abdominal processes 2—8 (ventrolateral view). 12, Posterior margin of sternum 9. Legs with coxae and trochanters brown; femora without a black dorsodistal macula; foreleg (Fig. 15) with femur pale brown, with brown stripes, and with tibia and tar- sus pale; midleg and hindleg pale. Ratios of length of body: foreleg : midleg : hindleg, 2.5:2.3:1.0:1.1. Ratios of length of forefe- mur: tibia: tarsus, 1.0:2.9:1.9. Ratios of length of foretarsus segment [:I:III:1V:V, Ort 27257328 30:20. Forewing: (Figs ob6) with ratio of length to width at widest por- tion 1.9, with Sc, R1 and adjacent area pale brown; other veins pale. Abdomen: Color- ation generally pale, without maculae, with terga 3—6 somewhat palest. Segments 3—6 with distinct vestiges of larval lateral pro- cesses. Genitalia (Fig. 17) with penes lobes moderately convex laterally, and with for- ceps slightly bowed, with apex pointed. Caudal filaments pale. Female adult (in alcohol).—Body length 4.0 mm. Caudual filaments length 1.2 mm. Forewing length 4.5 mm, with ratio of length to width at widest portion 2.0. Ratio 316 Figs. 13-18. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 18 Brachycercus capnicus, larva, male imago, and egg. 13, Gill 2 (dorsal), larva. 14, Habitus, male imago. 15, Foreleg (dorsal), male imago. 16, Wing (dorsal), male imago. 17, Sternum 9 and genitalia (ventral), male imago. 18, Egg (lateral). of length of body : foreleg : midleg : hindleg, 2.2:1.0:1.3:1.4. Coloration similar to male. Egg (dissected from mature larva).— Length approximately 160 m. Shape (Fig. 18) narrow ovate, with single polar cap ca. one-third length of entire egg. Chorion (Fig. 18) with 10—15 broad costae in lateral view. Material examined.—Holotype: ¢ adult, Wei- Yuan River (23.30 N, 100.41 E), Feng- Shan Village, Jing-Gu County, Yunnan Province, China, IV-8-2001, Chang-Fa Zhou (deposited in Nanjing Normal Uni- versity, Nanjing, China). Paratypes: 17 d adults, one 2 adult, 15 d subimagos, 3 ° subimagos (reared from larvae), 18 larvae, data and deposition same as holotype; 3 d adults; two larvae, data same as holotype (deposited in the Purdue Entomological Re- VOLUME 106, NUMBER 2 search Collection, West Lafayette, Indiana, U.S.A.). Other Material: two larvae, Sang- Lang, Wang-Mu County (25.10 N, 106.06 E) Gui-Zhou Province, China, [X-15-2000, Chuan-Ren Li and Chang-Fa Zhou (depos- ited in Nanjing Normal University, Nan- jing, China). Etymology.—The specific epithet is an adjective of masculine gender latinized from the Greek word “‘kapnikos,” which means smoky. It is an allusion to the diffuse black stain on the thoracic sterna in the lar- vae and adults of the new species. Discussion.—Among Asian species, lar- vae of Brachycercus capnicus and those of B. tubulatus, B. corniger Kluge, B. japon- icus Gose, and B. gilliesi Soldan and Landa are relatively similar. Larvae of these spe- cies share the absence of thoracic sternal protrusions and a posterolateral protruding edge on the operculate gill (Fig. 13), and as such are distinguished from B. harrisella. Also, they all do not have dense, long setae along the anterior margin of mesosternum, which is a distinctive characteristic of B. petersorum Soldan and B. minutus Tsher- nova. Additional diagnostic features in lar- val B. capnicus are as follows. The lateral margin of abdominal segment 2 (Figs. |, 11) is slightly produced and does not form a proccess subequal to that of segment 3, as is the case in B. corniger. The lateral ocellar tubercle (Figs. 1—2) is triangulate in lateral view, slightly shorter than the com- bination of scape and pedicel; the middle ocellar tubercle length is ca. 1.5% its basal width, and less than that of the lateral ones by ca. one-third; whereas in B. japonicus the lateral ocellar tubercle is fingerlike in lateral view and longer than the combina- tion of scape and pedicel; the middle ocellar tubercle length is ca. 2.5 its basal width, and subequal to that of the lateral ones; and in B. tubulatus the middle ocellar tubercle is distinctively longer than the lateral ones. The pedicel (Fig. 3) has ca. 10 setae sube- qual to, or longer than one-half the length of the pedicel, and the abdominal terga (Fig. 1) lack maculae; whereas in B. gilliesi Sly the pedicel has over 20 long setae, and the abdominal terga 1, 2, and 7—9 have a pair of brownish bands (Soldan and Landa 1991): Four species of Asian Brachycercus had previously been known in the adult stage (Soldan 1986, Kluge 1991, Tojo 2001). The adults of Brachycercus capnicus can be dis- tinguished from them as follows. Male and female B. capnicus are no more than 4.0 mm in length; whereas the male of B. har- risella is more than 4.5 mm, and the female is more than 5.0 mm. The ratio of pedicel to scape length is greater than 2.0 in B. cap- nicus (Fig. 14); whereas this ratio is ca. 1.5 in B. minutus. Abdominal segment 2 lacks vestiges of larval lateral processes in B. capnicus (Fig. 14); whereas these vestiges are distinct in B. coniger. Thoracic sterna of B. capnicus are extensively stained with pale black; whereas B. japonicus lacks such shading. Brachycercus capnicus larvae were col- lected from a relatively clean-water river during the dry season when the river was 5—10 m in width. The larvae were found in association with a substrate composed of sand and silt, strewn with driftwood and leaf packs; a water depth of ca. 40 cm; and a current velocity less than 0.5 m/s. Other genera of mayflies collected in the same habitat included Ephemera Linnaeus, Po- tamanthellus Lestage, and Caenis. Subimagos and adults of B. capnicus were attracted to a collecting light in pre- dawn hours. The subimagos molted to adults while perched on stones near the at- tracting light. Many subimagos and adults of Caenis and Clypeocaenis Soldan were also collected at the same time by the same method. Similar observations for collecting alate stages of caenid mayflies were made by WPM in South Africa. ACKNOWLEDGMENTS We thank Dr. Chuan-Ren Li (Hubei Ag- ricultural College, China) for helping col- lect specimens. Scanning Electron Micros- 318 copy was conducted at the Life Science Mi- croscopy Facility, Purdue University. LITERATURE CITED Hwang, J. M. and Y. J. Bae. 1999. Systematics of the Caenidae (Ephemeroptera) in Korea. Korean Jour- nal of Entomology 29: 239-245. Kluge, N. J. 1991. Mayflies of the genus Brachycercus (Ephemeroptera, Caenidae) of the fauna USSR. Vestnik Zoologii 1991(2): 14—23. (In Russian.) McCafferty, W. P. and T.-Q. Wang. 2000. Phylogenetic systematics of the major lineages of pannote may- flies (Ephemeroptera: Pannota). Transactions of the American Entomological Society 126: 9-101. Quan, Y.-T., Y. J. Bae, J.-C. Jung, and J.-W. Lee. 2002. Ephemeroptera (Insecta) fauna of Northeast Chi- na. Insecta Koreana 19: 241—269. Soldan, T. 1986. A revision of the Caenidae with ocel- lar tubercles in the nymphal stage (Ephemerop- tera). Acta Universitatis Carolinae—Biologica (5— 6), 1982-1984: 289-362. e PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Soldén, T. and V. Landa. 1991. Two new species of Caenidae Ephemeroptera from Sri Lanka, pp. 235-243. In Alba-Tercedor, J. and A. Sanchez-Or- tega, eds. Overview and strategies of Ephemer- optera and Plecoptera. Sandhill Crane Press, Gainesville, Florida. Tojo, K. 2001. Redescription of Brachycercus japoni- cus Gose, 1980 (Ephemeroptera, Caenidae). En- tomological Science 4: 369-377. Tshernova, O. A. 1952. Mayflies (Ephemeroptera) of the Amur River Basin and adjacent waters and their role in the nutrition of Amur fishes. Trudy Amurskoj Ichtiologicheskoy Ekspedicii 1945— 1949: 230-360. (In Russian.) Zhou, C.-F, H. Gui, and C.-R. Su. 2000. The first re- cord of the genus Brachycercus in China with de- scription of a new species (Ephemeroptera: Caen- idae). Entomologia Sinica 72: 132-134. Zhou, C.-E and L.-Y. Zheng. In press. A preliminary study on the genus Caenis from mainland China, with description of a new species (Ephemeroptera: Caenidae). Entomotaxonomia. PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 319-323 A NEW SPECIES OF THE SAICINE ASSASSIN BUG GENUS CARAYONIA VILLIERS (HETEROPTERA: REDUVIIDAE) FROM INDOCHINA TADASHI ISHIKAWA AND SHUJI OKAJIMA (TI) c/o Dr. T. Yasunaga, Zoological Laboratory, Department of Science, Faculty of Education, Okayama University, Tsushimanaka 3-1-1, Okayama, 700-8530, Japan (e-mail: chu @cc.okayama-u.ac.jp); (SO) Laboratory of Insect Resources, Faculty of Agriculture, Tokyo University of Agriculture, Funako 1737, Atsugi-shi, Kanagawa, 243-0034, Japan (e-mail: okajima@nodai.ac.jp) Abstract.—Carayonia orientalis, n. sp., is described based on recently collected specimens from Vietnam, Laos, and Thailand. This new species inhabits wetlands with monocots. Key Words: Carayonia Villiers is a comparatively small assassin bug genus of the subfamily Saicinae, comprising nine species described from the Old World tropics and subtropics (Maldonado 1990, Malipatil 1990). The Asian fauna of the genus has been repre- sented only by two species, C. nitens (Mill- er) from the Philippines and C. culiciformis Usinger from Sri Lanka, while the others occur in Australia (one species) and Africa (six species). From recent fieldwork by us and our col- leagues in Indochina, we discovered an in- teresting species belonging to Carayonia. Upon examination, we concluded that it is undescribed. In this paper, we describe the species as new and provide habitus photo- graphs and illustrations, including the male genitalia. The diagnostic characters for the genus also are given. All the type material is housed in the Laboratory of Insect Resources, Faculty of Agriculture, Tokyo University of Agricul- ture, Atsugi, Japan. Carayonia Villiers Carayonia Villiers 1951: 278. Type spe- cles: Carayonia camerunensis Villiers 1951. Original designation. Insecta, Heteroptera, Reduviidae, Saicinae, Carayonia, Indochina Visayanocoris Miller 1952: 89 (type of Vi- sayanocorinae Miller 1952, synonymized by Villiers 1958). Type species: Visay- anocoris nitens Miller 1952. Original designation. Diagnosis.—Distinguished from other Saicine genera by a combination of the fol- lowing characters: Body shining; head pyr- iform, lacking dorsal transverse sulcus be- tween eyes, with conical anteoculus; pron- otum without spines at humeri; scutellar spine long; forecoxae elongate; forefemora and foretibiae with series of long, sparse, spiniform setae; foretibia apically with large, flattened, distally acute projection; tarsal segment II long; and abdomen ellip- tic. Detailed generic diagnoses and descrip- tions were provided by Villiers (1951, 1969), Miller (1952, as Visayanocoris) and Malipatil (1990, as Visayanocorinae). Remarks.—Taxonomic placement of Carayonia has been controversial. It has been put either in Saicinae (Villiers 1951, 1958, 1969; Maldonado 1990) or Visayan- ocorinae (Miller 1952, China and Miller 1959, Malipatil 1990, Cassis and Gross 1995). Wygodzinsky (1966) and Putshkov and Putshkov (1996) treated Visayanocori- 320 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON H ; i Carayonia orientalis, holotype (¢). 1, Dorsal view. 2, Lateral view. Figs. 1-2. nae as a tribe in Saicinae and placed this genus in Visayanocorini. We herein follow the Villiers and Maldonado’s opinions in the placement for convenience. The pyriform head without the dorsal transverse sulcus is a distinct feature sepa- rating this genus from the other saicine gen- era, and it is considered to be an autapo- morphy for Carayonia. The genus Warda- manocoris Malipatil is most similar to it in sharing such characters as the forefemur with long, sparse, spiniform setae, the for- etibia apically with a large, flattened, dis- tally acute projection, and tarsal segment II much longer than the remaining segments combined (Malipatil 1990). However, the reliable sister genera are currently not de- termined, and a comprehensive revision is required to ascertain the accurate systematic position. 1 | 2 Carayonia orientalis Ishikawa and Okajima, new species (Figs. 1-12) Description.—Measurements (in mm): d/ 2 (holotype in parentheses): Body length 3.30—3.50/3.65—3.90 (3.33). Head length in- cluding neck 0.72—0.74/0.73—0.80 (0.74): width across eyes 0.42—0.44/0.42—0.43 (0.44); interocular space 0.19—0.22/0.23 (0.21). Antenna length 2.93—3.55/3.00 (3.55); lengths of segments I, II, III and IV 1.03-1.13/1.06 (1.13), 0.37—0.41/0.37 (0.40), 0.62—0.73/0.57 (0.73) and 0.80—1.29/ 1.00 (1.29), respectively. Rostrum length 1.03—1.10/1.05—1.09 (1.10); lengths of seg- ments I, II and Il 0.49—0.51/0.52 (0.51), 0.23—0.26/0.23-0.24 (0.26) and 0.29-0.33/ 0.30—0.33 (0.33), respectively. Pronotum length 0.54—0.59/0.55—0.58 (0.54); width VOLUME 106, NUMBER 2 across humeri 0.69—0.70/0.71—0.73 (0.70). Hemelytron length 2.14—2.47/2.26—2.38 (2.39). Lengths of femur, tibia and tarsus of forelegs 1.14—1.22/1.21—1.23 (1.22), 1.16— 1.25/1.21-1.24 (1.25) and 0.60—0.65/0.62 (0.65); of midlegs 1.29—1.39/1.38-1.41 (1.39), 1.32—1.43/1.40—1.42 (1.43) and 0.54—0.66/0.68 (0.66); of hindlegs 1.64— 1.88/1.90-1.95 (1.88), 2.05—2.32/2.22—2.34 (2.30) and 0.63—0.71/0.66 (0.71), respective- ly. Abdomen length 1.70—1.84/1.96—1.98 CL76): Coloration (Figs. 1-2): Body generally dark brown to fuscous. Head and thorax shining. Anterior part of head, rostrum and areas above metacoxal cavities yellowish brown. Antennal segments I and II brown, except for apex of I] whitish; segment III brown, with many narrow, whitish annula- tions; segment IV brownish yellow, with many brown annulations on basal half. Legs brown; tibiae gradually pale toward apices; tarsi brownish yellow; mid- and hindcoxae yellowish brown. Hemelytron including veins brown to dark brown, except for pale subbasal parts. Abdomen dark brown, tinged with yellow posteriorly; lateroter- gites brownish yellow. Structure: Head (Fig. 3) elliptical, 1.7 times as long as width across eyes, ventro- laterally armed with 4 pairs of long spini- form setae, and with several, long, spini- form setae in ventral disc; anteoculus 0.7 times as long as postoculus, covered with short, decumbent setae; postoculus with long, sparsely distributed, suberect setae. Compound eye half as wide as interocular space in dorsal view. Antenna densely cov- ered with short reclining and suberect setae; proportion of segments I to [IV 14: 5: 9: 16. Rostrum (Fig. 3) bearing short, suberect se- tae; segment I with 2 pairs of long spini- form setae and a few slender setae; segment II with 3 pairs of long spiniform setae; seg- ment III with 2 pairs of long spiniform se- tae; proportion of segments I to III 20: 10: 13) Pronotum about 0.7 times as long as head, slightly more than 0.7 times as long 321 as humeral width, with long, sparsely dis- tributed, erect setae; posterior margin weak- ly arched; anterior lobe a little more than 0.75 times as long as posterior lobe. Scu- tellar spine (Fig. 3) about 1.3 times as long as scutellum, weakly curved upward, blunt- tipped. Foreleg (Fig. 4) densely covered with short, reclining and suberect setae; forefemur with 8 to 9 interolateral and 4 ventral, long spiniform setae (in Fig. 4, api- cal parts of former 3 setae visible); foretibia with 4 interolateral and 2 ventral, long spi- niform setae; foretarsus half as long as tibia. Mid- and hindlegs densely covered with short, decumbent and suberect setae. Hem- elytron slightly exceeding posterior apex of abdomen; veins on membrane ambiguous; venation shown as in Fig. 5. Abdomen wider than hemelytra, covered with short, decumbent setae except for an- terior part, apically rounded in male, sub- acute in female. Male genitalia: Pygophore (Figs. 9, 10) somewhat flattened dorsoventrally, with posterior process; posterior process taper- ing, slightly sinuate and apically obtuse in lateral view (Fig. 9), spatula-shaped and apically rounded in dorsal view (Fig. 10). Parameres (Figs. 11, 12) curved inward, weakly constricted dorsoventrally near mid- dle, rounded apically, covered with erect and suberect setae variable in length. Phal- lus (Fig. 6) elliptical, somewhat flattened dorsoventrally; phallotheca sclerotized on dorsum; endosoma long, membranous, with 2 rows of minutely spinulate areas. Female genitalia: Tergite VIII with rounded posterior margin (Fig. 8). Tergite IX declivous (Fig. 7). Valvifer I with round- ed margin (Figs. 7, 8); valvula I (Figs. 7, 8) large, nearly triangular, subacute at apex. Styloids (valvula HI) oblong, well visible between tergite IX and valvula I (Fig. 8). Holotyle.—d¢, Vietnam: Dambri, Bao Loc, Lam Dong Prov., 28.x11.2001, T. Ish- ikawa. Paratypes.—Vietnam: 1d, same data as for holotype; 1¢, 12, Loc Thang Ward, Bao Lam, Lam Dong Prov., 27.x11.2001, T. S22, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 3-12. Ishikawa. Laos: 1d, Vang Vieng, Vieng- tiang Prov., 1.11.2002, T. Kishimoto. Thai- land: 16, Mae Sa, 400-450 m alt., Mae Rim, Chiang Mai, 12.vili.2001, S. Nagash- ima; 12, Mae Sa, 450 m alt., Mae Rim, Chiang Mai, 18.viii.2001, light trap, T. Ish- ikawa. Distribution.—Vietnam, Laos, Thailand; this is the first record for the genus Cara- yonia from continental Asia. Etymology.—From the Latin, orientalis, referring to the type locality; an adjective. Carayonia orientalis (only long, spiniform setae shown on Figs. 3-10). 3, Head, prothorax and scutellum, lateral view. 4, Left foreleg. 5, Right hemelytron. 6, Phallus, dorsal view. 7—8, Apical part of female abdomen, lateral and posterior views. 9, Pygophore, lateral view. 10, Apical part of pygophore, dorsal view. 11— 12, Left paramere, lateral and dorsal views. Remarks.—This new species is distin- guished from other members of Carayonia by a combination of such features as the body 3.3—3.9 mm long, the head about 1.7 times as long as width across eyes, antennal segment IV much longer than segment III, rostral segment I twice as long as segment II, the venter of the forefemur with four long spiniform setae, the venter of the fore- tibia with two long spiniform setae, the scu- tellar spine 1.3 times as long as the scutel- lum, and the evenly curved parameres. VOLUME 106, NUMBER 2 This species inhabits marshes grown with monocots in Vietnam and Thailand, and was found on low parts of shrubs. A single individual was attracted to light. No other information 1s available on its biology. Key TO ASIAN SPECIES OF CARAYONIA 1. Pronotum 1.3 times as long as scutellum and Scutellar spine: COMbINed 7.6. .6 2.5 -. = C. culiciformis Usinger — Pronotum as long as scutellum and scutellar Spintex COMING Ye eyey. fecks clices, sp bes aka) fer ors 2 Scutellar spine twice as long as scutellum; ven- ter of foretibia with 4 long spiniform setae; parameres bent at middle . C. nitens (Miller) — Scutellar spine 1.3 times as long as scutellum (Fig. 3); venter of foretibia with 2 long spini- form setae (Fig. 4); parameres evenly curved mowyanGgudiie., 2) 2 es ees = s C. orientalis, n. sp. No ACKNOWLEDGMENTS We are grateful to Drs. T. Yasunaga (Okayama University), C. Weirauch (Freie Universitat Berlin) and D. Forero (Bogota, Colombia) for critically reading the manu- script with valuable comments, and Dr. T. Kishimoto (Japan Wildlife Research Cen- ter) and Mr. S. Nagashima (Okayama Uni- versity) for kindly providing the material used in this study. This study was partly supported by the Academic Frontier Co- operative Research Project, Tokyo Univer- sity of Agriculture. oS) i) ee) LITERATURE CITED Cassis, G. and G. E Gross. 1995. Hemiptera: Heter- optera (Coleorrhyncha to Cimicomorpha). /n Houston, W. W. K. and G. V. Maynard, eds. Zoo- logical Catalogue of Australia. Vol. 27.3A. CSI- RO Australia, Melbourne. xv + 506 pp. China, W. E. and N. C. E. Miller. 1959. Check-list and keys to the families and subfamilies of the He- miptera-Heteroptera. Bulletin of the British Mu- seum (Natural History). Entomology 8: 1—45. Maldonado, J. Capriles. 1990. Systematic catalogue of the Reduviidae of the World (Insecta: Heterop- tera). Caribbean Journal of Science (Special edi- tion), x + 694 pp. Malipatil, M. B. 1990. First record of Visayanocorinae (Hemiptera: Reduviidae) from Australia with de- scription of a new genus and two new species. Journal of the Australian Entomological Society 29: 31-36. Miller, N. C. E. 1952. Three new subfamilies of Red- uviidae (Hemiptera-Heteroptera). EOS 28: 85—90. Putshkov P. V. and G. V. Putshkoy. 1996. Family Red- uviidae Latreille, 1807—assassin-bug, pp. 148— 265. In Aukema, B. and C. Rieger, eds. Catalogue of the Heteroptera of the Palaearctic Region, Vol. 2. The Netherlands Entomological Society, Am- sterdam. xiv + 361 pp. Villiers, A. 1951. Sur deux Reduviidae Saicinae du Cameroun récoltés par J. Carayon. Bulletin du Musém National d’histoire Naturelle 23: 277-279. . 1958. Hémipteres Réuviides récoltés en An- gola (3e@me note). Publicacoes Culturais da Com- panhla de Diamantes Angola 38: 15—46. . 1969. Révision des Réduviidés africains. IV. Saicinae. Bulletin de |’ Institut Frangais d’ Afrique Noire (A) 31: 1186-1247. Wygodzinsky, P. 1966. A monograph of the Emesinae (Reduviidae, Hemiptera). Bulletin of American Museum of Natural History 133: 1-616. PROC, ENTOMOL. SOC. WASH. 106(2), 2004, pp. 324-338 SEXUAL DIMORPHISM AND SIZE OF AEDEAGI IN APIONID WEEVILS (COLEOPTERA: APIONIDAE) AND FLEA BEETLES (COLEOPTERA: CHRYSOMELIDAE): WHY SOME MASCULINE MALES HAVE SMALL AEDEAGI ALEXANDER S. KONSTANTINOV AND BorRIsS A. KOROTYAEV (ASK) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Insti- tution. PO, Box 37012, MRC 16%, Washington, DC 20013-7012, U.S.A. (e-mail: akonstan @sel.barc.usda.gov); (BAK) Zoological Institute, Russian Academy of Sciences, 199034, St. Petersburg, Russia Abstract.—A correlation between strongly developed sexually dimorphic external struc- tures and the size and mechanical properties of male genitalia are described and illustrated in Trichoconapion Korotyaeyv (Coleoptera: Apionidae) and Normaltica Konstantinoy (Co- leoptera: Chrysomelidae). Sexually dimorphic structures include legs and antennae in Trichoconapion and head and mouth parts in Normaltica. These sexually dimorphic char- acters in our examples are likely to function as clasping (restraining) or copulatory court- ship devices rather than male/male combat or precopulatory courtship devices. Key Words: courtship A monotypic Oriental apionid genus 771- choconapion Korotyaev with an unusually sharp sexual dimorphism in external char- acters was described without any reference to the male genitalia (Korotyaev 1985). In the type series, only a few males were pre- sent, and the content of their abdomens was partly decayed, Examination of additional material has revealed a small, membranous aedeagus with a weakly sclerotized median lobe, which was rather unexpected in a spe- cies with strongly developed external di- morphic characters manifested in various structures. The obvious disagreement be- tween the external characters and the de- sclerotized genital structures invited a search for similar trends in other beetles. The recently described West Indian flea beetle genus Normaltica Konstantinov (2002b) (Chrysomelidae) presented such an Apionidae, Chrysomelidae, sexual dimorphism, male genitalia, copulatory opportunity. The male of one of the only two known species of this genus differs dra- matically in having greatly enlarged head with all the mouthparts being extraordinari- ly long. Males and females of the other spe- cies are indistinguishable by their heads. Comparison of male genitalia of the Nor- maltica species revealed the same tendency as in the Oriental apionids. In the species without sexual dimorphism, the aedeagus was significantly larger and more complex in shape than in the one with sexual di- morphism. The difference in size was even greater when aedeagi were compared to body size. Unfortunately, the sexual behavior of Trichoconapion and Normaltica remains undocumented, so how their sexually di- morphic structures function is unknown. However, the fact that the structures under VOLUME 106, NUMBER 2 consideration are very species specific, sug- gests that they are under strong pressure of sexual selection (Eberhard 1985). Although we did not use a Statistical analyses, standardly used to establish cor- relation in comparative data, it is quite clear that smaller and mechanically weaker male genitalia occur in males with strongly de- veloped external, sexually dimorphic struc- tures in such distant groups as Curculion- oidea and Chrysomeloidea. An objectivity of this result is insured by comparing the genitalia of males with sexual dimorphism to those of males from closely related spe- cies (or genus) without sexual dimorphism. The structure and size of internal sac is not known for the species under consideration, so it is assumed that it correlates in size with the sclerotized portion of male geni- talia. Below we will describe external, sexu- ally dimorphic structures in Trichoconapion and Normaltica; propose an explanation for the observed correlation between sexual di- morphism and size of male genitalia; and propose hypotheses on the possible func- tion of these structures. Dissecting techniques and terminology for internal and external structures of these beetles follow Konstantinov (1998, 2002a) and Korotyaev et al. (2000). Illustrations were made with an AMRAY 1810 scanning electron microscope and ZEISS Stemi SV 11 stereo microscope with camera lucida at- tachment. SEXUAL DIMORPHISM IN APIONIDS AND FLEA BEETLES Sexual dimorphism is widespread among apionids. In addition to the characters ob- viously associated with oviposition (length and width of rostrum, which are sexually dimorphic in the great majority, if not in all Apionidae), males in many groups have more or less distinct legs. Males of many species in various regions of the world dif- fer from females in brighter coloration of the rostrum and dense white pubescence on the rostrum, underside of the head, sides of 325 the thorax and procoxae. Tibiae, especially protibiae, are often also more brightly col- ored in males. Femora in males are usually at least slightly thicker than in females; in species of the Palearctic genus Squamapion Bokor, the metafemur is often strongly swollen. Tibiae (usually protibiae) in males are often longer than in females and more or less inwardly (less frequently outwardly) curved apically, and meso- and metatibiae often have a variously shaped tooth (mucro) on the inner or, rarely, outer apical edge. Procoxae in some Protapion Schilsky are armed with spines. The apical part of the protibiae in Ceratapion penetrans Germar and C. basicorne Illiger are flattened, some- what twisted, and shallowly concave. Tarsi, especially in fore legs, are usually conspic- uously wider in males, except for the xe- romorphous species which lack or have re- duced sole hair brushes. In some groups lacking a tibial mucro, e.g., in several gen- era of the Palearctic tribe Ceratapiini, the first pro- or mesotarsomere is often armed with a sharp denticle on the ventral surface. The presence of mucro on male meso- and metatibiae also is very common in many Curculionidae, and in some cases of its re- duction on metatibia, for example, in sal- tatory Ceutorhynchinae, the first metatar- somere also may develop an acute dent on the ventral surface. Thus, both tibial mucro and tarsal denticles probably are used by males to hold themselves on the convex fe- male body. Apionidae have an additional structure probably for the same purpose, a denticle on the midline of the male meta- sternum (Eutrichapion Reitter) or first ven- trite (Hemitrichapion Voss); in several Asian species of Piezotrachelus Schoenherr the basal process of the first ventrite bears a pair of small denticles laterally (Koro- tyaev 1987). In many species of Apionidae, the male antennae are more or less conspicuously modified and usually more brightly colored, these changes being accompanied by alter- ation of the head proportions and structure. Antennae may become more or less strong- 1S) 26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Comparison of body and aedeagus size in Apionids. 1-3, Trichoconapion hirticorne. 1, Habitus. 2, Aedeagus in proportion to body size. 3, Aedeagus in details. 4-6, Vietapion paradoxum. 4, Habitus. Sy Ls Aedeagus in proportion to body size. 6, Aedeagus in detail. VOLUME 106, NUMBER 2 188 ym} Figs. 7-10. tatibia. 9, 10, Profemora in lateroventral view. ly elongated and covered with longer hairs (Protapion), the entire funicle may be thickened and pilose (many Ceratapion Schilsky) or the first antennomere (scape) is more or less strongly, sometimes pear- shaped swollen [Tatyanapion laticeps (Des- Legs of male, Trichoconapion hirticorne. 7, Ventral surface of protarsomeres. 8, Apex of me- brochers)]. The head capsule and basal part of the rostrum in species with strongly modified antennae become wider, the ros- trum often forming rather strong and more or less angular expansion at antennal inser- tion (Loborhynchapion Gy6rffy, Tatyana- 328 pion Legaloy, Ceratapion Schilsky); this expansion may be dentiform (Exapion Be- del, many Ceratapion) in species with mod- erately or weakly elongate antennae but with sharp difference between sexes in ros- trum length. The antennal club may be more or less strongly elongate (often in Ceratapion), sometimes similarly to what is common in the Dermestidae, apparently re- vealing a parallelism to the Cyladinae (Brentidae). Some really monstrous forms are known in the Mediterranean fauna of the genus Protapion, with strongly shortened and di- lated antennae, strongly flattened tarsi with angular or dentiform expanded tarsomeres, and inner margin of protibia produced into a long acute projection. In a recently found undescribed apionine from northeastern Turkey, usual modifications of the antennae and rostrum (resembling those in Lobo- rhynchapion) and apically outcurved protibiae are supplemented with an unusual position of the sharp tubercle (present in some Eutrichapion) at the anterior margin of the mesosternum and a unique modifi- cation of the middle legs which have dense and long erect pilosity on both femora and tibiae. In no examined species with sharply dimorphic external male characters, includ- ing the most peculiar ones, has any con- spicuous modification of the genitalic struc- tures been found. The most bizarre male ap- pearance is known for taxa from the tem- perate, predominantly Mediterranean fauna. No particular example of sexual dimor- phism has been reported from the Australo- Pacific apionids (Wanat 2001). That mani- fested by Trichoconapion hirticorne Koro- tyaev may be the most pronounced one among the known Oriental species of the family. In addition to those of Trichoconapion hirticorne, the only disproportion of the genitalic structures reported in the Apioni- dae is the small, weakly sclerotized aedea- gus in Aspidapion grisescens Korotyaev, from India and Vietnam, which is only half as long as the normally sclerotized aedea- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gus with strengthened apex (Korotyaev 1985: figs. 5, 6, 7) in Aspidapion vietna- mense Korotyaev, a species co-occurring with A. grisescens in Vietnam and of ex- actly the same body size as in A. grisescens. In contrast to Trichoconapion hirticorne, the mechanically weak aedeagus in A. gri- sescens is not accompanied by considerable development of the external secondary di- morphic structures. The fifth to seventh fu- nicular antennomeres in A. grisescens each bear one erect hair longer than the usual pubescence; the protibia is slightly incurved apically and armed with a small mucro; the metasternum has a sharp, rather high me- dian keel at the base, which is not found in any other Oriental Aspidapion Schilsky, but the meso- and metatibiae lack the mucro, which is well developed in A. vietnamense. Although not so conspicuous as in T. hir- ticorne, these external distinctions of A. gri- sescens may also be correlated with the small, poorly sclerotized aedeagus and pro- vide additional evidence in support of the hypothesized existence of the interrelation between the genitalic and external second- ary dimorphic male structures. Sexually dimorphic structures of Tricho- conapion include the antennae and front and hind legs. The male of Trichoconapion hirticorne (Figs. 1-14) can be easily sepa- rated from the female (Figs. 15—23) by the thickened and pilose antennae with en- larged and angular antennomere | (Figs. 13, 14, 20—23). The ventral side of male anten- nomeres 3 to 7 (which constitute most of the funicle) is covered with long setae, some of which are as long as two funicle antennomeres together (Fig. 14). Female antennae lack such setae (Fig. 23). The pro- femur of Trichoconapion is only slightly longer in males than in females, but in males its ventral surface bears two rows of stiff, closely placed, semierect setae (Figs. 9—11). In the female this surface of the pro- femur has two rows of thin, sparse, less erect setae (Figs. 18, 19). The latter char- acter is unique among the Oriental Apion- idae. The protibiae in the male are much VOLUME 106, NUMBER 2 Figs. 11-12. Protibia. longer than in the female, they are also strongly incurved near the apex (Figs. 12, 15, 17). The protarsi are only slightly dif- ferent in the male with setae slightly longer than in the female. The male metatibia has a unique dent on the inner-dorsal (instead of the usual inner-ventral) angle of the apex. Trichoconapion is a monotypic genus so we chose Vietapion paradoxum Korotyaev, which is not very closely related to Tricho- conapion, but similar in size and with only moderately expressed sexual dimorphism to Legs of male, Trichoconapion hirticorne. 11, Larger view of profemora, ventral surface. 12, compare their aedeagi. The median lobe of the aedeagus of Vietapion paradoxum is solid, strongly sclerotized, strongly curved near the apex with a wide abrupt hook at the apex (Fig. 5, 6). The paired apodemes, a part of the median lobe which is much weaker mechanically than the solid part of the median lobe, are not longer than % of the total length of the median lobe. The me- dian lobe of Trichoconapion (Figs. 2, 3) is mostly membranous, clearly consisting of two weakly connected parts: a ventral, flat- tened, short cylinder and a dorsal, flat at 330 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 13-14. apex, continuation of the apodemes. The apodemes are nearly as long as half of the total length of the median lobe. The aedea- gus of Trichoconapion is noticeably smaller than the aedeagus of Vietapion Korotyaevy, and the difference in size is even more sig- nificant if the solid, sclerotized parts of the aedeagi are measured in relation to the body sizen(Biagsnil2; 45.5)! Moderately developed sexual dimor- phism is also common in flea beetles. In the great majority of taxa males can be sepa- rated from females by their wide first pro- and mesotarsomeres. In some Aphthona Chevrolat and Phyllotreta Chevrolat spe- cies, males have strongly and variously modified antennomeres (Konstantinov 1998). Males of some species of Chaloenus Westwood have greatly enlarged heads with Antennae of male, Trichoconapion hirticorne. 13, Dorsal view. 14, Lateral view. the facial part longer and wider, but their mouthparts do not seem longer than those of females. The male of Normaltica obrieni Konstantinov has a similar kind of sexual dimorphism with enlarged heads, but its mouthparts are much longer. The head of the female of Normaltica obrieni has lateral parts converging below the eyes, but in the male the sides continue to be parallel to each other (Figs. 28-30). The clypeus and labrum together are as long as the facial part of the head to the dorsal margins of the eyes (Fig. 30); in the female, the clypeus and labrum reach to the middle of the eye (Fig. 29). In lateral view, male mandibles are particularly long (as long as the head capsule). The ventral view of the male head is even more bizarre (Fig. 31). The pre- mentum is long, wide at the base and in the VOLUME 106, NUMBER 2 middle (nearly as wide as % of the head width), narrowing at the apex, and deeply concave. The mentum is narrow and long, as long as the prementum. We compare Normaltica obrieni with N. iviet Konstantinov which lacks sexual di- morphism and is the only other species in the genus. Normaltica iviei is slightly smaller than N. obrieni, however its median lobe is actually larger than that of N. obri- eni (Figs. 25-27). It is also more complhi- cated in shape (Figs. 24, 26). The difference is even more impressive if the genitalia are compared in relation to the body size. Both examples of apionid and flea beetle clearly show aedeagi to be significantly smaller and generally mechanically weaker in males with strongly developed sexual di- morphism. DISCUSSION A negative correlation between sexually dimorphic characters and the size of male genitalia is spread across a wide array of structures in a variety of taxa at least among phytophagous beetles, since secondary sex characters occur in flea beetle heads and mouthparts and apionid legs and antennae. However, a search of available literature failed to find a described example of this tendency in other animals. Correlation of sexual behavior and gen- italic characters has been noted before. For example, Eberhard (1985: 82) suggested that elaborate premating courtship “‘imper- fectly”’ correlates with “relatively simple and uniform genitalia.’’ Rapid diversifica- tion of male genitalia and nongenitalic structures specialized to function in sexual contexts is also well known (Eberhard 1996). “Those nongenitalic portions of the male which consistently contact the female and become specialized for such contact should tend to evolve like genitalia and di- verge rapidly; they should also often func- tion as courtship devices to influence fe- male choice’? (Eberhard 1996: 354). It is therefore pretty obvious that dimorphic structures of Trichoconapion and Normal- 331 tica function in a sexual context and an ex- planation of negative correlation between secondary sex characters and the size of male genitalia can be found in the fact that male secondary sex characters and genitalia function similarly, at least to a degree. Our argument is based on the assumption that it is not evolutionary advantageous to invest energy into morphological structures which function redundantly. Further, we would ar- gue that sexually dimorphic characters in our examples are likely to function as clasp- ing (restraining) or copulatory courtship de- vices rather than male/male combat devices. All feasible explanations of functional significance of male sexually dimorphic structures are limited to four general pos- sibilities: male/male combat devices; pre- copulatory courtship devices; copulatory courtship; and/or clasping (restraining) de- vices. Most commonly, development of sexual dimorphism is explained as a result of male/ male competition (Medvedev and Pavlov 1987, Windsor 1987, Eberhard and Gutier- ez 1991, Zeh et al. 1992). In groups where reproductive success depends on domina- tion (e.g., elephant seals), large male body size is strongly favored by sexual contests competition (Andersson 1995). In leaf bee- tles, most known sexually dimorphic struc- tures are used in male combat (Beaman 1980, Eberhard 1981, Windsor 1987, Kon- stantinov 2002c). Leaf beetles without sex- ual dimorphism fight much less commonly (Konstantinoy, in press). Sexually dimor- phic structures used in fights include ce- phalic and thoracic horns, enlarged mandi- bles, fore and hind legs, appendages on the pronotum, etc. In our examples, the large legs and mandibles of Trichoconapion and Normaltica potentially can be used in male fights. However, that would not explain the smaller size of the genitalia in sexually di- morphic males since genitalia have no role in male/male combat. A male which won the precopulatory battle still faces obstacles provided by “‘cryptic female choice” (Eberhard 1996) during and after copula- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1 4 A Lm 14 pm Figs. 15-19. Legs of female, Trichoconapion hirticorne. 15, Apex of metatibia. 16, Ventral surface of protarsomeres. 17, Protibia in frontal view. 18, Profemora in ventral view. 19, Ventral surface of profemora. tion, and still competes with other males if tory stimuli which ensure successful insem- the female copulates with multiple males. ination. In that competition larger male genitalia Another possible explanation would be may be an advantage in providing copula- use of sexually dimorphic structures in pre- VOLUME 106, NUMBER 2 108 ym Figs. 20-23. 555 Antennae of female, Trichoconapion hirticorne. 20, Antenna in dorsal view. 21, Scape in lateral view. 22, Scape in ventral view. 23, Antenna in ventral view. copulatory courtship for recognition or/and as visual or tactile stimuli indicating ““good viability genes.”’ Andersson (1995) sug- gested that large head size in certain flies may help to impress rivals. In Normaltica a large head may be used for this reason, but it can be a base of large, widely sepa- rated mandibles. Long antennal setae and stiff setae on the profemora of Trichocon- apion can function as precopulatory tactile stimuli. However, genitalia play no role in precopulatory courtship, so no function of them can be substituted by the aforemen- tioned structures. In addition, cryptic fe- male choice and competition with other males during and after copulation would still be an obstacle to overcome. Copulatory courtship is described in many insects, including leaf beetles (Eber- hard 1994). Also well documented is that male courtship influences the male’s chanc- es for fertilization success (Eberhard and Kariko 1996). For example, in Macrohal- tica jamaicensis (F.) movements of anten- nae and legs of males during copulation are synchronized with aedeagal movements in- side the female, both stimulating the fe- male. Clearly, long antennal setae and setal 334 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 24 Figs. 24-27. f Comparison of body and aedeagus size in Normaltica. 24-25, N. obrieni. 24, Aedeagus in proportion to body size. 25, Habitus. 26—27, N. iviei. 26, Aedeagus in proportion to body size. 27, Habitus. rows on the profemora of Trichoconapion can function as courtship devices during copulation, but the other sexually dimor- phic characters of Trichoconapion and Nor- maltica do not look like obvious stimula- tors. Nevertheless, is not too difficult to imagine that the stimulating function of male genitalia can be diminished by the stimulating ability of their external sexually dimorphic structures, which may result in smaller size and more simple shape of the genitalia. An ability of males to remain mounted on females during (and sometimes after) copulation is very important for reproduc- tive success. It prevents other males from copulating with the female and assures sperm transfer to the spermatheca or insem- ination site. Copulating pairs are often at- tacked by single males in donaciine and chrysomeline leaf beetles (Konstantinov 2002c, Shimizu and Fujiyama 1986). That would probably constitute “direct male- male competition via mechanical properties of their genitalia” (Eberhard 1985: 92). A male structure which would help to main- tain control over the female is important to assure reproductive success in male-female conflict of interests (Eberhard 1996). Doc- umented episodes of this conflict include the female countering male mating attempts (Eberhard 1996), or forcefully terminating copulation (known in some _ cicindelids, Eberhard 1996). VOLUME 106, NUMBER 2 Sexual selection would favor develop- ment of characters functioning as clasping devices. Such devices are described in the waterstrider Gerris odontogaster (Zetter- stedt), for which it was shown that larger devices in the male is actually favored by sexual selection (Arnqvist 1989). Male gen- italia are also reported to be used for clasp- ing (Eberhard 1996). The male of Chryso- lina aurichalcea (Mannerheim) uses its ae- deagi to anchor on a female while being attacked by another male (Shimizu and Fu- jiyama 1986). Thornhill and Alcock (1983) suggested that several characteristics of male genitalia are particularly important for clasping: length, presence of spines and in- terlock features. The long and bent protib- iae, strongly setose profemora and antennae of Trichoconapion and the giant mouthparts of Normaltica can be used to remain mounted on top of the female and/or main- tain control over her during and after cop- ulation. This would lessen the clasping role of male genitalia, and as a result the geni- talia could evolve to become smaller and mechanically weaker. In the last two examples, sexually di- morphic, external characters and male gen- italia work together to provide stimuli and/ or maintain control over the female during copulation, so the genitalia are partly re- lieved from this function. However, that the smaller genitalia are less effective as stim- ulators is not really clear. They are undoubt- edly less effective as clasping devices. A single structure can be used for more than one function, but if we are to make a choice, we would conclude that the sexually dimorphic external characters in Trichocon- apion and Normaltica are more likely used during copulation as restraining and/or clasping devices. Further studies of sexual dimorphism can put this hypothesis to the test. Therefore, based on that reasoning, we would hypothesize that if dimorphic struc- tures are used in male combat, or precop- ulatory courtship, they will not correlate with the size of the male genitalia. Most of the above discussion, at least to ee) ee) Nn a degree, is based on an idea that larger male genitalia are more effective for assur- ing reproductive success, so if the genitalia in a related species are significantly smaller, something else is contributing to this suc- cess. Eberhard (1996) described three ex- perimental tests of the hypothesis that larger genitalia are associated with greater fertil- ization success. Test of the spider Nephila clavipes (Linnaeus) showed no relation be- tween size of the male genitalia and copu- latory success. The latter was measured by the amount of sperm that reached the sper- matheca and the frequency of rejections of males with shortened genitalia. In the cas- sidine leaf beetle Chelymorpha alternans Boheman, males with a longer aedeagal fla- gellum had greater paternity. Apparently the flagellum threads up the spermathecal duct of the female delivering sperms as close to the spermathecal receptacle as pos- sible (Eberhard 1996). The last example of rats, Rattus norvegicus (Berkenhout), in an analysis of Eberhard (1996) also suggested that the size of male genitalic structures is important in determining fertilization suc- cess. Conversely some studies of the allometry of male genitalia and their relation to the size of other male structures show contra- dictory results and produced debate (Eber- hard et al. 1998, Eberhard et al. 1999, Green 1999). Eberhard et al. (1998) found overall low allometric values of male gen- italia, even in Chelymorpha alternans men- tioned above, for which previously it was suggested that selection would favor larger genitalia. Based on that, they supported the earlier idea (Eberhard 1985: 79) that “‘male genitalia may not be consistently good in- dicators of male size’’ (Eberhard 1996: 76, 200). They have also concluded that selec- tion would favor an intermediate size of male genitalia which would fit the typical size of the opposite sex, and suggested that the hypothesis of forceful male/female con- flict cannot explain genitalic evolution. Green (1999) reanalyzed the ,data of Eber- hard et al. (1998) using different statistical 336 Figs. 28-31. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 188 ym & Head of Normaltica obrieni. 28-29, Female. 28, Ventral view. 29, Dorsolateral view. 30-31, Male. 30, Dorsolateral view. 31, Ventrolateral view: mn—mentum, pm—prementum, sbm—submentum. analyses and concluded that their results are dependent on the techniques used and therefore inconclusive. In a reply to Green (1999), Eberhard et al. (1999) insisted on the validity of their results. Unable to comment on the statistical techniques used by Eberhard et al. (1998 and 1999) and Green (1999), we strongly suggest against generalization in this issue. Based on the examples provided by Tricho- conapion and Normaltica, we would sug- gest that the results of the analyses are spe- cies specific. In groups in which clasping is important for assuring reproductive success, with strongly developed sexually dimorphic structures which work as restraining/clasp- ing devices, the genitalia are less important for clasping becoming smaller and mechan- ically weaker, possibly having low allome- tric value (Trichoconapion and Normaltica obrieni). In comparison, in groups with vi- tal clasping, but with the male genitalia as VOLUME 106, NUMBER 2 the only clasping device, the larger genitalia may be selectively advantageous with high allometric values (Vietapion paradoxum and Normaltica iviei). ACKNOWLEDGMENTS We are grateful to S. Braden and S. Whit- taker (SEM Laboratory, National Museum of Natural History, Smithsonian Institution) for assisting with scanning electron micros- copy. We are greatly thankful to W. G. Eberhard (Universidad de Costa Rica, Ciudad Universitaria, Costa Rica), S. W. laneatelte:, As ie, INorrbom, and EC. Thompson (Systematic Entomology Labo- ratory, USDA, Washington, DC) and C. L. Staines (Department of Systematic Biology, Smithsonian Institution) for reviewing this manuscript and providing valuable sugges- tions. LITERATURE CITED Andersson, M. 1995. Sexual Selection. Princeton Uni- versity Press, Princeton, 599 pp. Arnqvist, G. 1989. Sexual selection in a water strider: The function, mechanism of selection and herita- bility of a male grasping apparatus. Oikos 56: 344-350. Beaman, N. J. 1980. Dispersion, dispersal and phe- notypic variability of Xenarescus monocerus, a Neotropical rolled-leaf hispine beetle. Thesis, Fac- ulty of the Graduate School of Arts and Science of the George Washington University. September 30. 64 pp. Eberhard, W. G. 1981. The natural history of Dory- phora sp. (Coleoptera, Chrysomelidae) and the function of its sternal horn. Annals of the Ento- mological Society of America 47: 445—448. . 1985. Sexual Selection and Animal Genitalia. Harvard University Press, Cambridge, Massachu- setts, and London, England, 244 pp. . 1994. Evidence for widespread courtship dur- ing copulation in 131 species of insects and spi- ders, and implications for cryptic female choice. Evolution 48: 711-733. . 1996. Female Control: Sexual Selection by Cryptic Female Choice. Princeton University Press, Princeton. 501 pp. Eberhard, W. G. and E. E. Gutierez. 1991. Male di- morphism in beetles and earwigs and the question of developmental constraints. Evolution 45: 18— 28. Eberhard, W. G. and S. J. Kariko. 1996. Copulation behavior inside and outside the beetle Macrohal- 313i7/ tica jamaicensis (Coleoptera, Chrysomelidae). Journal of Ethology 14: 59-72. Eberhard, W. G., B. A. Huber, R. L. Rodriguez, R. D. Briceno, I. Salas, and V. Rodriguez. 1998. One size fits all? Relationships between the size and degree of variation in genitalia and other body parts in twenty species of insects and spiders. Evolution 52(2): 415-431. Eberhard, W. G., B. A. Huber, and R. L. Rodriguez. 1999. Don’t forget the biology: A reply to Green. Evolution 53(5): 1624-1627. Green, A. J. 1999. Allometry of genitalia in insects and spiders: One size does not fit all. Evolution 53(5): 1621-1624. Konstantinov, A. S. 1998. Revision of the Palearctic Species of Aphthona Chevrolat and Cladistic Classification of the Aphthonini (Coleoptera: Chrysomelidae: Alticinae). Memoirs on Entomol- ogy, International, Associated Publishers, Gaines- ville. 429 pp. . 2002a. New data on the structure of the fe- male genitalia of flea beetles (Coleoptera: Chry- somelidae). Proceedings of the Entomological So- ciety of Washington 104(1): 237-239. . 2002b. A new genus of flea beetles from the Greater Antilles (Coleoptera: Chrysomelidae). Zootaxa 124, 24 pp. . 2002c. New data on aggressive behavior of donaciines (Coleoptera, Chrysomelidae, Donaci- inae), pp. 74-79. In Yurin, V. M. et al., eds. Ad- vances in Modern Biology and Biological Edu- cation. Proceedings of the Second International Scientific Conference. November 29-30, 2002. Minsk. Minsk, Izdatel’skii centr BGU, 332 pp. (in Russian) . In press. Male combat and mating behavior of Donacia crassipes EF and other chrysomelids (Coleoptera: Chrysomelidae: Donaciinae). Jn Jo- livet, P, M. Schmidt, and J. Santiago-Blay, eds. New Developments on the Biology of Chryso- melidae. SPB Academic Publishing by, The Hague, The Netherlands. Korotyaev, B. A. 1985. New species of weevils of the subfamily Apioninae (Coleoptera, Apionidae) from Vietnam, China and India, pp. 133-156. Jn Medvedev, L. N., ed. Nasekomye Vietnama (In- sects of Vietnam), Moscow, Nauka. 182 pp. (in Russian). . 1987. Review of Asiatic species of the genus Piezotrachelus Schonh. (Coleoptera, Apionidae), pp. 80-94. In Medvedev, L. N., ed. Entomofauna Vietnama (Entomofauna of Vietnam), Moscow, Nauka. (in Russian). Korotyaev, B. A., A. S. Konstantinov, and C. W. O’Brien. 2000. A new genus of the Orobitidinae and discussion of its relationships (Coleoptera: Curculionidae). Proceedings of the Entomological Society of Washington 102(4): 929-956. 338 Medvedev, L. N. and S. I. Pavlov. 1987. Mating be- havior of leaf beetles (Coleoptera, Chrysomeli- dae). Entomologicheskoe Obozrenie 66(4): 746— 753 (in Russian). Shimizu, N. and S. Fujiyama. 1986. Aggressive en- counters between males in Chrysolina aurichal- cea. Journal of Ethology 4(1): 11-15. Thornhill, R. and J. Alcock. 1983. The Evolution of Insect Mating Systems. Harvard University Press, Cambridge, Massachusetts, 547 pp. Wanat, M. 2001. Genera of Australo-Pacific Rhadi- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nocybinae and Myrmacicelinae, with Biogeogra- phy of the Apionidae (Coleoptera: Curculionoi- dea) and Phylogeny of the Brentidae (s. lato). Mantis, Olsztyn, Poland, 432 pp. Windsor, D. M. 1987. Natural history of a subsocial tortoise beetle, Acromis sparsa Boheman (Chry- somelidae, Cassidinae) in Panama. Psyche 94(1— 2): 127-150. Zeh, D. W., J. A. Zeh, and G. Tavakilian. 1992. Sexual selection and sexual dimorphism in the harlequin beetle Acrocinus longimanus. Biotropica 24: 86— 96. PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 339-345 A NEW GENUS OF PRIMITIVE CRANE FLIES (DIPTERA: TANYDERIDAE) IN CRETACEOUS BURMESE AMBER, WITH A SUMMARY OF FOSSIL TANYDERIDS GEORGE POINAR, JR. AND ALEX E. BROWN (GP) Department of Zoology, Oregon State University, Corvallis, OR 97331, U.S.A. (e-mail: poinarg@casco.net); (AEB) 629 Euclid Avenue, Berkeley, CA 94708, U.S.A. Abstract.—A new genus and species of primitive crane flies, Dacochile microsoma Poinar and Brown (Tanyderidae) is described from Cretaceous Burmese amber. It differs from extant and extinct members of the family by the following combination of characters: small size (wing length, 2.8 mm), reduced anal lobe, hyaline wing membrane, crossvein cua-a, forming cell cua, very short vein R,,;, very long terminal maxillary palps, and mandibles. The well-developed mandibles indicate that the species obtained food by pierc- ing and sucking. A list of fossil tanyderids is presented. Key Words: ber, Cretaceous The Tanyderidae, or primitive crane flies, possess many plesiomorphic characters, es- pecially regarding the wing venation. The family consists of 12 extant genera, most in the Southern Hemisphere. The adults occur in moist terrestrial habitats while the larvae are found in shallow water or along the margins of lakes, rivers and streams, usu- ally in sediment or rotting vegetation (Al- exander 1981). Origin of the family and its relationship with other Nematocera has been of interest, and fossils from the Eo- cene and Jurassic have been reported (Table Ii). A female tanyderid discovered in Creta- ceous Burmese amber, described below in a new genus, provides insight into the mor- phology of early members of this interest- ing group. Amber from Burma occurs in lignitic seams in sandstone-limestone de- posits in the Hukawng Valley (Poinar 1992, Zherikhin and Ross 2000). Nuclear mag- netic resonance (NMR) spectra of amber samples taken from the same locality as the Dacochile n. gen., Dacochile microsoma n. sp., Tanyderidae, Burmese am- fossil indicated an araucarian (possibly Agathis) source of the amber (Lambert and Wu, unpublished research 2002). Palyno- morphs obtained from the amber beds where the fossil originated have been as- signed to the Upper Albian (~100—110 mya) of the Lower Cretaceous (Cruick- shank and Ko 2002). MATERIALS AND METHODS The amber was recut and polished in or- der to view the specimen better. The amber piece containing the fossil is rectangular in outline, measuring 7 mm in greatest length, 5 mm in greatest width and | mm in great- est depth. The specimen is well preserved and nearly complete, with only the tips of the tibiae and ovipositor missing. Obser- vations, drawings and photographs were made with a Nikon SMZ-10 stereoscopic microscope and Nikon Optiphot optical mi- croscope (with magnifications up to 650). In the following description, terminology is 340 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. Dacochile microsoma. 1, Holotype in Burmese amber. Bar = 836 pm. 2, Proboscis with well- developed mandibles (arrows). Bar = 95 pm. 3, Anal region of wing showing long setae emerging from costal vein (arrow). Note microtrichia on wing membrane. Bar = 103 pm. VOLUME 106, NUMBER 2 that presented in the ““Manual of Nearctic Diptera” (McAlpine 1981). The presence of a radius with 5 branches all ending at the wing margin, the subcosta bifurcate at its extremity, with the basal branch appearing as a crossvein ending in R,, a single anal vein reaching the margin, the costa continuing around the wing mar- gin, setae between the ommatidia and cell br (first basal cell) longer than cell bm (sec- ond basal cell) place the fossil in the family Tanyderidae Osten-Sacken 1879. Dacochile Poinar and Brown, new genus Type species: Dacochile microsoma Poinar and Brown. Description.—Body (length, 2.6 mm) and wing (length, 2.8 mm) small, anal lobe greatly reduced, anal margin slightly con- cave; wing membrane hyaline, crossvein cu a-a, forming cell cu a, vein R,,, short, man- dibles well developed. Etymology.—Daco is from the Greek ‘““dakno”’ for bite; chile is from the Greek “chilo”’ for lip; the gender is neuter. Diagnosis.—In Williams’ (1933) key to the extant and fossil Tanyderidae, D. micro- soma would align with the Baltic amber fossil Macrochile Loew in the couplet ‘““wings immaculate.’’ Whether the wings of Macrochile are completely hyaline is ques- tionable since Alexander (1931) noted vague indications of dusky spots at the end of vein Sc, on r-m and at the fork of M3 and CuA1 on M. spectrum Loew, and Po- denas (1997) noted dark areas on the wings of M. baltica Podenas. The two described species of Macrochile are large (body length from 8.2 mm to 12 mm and wing length from 4.5 mm to 8.5 mm) and have distinct anal lobes. The Rs varies from shorter to approximately the same length as R,.; in Macrochile while it is 3 times as long as R,,; in Dacochile. The extant and extinct Jurassic species of Protanyderus Handlirsch differ from Dacochile in having pictured wings with an anal lobe and in lacking the cu a-a, crossvein. The presence 341 of a distinct cu a-a, crossvein, forming a cu a cell is rare in tanyderids. However, it oc- curs in the Patagonian species Neoderus pa- tagonicus Alexander (Williams 1933). The extinct genus Praemacrochile Ka- lugina differs from Dacochile in having Sc end opposite the middle of R,,;, R, short, almost % of Sc and % of R,,, and having R,,; as long as or slightly longer than R,. Dacochile shares some characters with Nannotanyderus Ansorge 1994, namely the small size and reduced anal lobe, but the latter genus is characterized by a short Rs vein, only half as long as R,,;, a large cell bm and a rounded anal lobe (Ansorge 1994, Ansorge and Krzeminski 2002). Dacochile microsoma Poinar and Brown, new species (Figs. 1-8) Description.—Holotype female; with characters listed under the generic diagno- sis. General coloration brown; body length 2.6 mm. Head: Length: 315 wm (not in- cluding rostrum), eyes dichoptic, with short hairs arising between facets of ommatidia; antenna 17-segmented, 2.37 mm long, seg- ment. 1: 63 jum: 2: 113 4am: 3:63: pun,. 4: 283 jum, 52/214 pom, 6: 139°m, 7: E89 jum, $2145: pum, 92> 170) om 10-95 pam 1; 107 pm, 12: 107 ems 13 101 ums 142 807 jm, 15: 107 wm, 16: 107 pm, 17: 107 pm. An- tennal segments with numerous uniform se- tae; conspicuous verticils equal or exceed corresponding segments in length, apical segment as long as preceding segment, postpedicel short, equal in length to pedicel: maxillary palp 5-segmented, 850 ym long, with 5th segment 2.6 times as long as 4th, segment 1: 50 wm, 2: 132 pm, 3: 145 pm, 4: 145 wm, 5: 378 wm; head bearing dis- tinct, short rostrum, 265 jm long, with strong, pointed mandibles lacking notice- able teeth on inner surface (details shown in Figs. 2, 7). Thorax: Brown, 790 wm long: legs yellowish brown, coxa 1: 334 wm, 2: 410 pm, 3: 422 pm, trochanter 1: 113 wm, 220126: pom, 32 113) jm, femur 'T:+1:26mm; 2: 1.43 mm, 3: 1.42 mm; all tarsi and tips PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 4—S. Dacochile microsoma. 4, Flagellomere showing verticils at the base of each segment and dense setae uniformly covering each segment. Bar = 89 ym. 5, Extremely long terminal segment of maxillary palp. Bar = 48 pm. of tibiae missing; wing hyaline, all veins bearing setae, long setae on costa in anal area, wing membrane covered with micro- trichia, Sc relatively short, less than half wing length, ending before origin of R,,;; cell br over twice as long as cell bm; cross- vein m-cu absent; Cu P vein faint, arising from A, and curving toward wing margin; base of Cu A, appears as crossvein; tip of veins R,, R, and R, bent anteriorly at wing margin; anal lobe greatly reduced or pos- sibly absent, replaced with long setae; hal- ter brownish yellow, 258 pm long, covered with short setae; further details of venation VOLUME 106, NUMBER 2 Oo pas eS) Figs. 6-8. Dacochile microsoma. 6, Wing showing veins and cells referred to in this study. Bar = 627 ym. Setae and microtrichia omitted. 7, Lateral view of head. | = labrum, la = labium, Ib = labellum, m = mandible. Bar = 78 wm. 8, Lateral view of thorax. ke = katepimeron + meron, kt = katepisternum, m = metanotum, mk = metakatepisternum, p = prosternum, ps = postscutellum, s = scutellum, sc scutum. Bar = 165 wm. 344 Table 1. Fossil species of Tanyderidae. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Species Locality Dacochile microsoma, Nn. sp. Burma Macrochile baltica Podenas Macrochile spectrum Loew Nannotanyderus grimmenensis A. & K. Germany Nannotanyderus krzeminskii Ansorge Germany Praemacrochile chinensis K. & R. China Praemacrochile decipiens (Bode) Germany Praemacrochile dobbertinensis A. & K. | Germany Praemacrochile stackelbergi Kalugina Siberia Protanyderus mesozoicus Kalugina Mongolia Protanyderus senilis Kalugina Mongolia Protanyderus vetus Kalugina Mongolia shown in Fig. 6 and of thorax in Fig. 8. Abdomen: Brown, extended, 1.64 mm long, twisted unnaturally about 90° at approxi- mately *% of its length, tip of ovipositor missing. Male.—Unknown. Material examined.—Holotype female in Burmese amber, deposited in the Poinar amber collection (accession # B-D-12) maintained at Oregon State University. Etymology.—Micro is from the Greek ‘“‘mikros”’ for small and somus is from the Greek “‘soma” for body. Diagnosis.—Dacochile microsoma is one of the smallest known members of the fam- ily. Other distinguishing characters are the short antennal postpedicel, the very short R,,; and the extremely short bm cell. While members of the genus Radinoderus also have a relatively short R,,,, they lack a cu- a cell, possess an anal lobe and have pic- tured wings. Most tanyderids have the ter- minal flagellomere shorter than the penul- timate, and the terminal maxillary palp not more than twice the penultimate one (Wil- hams 1933, Alexander 1981). However the terminal and penultimate flagellomeres are equal in D. microsoma and the terminal maxillary palp is 2.6 times longer than the penultimate. Unfortunately, head structures are not noted for the other Mesozoic fossil tanyderids (Table 1). Baltic amber Baltic amber Age Reference Cretaceous present work Eocene Podenas 1997 Eocene Loew 1850 Lower Jurassic Lower Jurassic Middle Jurassic Lower Jurassic Lower Jurassic Upper Jurassic Upper Jurassic Upper Jurassic Upper Jurassic Ansorge & Krzeminski 2002 Ansorge 1994 Krzeminski & Ren 2001 Ansorge & Krzeminski 2002 Ansorge & Krzeminski 2002 Kalugina & Kovalev 1985 Kalugina 1988 Kalugina 1992 Kalugina 1992 DISCUSSION The presence of well-developed mandi- bles in D. microsoma is interesting because these structures have only been reported in the Australian genus Radinoderus Han- dlirsch and the North American Protoplasa Osten-Sacken (Downs and Colless 1967). In Dacochile, the size of the mandibles sug- gests that a piercing-sucking habit provided the major means of obtaining nourishment. The blood-sucking habit is well established in the Diptera, with each group evolving their own specialized type of mouthpart modification in order to obtain the food source (Downs 1970, Kalugina 1991). It is interesting that the mouth structures closely resemble those of the Psychodidae (Quate and Vockeroth 1981). The similarity between the wing structure of Mesozoic Tanyderidae and Psychodidae has been not- ed, and the similarity of the mouth structure reported here supports the contention that the two families are closely related and may have originated from a common stem (Crampton 1926, Krzeminski and Evenhuis 2000). A list of described fossil tanyderids is presented in Table |. Twelve fossil species in 5 genera (one extant and 4 extinct) have been described, all from Old World depos- its. The oldest members are from the Lower Jurassic and the youngest from the Eocene. VOLUME 106, NUMBER 2 Only three species have been described from amber deposits. It is interesting that some of the Upper Jurassic forms have been placed in the extant genus Protanyderus Handlirsch. Since the Jurassic fossils were identified only on the basis of wing char- acters, we feel that their placement in an extant genus is premature and should be made only after an examination of body characters. If confirmed, these would rep- resent the oldest terrestrial generic lineages, thus far limited to four insect genera found in Cretaceous Lebanese amber (Poinar and Milki 2001). ACKNOWLEDGMENTS We thank Neal Evenhuis, Jorg Ansorge, and Wieslaw Krzeminski for supplying per- tinent literature on fossil tanyderids and Roberta Poinar for comments on an early draft of the manuscript. LITERATURE CITED Alexander, C. P. 1931. Crane-flies of the Baltic amber (Diptera). Bernstein-Forschung 2: 1—135. 1981. Tanyderidae, pp. 149-151. In Mc- Alpine, J. E et al., eds. Manual of Nearctic Dip- tera. Monograph 27, Vol. 1. Agriculture Canada, Research Branch, Ottawa. Ansorge, J. 1994. Tanyderidae and Psychodidae (In- secta: Diptera) from the Lower Jurassic of north- eastern Germany. Palaéontologische Zeitschrift 68: 199-210. Ansorge, J. and W. Krzeminski. 2002. Lower Jurassic tanyderids (Diptera: Tanyderidae) from Germany. Studia Dipterologica 9: 21—29. Crampton, G. C. 1926. The external anatomy of the primitive tanyderid dipteran Macrochile spectrum Loew, preserved in Baltic amber. Bulletin of the Brooklyn Entomological Society 21: 1—14. Cruickshank, R. D. and K. Ko. 2002. Geology of an amber locality in the Hukawng Valley, northern Myanmar. Journal of Asian Earth Sciences 21: 441-445. Downs, J. A. 1970. The ecology of blood-sucking Dip- tera: An evolutionary perspective, pp. 232—258. In Fallis, A. M., ed. Ecology and Physiology of Par- asites, University of Toronto Press, Toronto. Downs, J. A. and D. H. Colless. 1967. Mouthparts of the biting and blood-sucking type in Tanyderidae and Chironomidae (Diptera). Nature 214: 1355— 1366. Kalugina, N. S. 1988. Mesozoic Diptera Psychodo- morpha and Tipulomorpha (Tanyderidae, Eop- tychopteridae, Limoniidae: Diptera), pp. 81—88. Jn Rozanoy, A. Yu., ed. New Species of Fossil In- vertebrates of Mongolia. Trudy Sovetsko-Mon- golskaia paleontologeskaia Ekspeditsiia 33. (In Russian. ) . 1991. New Mesozoic Simuliidae and Lepto- conopidae and the origin of bloodsucking in the lower dipteran insects. Paleontological Journal 25: 66-77. . 1992. Psychodomorph Dipterans from the Ju- rassic of the Mongolian Altai (Diptera: Tanyderi- dae, Eoptychopteridae). Paleontological Journal 26: 142-146. Kalugina, N. S. and V. G. Kovalev. 1985. Dipterous insects of Jurassic Siberia. Paleontological Insti- tute, Akademia Nauk, Moscow, 198 pp. (In Rus- sian.) Krzeminski, W. and N. L. Evenhuis. 2000. Review of Diptera palaeontological records, pp. 535-564. Jn Papp, L. and B. Darvas, eds. Manual of Palaearc- tic Diptera. Science Herald, Budapest, Vol. 1. Krzeminski, W. and D. Ren. 2001. Praemacrochile chinensis sp. n. from the Middle Jurassic of China (Diptera: Tanyderidae). Polskie Pismo Entomolo- gica 70: 127-129. Loew, H. 1850. Ueber den Bernstein und die Bern- steinfauna. E. S. Mittler & Sohn, Berlin, 44 pp. McAlpine, J. FE 1981. Morphology and terminology— Adults, pp. 9-63. In McAlpine, J. E et al., eds. Manual of Nearctic Diptera. Monograph 27, Vol. 1, 674 pp. Agriculture Canada, Research Branch, Ottawa. Podenas, S. 1997. New Macrochile Loew, 1850 (Dip- tera, Tanyderidae) from the Baltic amber. Mittei- lungen der Geologisch-Palaéontologischen Institut der Univeristaét Hamburg 80: 173-177. Poinar, Jr., G. O. 1992. Life in Amber. Stanford Uni- versity Press, Palo Alto, 350 pp. Poinar, Jr., G. O. and R. Milki. 2001. Lebanese amber. The oldest insect ecosystem in fossilized resin. Oregon State University Press, Corvallis, 97 pp. Quate, L. W. and J. R. Vockeroth. Psychodidae, pp. 153-190. In McAlpine, J. FE et al., eds. Manual of Nearctic Diptera. Monograph 27, Vol. 1, Agricul- ture Canada, Research Branch, Ottawa. Williams, I. 1933. The external morphology of the primitive tanyderid dipteran Protoplasa _fitchii O.S., with notes on the other Tanyderidae. Journal of the New York Entomological Society 41: 1—36. Zherikhin, V. V. and A. J. Ross. 2000. A review of the history, geology and age of Burmese amber (Bur- mite). Bulletin of the Natural History Museum (London) (Geology) 56: 3-10. PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 346-351 ALLOEORHYNCHUS TRIMACULA (STEIN) (HETEROPTERA: NABIDAE: PROSTEMMATINAE), A PREDATOR OF RHYPAROCHROMIDAE (LYGAEOIDEA) ASSOCIATED WITH FIGS IN MEXICO Luts CERVANTES PEREDO Instituto de Ecologia, A.C., Km. 2.5 Antigua Carretera a Coatepec #351, CP 91070 Xalapa, Veracruz, México (e-mail: cervantl @ecologia.edu.mx) Abstract.—The nabid Alloeorhynchus trimacula (Stein) was found preying on adults and nymphs of several species of Rhyparochromidae, including members of the tribes Antillocorini, Myodochini, and Ozophorini. All records correspond to rhyparochromids that are associated with several native species of figs in Mexico. Descriptions of the adult and immature stages and information on the biology of A. trimacula are included. Resumen.—E] nabido Alloeorhynchus trimacula (Stein) fue encontrado depredando adul- tos y ninfas de varias especies de Rhyparochromidae, incluyendo miembros de las tribus Antillocorini, Myodochini y Ozophorini. Todos los registros corresponden a rhyparochro- midos que estan asociados a especies nativas de higueras en México. Se incluyen descrip- ciones del adulto, estados inmaduros e informacion sobre la biologia de A. trimacula. Key Words: Harris (1928) gave descriptions of all spe- cies of Nabidae known from North and Cen- tral America and the West Indies. He in- cluded notes about the biology of many spe- cies and their economic importance. Within this family, members of Nabinae seem to be general predators on small arthropods. Spe- cies of the Prostemmatinae were reported to prey exclusively on other Heteroptera (Pe- ricart 1987, Lattin 1989). Carayon (1970) re- ported other species of Al/loeorhynchus Fie- ber known to prey on lygaeids in Africa, but little work on this group of nabids with its prey has ever been documented. Information about the biology and descriptions of all in- stars of Alloeorhynchus trimacula (Stein) are given in this study. A list of the species of rhyparochromid bugs and their host plants of the associated bugs, and geographical re- cords from several states in Mexico are pre- sented. Nabidae, Alloeorhynchus, Rhyparochromidae, Moraceae, Ficus MATERIALS AND METHODS Monthly collecting trips during 2001 and 2002 were made to several localities in the Mexican states of Campeche, Puebla, Ta- maulipas, and Veracruz. The objective was to collect rhyparochromids associated with fruiting fig trees. While collecting lygaeids, other insects associated with the bugs or preying on them were collected. Around 30 fig species were sampled from localities at sea level to an altitude of 1,000 m. Several types of vegetation were included: low tropical dry forest, medium tropical forest, high tropical rain forest, and cloud forest. Nabids were collected alive and put into plastic containers (9 X 8 cm) covered with muslin to avoid condensation. A dry fig leaf and a small humid cotton ball were put in each container; living rhyparochromids of several species were offered as food. Con- VOLUME 106, NUMBER 2 tainers were checked daily for the presence of eggs, and more prey added when nec- essary. Individuals were kept under labo- ratory conditions at about 20° C and 70% RH. Individuals were placed in 70% alco- hol and used for illustrations and descrip- tions. Measurements are given in mm, +SD. Material collected during 2001 and 2002 is deposited in the Insect Collection of Instituto de Ecologia, A.C. TEXA). Ad- ditional material in the Insect Collection of Instituto de Biologia, U.N.A.M. (CNIN) was studied. Alloeorhynchus trimacula (Stein) (Eas. 1) Descriptions.—Egg (Fig. 1A) (n = 3): Elongate, 0.8 + 0 mm long, 0.23 = 0.03 mm wide, anterior pole slightly bent toward one side, anterior pole with grayish chori- onic ring, posterior pole round. White when laid, turning yellowish in two to three days; red eye spots appear after day seven. First instar (Fig. 1B): Body elongated, with maximum width across abdominal segment III. Head, thorax, and a _heart- shaped macula over dorsal abdominal seg- ments IV to IX pale brown. Eyes reddish brown; antennal segments pale gray, with joints white. Rostrum and legs grayish yel- low. Thoracic pleurae reddish. Abdomen pinkish gray, except for pale brown macula on last abdominal segments. All dehiscent lines pale yellow. Head slightly declivent, with two long setae on tylus; rostrum reach- ing posterior part of procoxae. Postero-lat- eral angles of pro-, meso-, and metanotum with a long seta. Front and middle femora with thick black mid spine on ventral sur- face. Scent gland openings barely apparent on segments III-IV, IV—V, and V—VI, the first two as a pair of openings, the third as a single opening. Measurements (n = 3). Body length 1.45 + 0.14; head length 0.19 + 0.04; width across eyes 0.31 + 0.04; in- terocular distance 0.16 + 0.02; postocular distance 0.02 + 0.0; antennal segments: I Ole =- 0. HE 0102) 0, 10:17) 0:03 7-5V 0.2 + 0, V 0.39 + 0.01; rostral segments: 347 1002. =: 0.006, FD OG r= O:OF EEE (O:08: 2 0.006; pronotum: length 0.25 + 0.02, width across humeral angles 0.32 + 0.05, width across anterior margin 0.26 + 0.04; fore leg: femur length 0.31 + 0.01, tibia length O:27 = 0005, tarsi length 0.15 == .0:005. Second instar (Fig. 1C): Similar to first instar. Head and pronotum slightly darker and macula on abdomen not always appar- ent. Antennal segments turn grayish yellow and first antennal segment with a reddish band on external margin; first rostral seg- ment reddish brown. Spine of fore and mid- dle femora more apparent and two or three more spines present near longer one. Tri- chobotria evident on dorsum of segments II to VIII. Scent gland opening on segments V—VI not always visible. Measurements (n = 4). Body length 1.8 + 0.11; head length 0.25 + 0.04; width across eyes 0.37 + 0.01; interocular distance 0.18 + 0.01; postocular distance 0.02 + 0.02; antennal segments: I OS = 0,03,51-0:02522)0 7 O2572.0.013 IV 0.26 + 0.02, V 0.51 + 0.01; rostral seg- ments: 0312) =WOOR 02 fy 70:02 iit O:22 0.005; pronotumz leneth) O33" 0.02, width across humeral angles 0.46 + 0.02, width across anterior margin 0.31 + 0.03; fore leg: femur length 0.46 + 0.01, tibia length 0.42 + 0.005, tarsi length 0.19 eeCHO: Third instar (Fig. 1D): Similar to second instar. Head and thorax with irregular red markings, region anterior to eyes brown; eyes also darker, almost black. Spines of fore and middle femora longer and more numerous. Meso- and metathoracic wing pads start developing. Otherwise as in sec- ond instar. Measurements (n = 10). Body lensth 2.28 = ©.13:; head length) 026: 0.03; width across eyes 0.45 + 0.02; inte- rocular distance 0.24 + 0.02; postocular distance 0.03 + 0.03; antennal segments: I O:23t= 0103 .TRO:02"2 0, TO S52 O02; IV 0.38 + 0.02, V 0.6 + 0.04; rostral seg- ments:; 1.0.13 2.0.02. 1h. 0.34) 0.01, 8 0.14 + 0.008; pronotum: length 0.41 0.02, width across humeral angles 0.58 0.03, width across anterior margin 0.37 he Whe War 348 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Alloeorhynchus trimacula. A, Egg. B, First instar. C, Second instar. D, Third instar. E, Fourth instar. E Fifth instar. G, Adult male. Scale lines = 1 mm. VOLUME 106, NUMBER 2 0.02; fore leg: femur length 0.55 + 0.05, tibia length 0.46 + 0.01, tarsi length 0.24 a O01: Fourth instar (Fig. 1E): Pyriforme. Head and pronotum brown, posterior areas red- dish. Antennal segments gray, although joints still white. Abdomen yellowish, es- pecially near lateral margins; two sclero- tized rectangular areas appear on segments I and II; heart-shaped macula on last ab- dominal segments turn darker and divisions between segments become clearer; scent gland opening on segments V—VI obscure. Spines of front and middle femora double- ranked; internal margin of front and middle tibiae with a longitudinal line of flat black teeth extending half way onto abdomen. Meso- and metathoracic wing pads cover- ing half of following segment. Measure- ments (n = 10). Body length 3.2 + 0.26; head length 0.32 + 0.04; width across eyes 0.54 + 0.03; interocular distance 0.25 + 0.006; postocular distance 0.02 + 0.02; an- fennalascements:.1-0:3. 2: 0:02, 0:03. MOOS) th O49. 0:06, TV 0.532: 10:02.V O71. 25. 0:06; rostral: segments: 10:16 = OOP O27, 2220:02,, TIL.0.22, = 0.025 TV 0.16 + 0.01; pronotum: length 0.58 + 0.06, width across humeral angles 0.76 + 0.06, width across anterior margin 0.44 + 0.05; scutellum: length 0.32 + 0.04, width 0.53 + 0.04; fore leg: femur length 0.77 + 0.06, tibia length 0.7 + 0.07, tarsi length 0.31 + O:02: Fifth instar (Fig. 1F): Similar to fourth instar. Head dark brown; eyes black, slight- ly reddish near their base. Pronotum, lateral areas of meso- and metanotum and thoracic pleurae brown. Pinkish coloration of abdo- men completely disappeared, changed to creamy yellow; heart-shaped macula visi- ble, four small black maculae appear on connexivum of segments II to V. Ventral abdominal segments creamy yellow. Me- sothoracic wing pads covering metanotum and reaching middle of abdominal segment Il. Measurements (n = 10). Body length 4.32 + 0.29; head length 0.45 + 0.09; width across eyes 0.69 + 0.04; interocular 349 distance 0.29 + 0.03; interocellar distance 0.06 + 0.06; postocular distance 0.04 + 0.04; antennal segments: I 0.37 + 0.02, II 01055 0:004;51He-O:69. 20:05. TV e057" 0.02, V 0.86 + 0.04; rostral segments: I 0.2 0:02) Th033y= 0:04. Ti O26 45) 01022 TV 0.19 + 0.02; pronotum: length 0.85 + 0.06, width across humeral angles 1.08 + 0.05, width across anterior margin 0.54 + 0.03; scutellum: length 0.52 + 0.02, width 0.81 + 0.04; fore leg: femur length 0.97 + 0.08, tibia length 0.85 + 0.06, tarsi length 0.39 #0102: Adult (Fig. 1G): Elongate, body densely covered with semi-erect black and pale yel- low hairs. Head brownish black and shiny; antennal segment I grayish-yellow, antennal segments II to IV dark brown to gray, slightly paler distally; rostrum grayish brown. Pronotum reddish with one black spot on each humeral angle, another black, triangular spot on midline of posterior mar- gin; scutellum and hemelytron dull brown, costal margin varying from pale yellow to reddish. Most individuals with tarsi, tibiae, and apical one third to one half of all fem- ora blackish brown, basal half of femora pale yellow; a few individuals had lighter tibiae and tarsi. Propleura reddish, meso- pleura dark shiny brown, and metapleura dark dull brown. Abdominal venter creamy yellow with broad submarginal dark brown stripe on each side. Head broader than long, distinctly broader than collar; eyes promi- nent; antennal segment IV sometimes twist- ed; rostrum reaching mesosternum. Prono- tum clearly divided into anterior and pos- terior lobe; scutellum with three small de- pressions, two on disc and one near apex, apex distinctly bifid. Fore and mid femora with double row of spines, fore femora ro- bust. Female measurements (n = 10). Body length 5.69 + 0.61; head length 0.64 + 0.1; width across eyes 0.8 + 0.05; interocular distance 0.29 + 0.02; interocellar distance 0.09 + 0.09; postocular distance 0.09 + 0.04; antennal segments: I 0.46 + 0.03, II 0:05. +,0:005; TH: 0:96 0.09; TV 0:88 0.06, V 1.03 + 0.12; rostral segments: I 350 0322 es 0102, 21 0:55 2810105; OSirs 0.03, IV 0.21 + 0.02; pronotum: length 1.22 + 0.13, width across humeral angles 1.7 + 0.12, width across anterior margin 0.6 = 0.03; scutellum: length 0.83 = O.TL, width 0.98 + 0.08; fore leg: femur length 1.32 + 0.13, tibia length 1.13 + 0.08, tarsi length 0.51 + 0.04. Male measurements (n = 10): Body length 4.93" = 0:35; shead length 0.56 + 0.08; width across eyes 0.73 + 0.04; interocular distance 0.27 + 0.02; interocellar distance 0.09 + 0.09; postocu- lar distance 0.11 + 0.03; antennal seg- ments: I 0.41 + 0.04, II 0.05 + 0.006, III 0:83 500103, EV (O:7602270:04,'V 0:88. 0.04; rostral segments: I 0.22 + 0.02, II 0.5 ENOLO2S THO:32 122 10.03), PV Ost: 10:02; pronotum: length 1.06 + 0.05, width across humeral angles 1.54 + 0.06, width across anterior margin 0.51 + 0.02; scutellum: length 0.75 + 0.06, width 0.83 + 0.03; fore leg: femur length 1.13 + 0.06, tibia length 0.98 + 0.06, tarsi length 0.44 + 0.03. Biology.—Alloeorhynchus trimacula preyed on adults and nymphs of several species of Rhyparochromidae in the tribes Antillocorini, Myodochini, and Ozophorini. Species of lygaeids that A. trimacula cap- tured varied according to fig species and lo- cality. In Antillocorini it preyed on several unidentified species of Botocudo Kirkaldy, Cligenes distinctus Distant, and on a new genus; in Myodochini, it preyed on Myo- docha unispinosa Stal and Neopamera bi- lobata (Say); and in Ozophorini, on Ozoph- ora atropictoides Slater & Baranowski, O. baranowskii Slater & O’Donnell, O. con- cava Distant, O. consanguinea Distant, and O. maculata Slater & O’ Donnell. Adults of Alloeorhynchus trimacula ar- rived soon after the fig tree started to drop its fruits, also corresponding to the time when the first adult lygaeoids were first found around the trees. Counts of rhypa- rochromid and nabids two weeks after the first fruit had dropped reached 106.94 + 18.64 and 10.28 + 2.03 individuals, re- spectively per square meter (Cervantes, un- published data). Adults and nymphs of the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON two groups of bugs were abundant. Alloeorhynchus trimacula was observed preying on adults and nymphs of all the species mentioned above. They appeared to catch more nymphs of the species of lygaeoids that are good flyers, but they also preyed on the less mobile adults of Antillo- corini. Nabids run rather quickly among the litter looking for lygaeoids. Under labora- tory conditions, nymphs and adults of A. trimacula captured the rhyparochromids of- fered as soon as they were put in the con- tainers, catching at least one rhyparochro- mid every two days. Cannibalistic behavior was not observed, although it is possible that it could occur if not enough rhyparoch- romids were available. Female A. trimacula laid isolated eggs on leaf litter at the same time when several species of rhyparochro- mids were doing the same. The life cycle for both groups was completed between 50 to 60 days. It is possible that nabids and rhyparochromids move to different fruiting trees as the food resources for the bugs de- crease. Due to the asynchronous fruiting of the fig species, nymphs and adults of rhypa- rochromids and A. trimacula were present all year around. Alloeorhynchus trimacula adults sometimes were attracted to light. From the 30 native species of figs sam- pled, A. trimacula was recorded on Ficus calyculata Miller, F. cotinifolia (Kunth), F. insipida (Willd), F. lapathifolia (Liebm.) Miq., F. obtusifolia (Kunth), F. pertusa L.f., F. tecolutensis (Liebm.) Miq., and F. trigonata L. A few species are widely dis- tributed within Mexico, such as F. cotini- folia and F. insipida; although F. calyculata and F. lapathifolia have more restricted dis- tributions. Ficus cotinifolia can be found at sea level, while F. calyculata can reach 1,500 m. This wide range of distribution will show that A. trimacula will have a broader host list if a higher number of sam- ples for each individual fig species are tak- en. Distribution.—In the literature, A. ¢ri- macula has been reported from México, Guatemala, Panama, and Brazil (Champion VOLUME 106, NUMBER 2 1900, Harris 1928). Henry and Brambila (2003) reported A. trimacula from Florida in the United States of America. New records: MEXICO: Campeche: Ca- lakmul, Balamku; Calakmul, El Hormigue- ro; Calakmul, Km 25 to Calakmul Ruins; Calakmul, Calakmul Ruins, 172 m. Nayarit: Los Sabinos. Oaxaca: Km 55 Tuxpec-Oa- xaca; Km 11 Teotitlan-Huautla, 1,850 m; Km 14 Mitla-Albarradas. Puebla: Km 4 San Jose Acateno-Poza Rica, 288 m. Queretaro: Puente Escandilla. Tamaulipas: Gomez Fa- rias, Ejido el Azteca, 350 m; Gomez Farias, 276 m; Gomez Farias, Km 7 to Ciudad Mante, 320 m; Gomez Farias, Km 5 to Al- tacima, 600 m. Veracruz: Ozuluama, Km 40 Matatampico-Ozuluama, 3 m; Naranjos, Km 10 Naranjos-Chontla, 118 m; Naranjos, Km 30 San Sebastian-Naranjos, 262 m; Mi- santla, Km 14 Misantla-Martinez de la Tor- re, 80 m; Huatusco, Km 21 Jalcomulco- Huatusco, 546 m; Actopan, La Mancha, sea level; Actopan, Quiahuiztlan; Cuitlahuac, Km 44 Cuitlahuac-Tierra Blanca, 204 m; San Andres Tuxtla, Los Tuxtlas, 150 m; Ca- temaco, Km 10 to Tebanca; Isla, Km 13 Benito Juarez-Villa Juanita, 137 m; Isla, Km 45 Isla-Playa Vicente, 49 m; Jesus Car- ranza, Km 54 Boca del Monte-Poblado Doce, 102 m; Orizaba, Buena Vista, 1,200 m; Atoyac; Montepio, Quetzalan, 1,160 m. (IEXA) (CNIN). 351 ACKNOWLEDGMENTS I thank Harry Brailovsky (Instituto de Biologia, UNAM), Thomas Henry (Sys- tematic Entomology Laboratory, USDA, National Museum of Natural History), and John Lattin (Oregon State University) for their comments on the manuscript. Finan- cial support was provided by a CONACYT grant (34238-V), and is part of the project “Lygaeidae (Hemiptera-Heteroptera) asso- ciated with Ficus spp. (Moraceae) and their effect on seed predation, in the region of the Gulf of Mexico.” LITERATURE CITED Carayon, J. 1970. Etude des Alloeorhynchus D’ Afrique Centrale avec quelques remarques sur la classifi- cation des Nabidae (Hemiptera). Annales de la So- ciété Entomologique de France 6: 899—931. Champion, G. C. 1900. Biologia Centrali Americana. Heteroptera II p. 300, Plate XVIII, fig. 20. Harris, H. M. 1928. A monographic study of the he- mipterous family Nabidae as it occurs in North America. Entomologica Americana 9(1&2): 1—90. Henry, T. J. and J. Brambila. 2003. First report of the Neotropical damsel bug Alloeorhynchus trimacula (Stein) in the United States, with new records for two other nabid species in Florida (Heteroptera: Nabidae: Prostemmatinae). Proceedings Entomo- logical Society of Washington 105: 801—808. Lattin, J. D. 1989. Bionomics of the Nabidae. Annual Review of Entomology 34: 383—400. Pericart, J. 1987. Faune de France. Hemipteres Nabi- dae d’ Europe Occidentale et du Maghreb. Vol. 71. 185 pp. PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 352-360 A REVIEW OF THE GENUS DOIRANIA WATERSTON (HYMENOPTERA: TRICHOGRAMMATIDAE), WITH A DESCRIPTION OF A NEW SPECIES FROM NORTH AMERICA JOHN D. PINTO Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e- mail: john.pinto @ ucr.edu) Abstract.—The trichogrammatid genus Doirania is reviewed. The genus, previously known only from species in Japan and New Guinea, has a considerably more widespread distribution which includes North America and other areas of the Palaearctic. Three spe- cies are assigned. The North American species (D. elegans, n. sp.) is described and compared to congeners. The limits of Doirania and its relationship to other members of the tribe Oligositini are discussed. Key Words: As is the case with most trichogrammatid genera, Doirania is uncommonly collected and poorly known. It was described by Wa- terston (1928) for a single species, D. leef- mansi, a parasite in eggs of Tettigoniidae (Orthoptera), from Ambon, Indonesia. A second species, D. longiclavata Yashiro, from Japan, was added in 1980. Recent col- lections show the genus to be considerably more widespread. This paper presents a brief review of the genus and the descrip- tion of an additional species, D. elegans, which is widespread in eastern North America. Doirania is assigned to the tribe Oligo- sitini (Viggiani 1971). It has been consid- ered close to Oligosita (Doutt and Viggiani 1968, Yashiro 1980), and possibly deserv- ing only subgeneric status (Viggiani 1971). The primary feature separating the two gen- era historically is antennal club segmenta- tion (one in Doirania, two or three in Oli- gosita). Although additional traits separat- ing these taxa are proposed here, generic limits remain questionable, and certain spe- cies currently assignable to Oligosita ap- Hymenoptera, Trichogrammatidae, Doirania, taxonomy pear to be closer to Doirania. The current definition of Doirania is retained pending a complete evaluation of oligositine relation- ships. Terminology employed for most morpho- logical traits follows Doutt and Viggiani (1968) and Gibson (1989, 1997). Terms for antennal sensilla follow Olson and Andow (1993), and Pinto (1999). Doirania Waterston Doirania Waterston 1928: 386. Type spe- cies: Doirania leefmansi Waterston 1928, original designation. Doutt and Viggiani 1968: 499; Viggiani 1971: 208; Hayat and Subba Rao 1985: 241, 304; Pinto 1997773. Diagnosis.—Ranging from 0.4—0.8 mm in length; color light to dark brown. Head: Foramen magnum near top of head, at or above level of dorsal margin of eyes (Fig. 1). Antennal formula (Fig. 4): 2 anelli (2nd anellus discoid, inconspicuous), funicle 1- segmented, club 1-segmented; funicle well separated from club, transverse; club with 6 placoid sensilla (2 at middle and 4 api- VOLUME 106, NUMBER 2 cally), with at most | or 2 unsocketed setae, these at extreme base when present; length of scape and pedicle combined greater than that of flagellum. Mesosoma: Finely etched longitudinal line present medially on scu- tum and scutellum (correlated with a white line visible in dried specimens). Mesopleu- ron without a pleural suture. Venter of me- sothorax with distinct transepisternal sulci (see Gibson 1989). Forewing (Fig. 15) 2.5— 3X as long as wide; longest fringe setae length ca. %—% greatest wing width. Meta- soma: At least the anterior 3 terga longitu- dinally striate posteriorly (Figs. 13, 14). Two or more metasomal sterna completely divided longitudinally and ovipositor short, not extending beyond apex of metasoma or only slightly so (Figs. 8, 9). Male with a relatively broad, apically truncate ventro- medial projection on antepentultimate ster- num (Fig. 10). Male genitalia simple, as in several other oligositine genera, reduced to a single tube with two short apodemes at base (Figs. 10, 16, 17). Remarks.—The Oligositini are recog- nized by features of the male genitalia (Vig- giani 1971) as well as by the black rather than red compound eyes and the presence of one instead of two pair of setae each on the scutum and scutellum (Fig. 7). Genera assigned to the tribe in addition to Doirania include Oligosita, Megaphragma, Prestwi- chia, Epoligosita, Prosoligosita, Chaetos- trichella, Hayatia, Eteroligosita, and Pro- brachista. De Santis (1997) also assigned his new genus Brachistagrapha to this tribe but, known only from females, its place- ment requires confirmation. Doirania is separated from all other oli- gositine genera by the following combina- tion of characters: Foramen magnum placed near top of head (Fig. 1); antenna (Fig. 4) with a single club segment and a transverse funicle; three or more metasomal terga with a longitudinally striate posterior section (Figs. 13, 14); ovipositor relatively short, not extending beyond apex of metasoma; male genitalia simple (Figs. 16, 17), tubular, without complex apodemes basally. Certain ie) Nn ee) species of Epoligosita also have a one-seg- mented club (Doutt and Viggiani 1968) but in that genus, as in other oligositines, the foramen is placed near the middle of the head (Fig. 2), much closer to the mouth- parts, and the metasomal terga are uniform, lacking a striate posterior section. Chaetos- trichella, also with a one-segmented club (Doutt and Viggiani 1968, as Brachista), has a more ventrally placed foramen mag- num, an elongate funicle, more complex genitalia, and an ovipositor which extends considerably beyond the apex of the meta- soma. Doirania can be separated unambiguous- ly from other described oligositine genera. Yet it is probable that certain undescribed species, currently assignable to Oligosita based on antennal segmentation, are actu- ally closer to Doirania. Unfortunately, these taxa are known only from females which makes their placement difficult at present. Oligosita is the largest genus of Trichogrammatidae and it is likely that it is composed of two or more unrelated line- ages which simply lack certain derived fea- tures of other oligositine genera. A detailed analysis of the entire tribe is required before generic limits are satisfactorily clarified. KEY TO SPECIES OF DOIRANIA (Traits pertain to females unless indicated) 1. Ovipositor elongate, 4—% longer than hind tib- ia, extending slightly beyond cerci (Fig. 9). Male genitalia elongate, ca. Y; longer than hind tibia, with apodemes curving laterally at base (Fig. 17). Forewings with a small but distinct fumate cloud near apex of stigma. Known from Papua New Guinea and Ambon, Indonesia . . D. leefmansi — Ovipositor shorter, its length ranging from dis- tinctly less than to subequal to hind tibial length, not quite attaining level of cerci (Fig. 8). Male genitalia shorter, length only 0.5—0.6 that of hind tibia, with apodemes straight, not curving laterally at base (Fig. 16). Forewing (Fig. 15) at most slightly darkened at apex of stigma, without a distinct fumate cloud ..... 2 . Metasomal terga with longitudinally striate posterior section distinct, at least second and third visible terga with striate section ca. twice the length of uniformly sclerotized anterior i) 354 Figs. 1—6. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1, Doirania elegans, posterior view of head. 2, Epoligosita sp. (same). 3, Doirania elegans, right maxillary palp (ventral). 4, D. elegans, antenna (lateral). 5, D. elegans, sensilla on dorsum of forewing disk anterior to retinaculum (see arrow in Fig. 15 for location). 6, D. leefmansi (same). section (Fig. 13). Male club distinctly longer than scape. New World — Metasomal terga with longitudinally striate D. elegans, new species posterior section relatively indistinct, sclero- tized striae poorly distinguished from adjacent membrane, terga with striate section, at most, subequal in length to uniformly sclerotized an- terior section (Fig. 14). Male club shorter than scape. Palaearctic D. longiclavata Doirania elegans Pinto, new species (Figse1, 352728) P1135 FS; Lo) Two forms (A & B) of this species are recognized (see Variation). The description is based on North American specimens of Form A. Unless indicated, quantitative data are based on five specimens from different locales; data represent means unless report- ed as a range. Description.—Female: Body length 0.44—0.75 mm. Medium to dark brown in color except face, legs and antenna lighter brown; forewing only slightly fumate be- neath venation. Head: Antenna (Fig. 4) with toruli at level of ventral margin of eyes; scape and pedicel together 1.2 com- bined length of funicle and club; club 1I- segmented; mean length/width of antennal segments as follows: scape, 2.76; pedicel, 1.95; -funicle, 0.67; .club,,2:24;, relative length of antennal segments in same order: 47/37/14/56; club 1.2X as long as scape, with terminal sensillum (= UPP trichodea D of Olson and Andow 1993) setiform (see Variation), 0.45 segment length (Fig. 11), placed on a small apical truncate pedestal; longest flagelliform setae (= MPP trichodea VOLUME 106, NUMBER 2 A) 0.65 club length. Maxillary palp narrow- ing asymmetrically to apex, with apical sen- sillum longer than half the length of adja- cent seta (Fig. 3). Mesosoma: Propodeum transverse (Fig. 7), slightly arcuate at mid- dle and slightly longer (<2) than meta- notum. Hind femur ca. 0.3 as wide as long; tarsi relatively short, pro-, meso- and meta- tarsi ca. 0.80, 0.75 and 0.75 the length of their respective tibia; segment I slightly shorter than II and III on fore- and middle legs, all tarsomeres subequal on hind legs. Forewing (Fig. 15) 2.5—3.0X as wide as long, longest fringe setae 0.4—0.5 maximum wing width; sensilla anterior to retinaculum on dorsal surface of disk relatively large, digitiform, usually clavate (Fig. 5). Hind wing with 2 complete tracks of setae on disk, a partial third track present at apex in some specimens. Metasoma: Terga anterior to VII clearly differentiated into an anterior, uniformly sclerotized section and a poste- rior longitudinally striate section; striate section elongate, twice as long as anterior section on most terga (Fig. 13). Venter (Fig. 8) with ail visible sterna except hypogyn- ium narrowly divided longitudinally; hy- pogynium broadly emarginate; first visible sternum with a distinct posterior longitudi- nally striate section, others obsolescently striate posteriorly. Ovipositor 0.76 (0.67— 0.89, n = 10) as long as hind tibia (see Variation). Male: As in female except antennal club with only 3 placoid sensilla (1 basal, 2 near apex), and fewer flagelliform setae; club longer than scape (1.1—1.4 as long); only anterior 3 metasomal terga with an obvious posterior striate section. Venter with a me- dial, posteriorly projecting lobe on antepen- tultimate sternum (as in Fig. 10, see Re- marks); lobe only slightly longer than wide. Genitalia (Fig. 16) gradually but distinctly widened to apex, length 0.5—0.6 that of hind tibia, with basal apodemes straight, not curved laterally. Variation.—There appears to be two sympatric forms (A & B) of this species. The description and types refer to Form A. 355 In Form A the terminal club sensillum in females is setiform (Fig. 11) and is similar in dimensions to the numerous flagelliform setae on the same segment; also the ovi- positor is relatively short (see above). In Form B, the terminal sensillum is broader and, although tapering, it remains slightly truncate at the apex rather than acuminate (Fig. 12). Also the ovipositor is longer in Form B [0.98 (0.87—1.14, n = 12) as long as hind tibia], and the discal forewing setae are usually shorter. I am unable to distin- guish the males of these variants. Although the two forms are not geographically dis- junct there is a degree of character overlap. Additional material and study is required to determine if this variation is taxonomically significant. A small series from Ecuador (Limonco- cha, Napo Province) consisting only of fe- males is questionably conspecific to D. ele- gans. In these specimens the propodeum is more distinctly produced at the middle, and the pedicel is subequal in length to the scape. Ovipositor length and the form of the terminal club sensillum are as in Form A. Types.—Holotype 2. UNITED STATES. Oklahoma: Red Oak (Latimer Co.); ix- 1993; flight intercept trap (FIT); K. Ste- phen. Paratypes, 2 6, 1 2 with same data. Holotype and one male paratype deposited in Canadian National Collection, Ottawa (CNC); the two additional paratypes depos- ited in the Entomology Department, Uni- versity of California, Riverside. Types are mounted in Canadian balsam on glass slides. Seven carded females from the same series are not designated as types. The slide-mounted types belong to Form A. Diagnosis.—Doirania elegans is closest to the Palaearctic species, D. longiclavata. It is separated by the more distinct and lon- ger striate section of the metasomal terga (see key to spp.). Also, in the male of D. elegans the club is distinctly longer than the scape, not shorter as in D. longiclavata. Sensilla anterior to the retinaculum on the dorsal surface of the forewing apparently provide an additional difference. In the new 356 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-10. Doirania. 7, D. elegans, mesosoma (dorsal). 8, D. elegans, metasoma (ventral). 9, D. leefmansi, metasoma (ventral). 10, D. leefmansi, apex of metasoma (ventral). species they are rather elongate, digitiform and often clavate (Fig. 5); in D. longicla- vata, as in D. leefmansi as well, they are shorter and typically acuminate apically (Fig. 6). Ovipositor length and shape of the apical antennal sensilla in females of D. longiclavata resemble Form A of D. ele- gans. Although very similar phenetically, it is not yet clear if D. elegans and D. longicla- vata are sister species. The only similarity which may be derived is the asymmetrically VOLUME 106, NUMBER 2 narrowed maxillary palp (Fig. 3). It occurs in both species but not in D. leefmansi. Geographic distribution.—Eastern Unit- ed States, Arizona, southeastern Canada, with a single record of questionable con- specifics from Ecuador. Remarks.—Hosts are unknown. Several of the records (see below) suggest that D. elegans occurs in relatively mesic habitats. Males of Doirania and of related genera are characterized by at least one medial pro- longation on the sternal region of the me- tasoma (Pinto and Viggiani, in preparation). The few males available of D. elegans and D. longiclavata are mounted on slides and do not allow adequate description of this structure. However, it appears to be similar to that occurring in D. leefmansi (Fig. 10). Material examined.—136 2, 8 6. Iden- tification of ‘form’ (A or B) is possible for slide-mounted females only. Consequently an indication of ‘A’ or/and ‘B’ does not fol- low those records based only on males or carded females. In cases where both slide- mounted and carded females are available, the number of individuals that identification of ‘form’ is based on is indicated (e.g., A/ 2, signifies that two of the females in the series were slide-mounted and identifiable to form). CANADA. Ontario: Ottawa (Innes Point); viii-20/27-1985; 1 2; L. Dumochel/ J. Denis (B). Powell’s Lake (105 km NE Lake Superior); viii-16-1980; 1 9; M. Kaulbars. Shirley’s Bay; vii-27/ix- 10-1985; 1 2; M. Sanborne/H. Goulet. UNITED STATES. Arizona: Brawley Wash (Pima Co.); viii-3-1982; 1 2; G. Gibson (A). No- gales (N edge of town); ix-27-1985; sweep- ing; | @; J. Pinto (A). Florida: Archbold Biological Research Station; x-27/xi-30- 1988; Malaise trap; 2 2; D. Wahl (A/1). Bradenton; x-19/26-1985; sweep; 1 2; C. Yoshimoto. Everglades National Park (Long Pine Key); vi-6/vili-26, & vili/xi- 986; Malaise/flight intercept trap; 6 &; S. & J. Peck (A). Gainesville; iv-17/23-1988; 1 (sex undetermined); D. Wahl. Gainesville; iv-8/14-1987, xii-1/7-1986; 2 2; W. Mason. 357 Highlands Hammock (Highlands Co.); ix- I8-1987isweep? 2. 2s 01, Masner 1(A/1): Georgia: Athens (Whitehall Forest); 1x-14/ 28-1987; flight intercept trap, hardwood forest (beaver swamp); 1 2; CNC Hym. Team (A). Sapelo Island; vii-18/ix-11-1987, ix-9/21-1987, x-15/xil6-1987; Malaise trap; live oak forest; 14 2, 1 6; CNC Hym. Team (A/4). Tifton, 13 km NW; ix-25/x-17- 1985; pan trap; 2 2; M. Keller (A/1). I1li- nois: Centralia (along roadside: Hwy 51 & Bethel Rd.); yellow pan trap ‘“‘among weeds ne soybeans 1x-I2/L7-1995; 2 ia Triapitsyn (A). Centralia, 3 mi N; ix-12/17- 1995; yellow pan trap “‘in grass nr. pond”; 1 @; S. Triapitsyn. Centralia, 2 mi S (of downtown); 1x-13/17-1995; yellow pan trap “onsedge-of forest; 21-2: Sv Tuapitsyn. Centralia, 8 mi E; ix-7-1993; sweeping “open field’; 2 2; J. Pinto. Litchfield; x-3- (O83:>sweep, rl 72) 1 6-5),/D; Huberr( A): Marion Co. (Myers Rd. nr. hwys. 57 & 161); ix-12/17-1995; yellow pan trap “‘in grass in swampy area”; 1 2, 1 6; S. Triap- itsyn. Kansas: Manhattan, 2 mi S; ix-6- 1983; 1 (sex undetermined); J./D. Huber. Wauconda Lake (2 mi W Glen Elder, Mitchell Co.); viii-28-1985; sweep; 1 @; J. Pinto. Maryland: Port Republic; viii/ix- 1986; flight intercept trap; 9 2, 1 6; M. Sharkey/Munroe (B/1). Prince Frederick, 7 km S; v-7/vii-7-1987; 1 2; CNC Hym. Team. Missouri: Columbia (Hinkston Creek); ix-8-1987; sweep; 1 @; J. Pinto. Williamsville (Wayne Co.); vii-16/viii-8- 1988, viti-1987, viili-8/31-1988, ix-1/20- 1988, ix-10/26-1987, ix-20/x-20-1988, x- 21/xi-11-1987; Malaise trap; 38 2, 2 6; J. Becker (A/3, B/5). Nebraska: Odessa, 6.8 mi E; viti-29-1983; 2 2; J. Pinto (A). Oklahoma: Red Oak (Latimer Co.); 9 2, 2 3d; (see Types). North Carolina: Whiteside Mtn. (Jackson Co.); ix-13-1987; sweep; 3 2; L. Masner (B/1). South Carolina: Fran- cis Beidler Forest (nr. Harleyville); ix-22- 1987; sweep; 1 2; L. Masner (A). Francis Beidler Forest (10 km NE Harleyville); v- 26/vi-11-1987; flight intercept trap; “bald cypress swamp”; | @. Pendleton (Tangle- 358 wood Springs; 34°38.7'S, 82°47.1'W; 225 m); vii-30/viii-20-1987, ix-1/9-1987, ix-15/ 30-1987, x-16/xi-3-1987; Malaise trap; 6 2: J. Morse (A/1). Pendleton (225 m); vii- 15/22-1987, vii-29/viii-5-1987; Malaise trap; 2 2; CNC Hym. Team (A/1). South Dakota: Pickstown (Charles Mix Co.); viii- 26-1985; “‘sweeping riparian’’; 4 2; J. Pin- to (A/1, B/1). Texas: Ben Bolt, 8 mi NW (La Copita Res. Stn.); ix-28/30-1990; Mal- aise trap; | 2; R. Wharton/J. Woolley. Cly- mer Meadow (Hunt Co.); vii-9-1991; sweep; 1 @; J. Woolley (A). College Sta- tion; viii-26-1987; 1 2; J. Woolley/G. Zol- nerowich (B). College Station (Lick Creek Park); vii-30-1987; sweep; 1 2; J. Woolley. College Station (Lick Creek Park); x-16/x1- 17-1987; sweep; 1 2; J. Woolley/J. Heraty. Hallsville, 2 mi W; iv-27-1984; sweep ‘roadside forbs’”’; | 2; W. Ewart (B). Hear- ne, 8 mi. E.; x-22/27/1990; Malaise trap; 1 2, 1 3; J. Woolley, et al. Park Hill Prairie (Collin Co.); vii-9-1991; 2 2; J. Woolley (B/1). Virginia: Blacksburg, 8 km NW (1000 m); vi-9/19-1987, vi-19/30-1987, vii- 13/19-1987; Malaise trap; 6 2; CNC Hym. Team (A/2). Doirania longiclavata Yashiro (Fig. 14) Doirania longiclavata Yashiro 1980: 131. Diagnosis.—Similar to D. elegans except for the more poorly defined striate section of the metasomal terga (cf. Figs. 13, 14), the smaller club/scape ratio in males, and the differently shaped sensilla at the base of the forewing (cf. Figs. 5, 6). Types.—Holotype @, from JAPAN, **Hatadera, Matsuyama City, Ehime Pref., Shikoku’”’; x-19-1977; N. Yashiro; “‘on turfs’; presumably in the Entomological Laboratory, University of Osaka Prefecture (inquiries regarding the holotype were un- answered). Geographic distribution.—Previously re- corded only from Japan, but probably wide- spread in Palaearctic. Currently known from Japan, eastern Russia, and France. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Material examined.—FRANCE. Dept. Gironde: St. Colombe (nr. Castillon-la-Ba- taille), 44°54’N, 00°02'W; viii-17-2000; suction trap; 1 2; M. van Helden. JAPAN. Kyushu: Fukuoka (Mt. Tachibana); vili-26/ 31-1979; yellow pan traps, “primarily ev- ergreen forest’; 4 2; K. Yamagishi. Hon- shu: Iwate (Mt. Hayachine); 400 m; viii-2/ 8-1989; Malaise trap; M. Sharkey. [Several additional Japanese records cited by Yosh- iro 1980]. RUSSIA. Primorskiy Krai: Ta- jvaza (30 mi NE Vladivostok); flight inter- cept trap “coastal forest”; viii-5-1992; 5 @, 1 3; B. Gill. Krasnodarskiy Krai: Krasno- dar, nr. (All Russian Research Institute of Biological Plant Protection); viii-30/31- 2001; yellow pan trap; 1 2; V. Kustjukov. Remarks.—Hosts are unknown. As with D. elegans, D. longiclavata apparently fre- quents mesic habitats. Yashiro (1980) states that the species has been collected in “‘pad- dy fields and grassplots.”’ Doirania leefmansi Waterston (Piss: 6, 90a) Doirania leefmansi Waterston 1928: 286; Yashiro 1980: 133; Doutt and Viggiani 1968: 499; Viggiani 1971: 209; Caudwell 2000: 218. Diagnosis.—Female. Body light brown in color. Antenna with scape subequal in length to club. Maxillary palp regular at apex, not narrowing asymmetrically. Fore- wing with a small fumate cloud directly be- hind stigma; sensilla anterior to retinaculum on dorsal surface of disk small, acuminate (Fig. 6). Ovipositor elongate, distinctly lon- ger than hind tibia, extending beyond cerci (Fig. 9). Male. Antenna with scape also subequal to club. Genitalia elongate, ca. ¥, longer than hind tibia, with basal apodemes directly laterally (Fig. 17). Types.—Holotype 2, ““Ambon (D.E.I.)” (currently Indonesia) from “eggs of Sexava coriacea,” iv-30-1925, S. Leefmans coll.; stated to be in The Natural History Muse- um, London (Waterston 1928) but not lo- VOLUME 106, NUMBER 2 359 Figs. 11-17. Doirania. 11, D. elegans, antenna (Form A) (arrow at apical sensillum). 12, D. elegans, antenna (same, Form B). 13, D. elegans, metasoma (dorsal). 14, D. longiclavata, (same). 15, D. elegans, forewing (arrow at location of disk sensillae; see Figs. 5, 6). 16, D. elegans, male genitalia (venter, arrow at basal apodemes). 17, D. leefmansi (same). 360 cated (4 2 paratypes with same data as ho- lotype examined). Geographic distribution —Known from Indonesia (Ambon) and Papua New Guin- ea. Material examined.—INDONESIA: Am- bon (see Types). PAPUA NEW GUINEA: Dami, West New Britain; ix-29-1991; T. M. Soluloz 11 27-156. New Hanover pst]: 1932; ‘ex. eggs of locustid’; J. L. Froggatt; 1 2. Culture material reared in Papua New Guinea, of unknown origin, 10 2, 1 d. Remarks.—Doirania leefmansi is known to attack eggs of species of Tettigoniidae (Segestes spp.), pests of oil palm, in Papua New Guinea. Although their role in affect- ing pest populations is questionable, these wasps continue to be cultured and released into oil palm growing areas (Caudwell 2000). ACKNOWLEDGMENTS Gary Platner was responsible for speci- men and plate preparation, and assisted with photographs. This study was supported by grants from the USDA (NRI) and NSF (PEET). Material studied came from nu- merous sources, the greatest number from J. Huber of the Canadian National Collec- tion and J. Woolley, of Texas A&M Uni- versity. LITERATURE CITED Caudwell, R. W. 2000. A sustainable IPM system for oil palm in Papua New Guinea. The British Crop Protection Council Conference, Pests & Diseases 1; 215-220. De Santis, L. 1997. Afelinidos y Tricogramatidos de la colecci6n del Dr. Alejandro A. Ogloblin (Insec- ta—Hymenoptera). I] Segunda Comunicacion. Academia Nacional de Agronomia y Veterinaria 51: 8-17. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Doutt, R. L. and G. Viggiani. 1968. The classification of the Trichogrammatidae (Hymenoptera: Chalci- doidea). Proceedings of the California Academy of Sciences (4th series) 35: 477—586. Gibson, G. A. P. 1989. Phylogeny and classification of Eupelmidae, with a revision of the world genera of Calosotinae and Metapelmatinae (Hymenop- tera: Chalcidoidea). Memoirs of the Entomologi- cal Society of Canada, No. 149, 121 pp. . 1997. Morphology and terminology. Chap. 2, pp. 16-44. In Gibson, G. A. P., J. T. Huber, and J. B. Woolley, eds. Annotated Keys to the Genera of Nearctic Chalcidoidea (Hymenoptera). NRC Research Press, Ottawa, Canada, 794 pp. Hayat, M. and B. R. Subba Rao. 1985. Family Tricho- grammatidae, pp. 239-245, 304-308. Jn Subba Rao et al., eds. The Chalcidoidea (Insecta: Hy- menoptera) of India and the adjacent countries, Part I. Reviews of families and keys to families and genera. Oriental Insects 19: 163-310. Olson, D. M. and D. A. Andow. 1993. Antennal sen- silla of female Trichogramma nubilale (Ertle and Davis) (Hymenoptera: Trichogrammatidae) and comparisons with other parasitic Hymenoptera. International Journal of Insect Morphology and Embryology 22: 507-520. Pinto, J. D. 1997. Trichogrammatidae. Chap. 22, pp. 726-752. In Gibson, G. A. P., J. T. Huber, and J. B. Woolley, eds. Annotated Keys to the Genera of Nearctic Chalcidoidea (Hymenoptera). NRC Re- search Press, Ottawa, Canada, 794 pp. . 1999 (1998). The systematics of the North American species of Trichogramma (Hymenop- tera: Trichogrammatidae). Memoirs of the Ento- mological Society of Washington, No. 22, 287 pp. Viggiani, G. 1971. Ricerche sugli Hymenoptera Chal- cidoidea XXVIII. Studio morfologico comparati- vo dell’armatura genitale esterna maschile dei Tri- chogrammatidae. Bollettino del Laboratorio di Entomologia Agraria ‘Filippo Silvestri’ di Portici 29: 181-222. Waterston, J. 1928. On a trichogrammid (Doirania leefmansi, gen. et sp. n.) reared from eggs of Sex- ava (Orth.) in the Dutch East Indies. The Annals and Magazine of Natural History (10)2: 386-388. Yashiro, N. 1980. A new species of the genus Doirania from Japan (Hymenoptera: Trichogrammatidae). Transactions of the Shikoku Entomological Soci- ety 15: 131-134. PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 361-395 NEW SPECIES OF CENTRAL AMERICAN CULICOIDES LATREILLE (DIPTERA: CERATOPOGONIDAE) WITH A SYNOPSIS OF SPECIES FROM COSTA RICA GUSTAVO SPINELLI AND ART BORKENT (GS) Departamento Cientifico de Entomologia, Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata, Argentina (e-mail: spinelli@museo.fonym.unip.edu.ar); (AB) Research Associate, Royal British Columbia Museum, American Museum of Natural History, and Instituto Nacional de Biodiversidad, 1171 Mallory Road, Enderby, British Columbia, VOE 1V3, Canada (e-mail: aborkent @jetstream.net) Abstract.—Ten new species of Central American Culicoides are described, illustrated and placed to subgenus or species group. Their position in previously published keys is indicated and their features discussed in light of the most recent revisions. Eight of the new species are known only from Costa Rica, one is known from Costa Rica and Panama and one is recorded from Honduras and El Salvador. The new species are named Culi- coides annettae, C. chaverrii, C. cummingi, C. hermani, C. hondurensis, C. monicae, C. picadoae, C. ronderosae, C. trifidus, and C. zumbadoi. A list of 148 Culicoides species known or suspected of being in Costa Rica is given in a table. Of these, 42, including the new species, are recorded from Costa Rica for the first time. Most of the new species are recorded from mid to high elevations. Resumen.—Se describen, ilustran y ubican en su respectivo subgénero o grupo de es- pecies, a diez especies nuevas de Culicoides de América Central. Se indica su posicion en las claves publicadas previamente, y sus caracteres distintivos se discuten a la luz de las revisones mas recientes. Ocho de las especies nuevas se conocen solo para Costa Rica, una para Costa Rica y Panama y la restante para Honduras y El Salvador. Las especies nuevas son nominadas Culicoides annettae, C. chaverrii, C. cummingi, C. hermani, C. hondurensis, C. monicae, C. picadoae, C. ronderosae, C. trifidus y C. zumbadoi. Se ofrece en una tabla, una lista de 148 especies de Culicoides conocidos 0 sospechados de ser hallados en Costa Rica, 42 de los cuales, incluyendo a las especies nuevas, son re- gistrados por primera vez para Costa Rica. Muchas de las especies nuevas se registran para altitudes medias a elevadas. Key Words: Ceratopogonidae, Culicoides, Costa Rica, Honduras, El Salvador, Central America, South America Species in the genus Culicoides Latreille world, from the tropics to far southern and are by far the most notorious members of northern areas, from coastal areas to very the 103 currently recognized genera of Ce- high altitudes (up to 4,200 m). Many of ratopogonidae. Not only is Culicoides the these are miserable pests of humans and do- most diverse genus in the family, with mestic animals and serve as vectors of a 1,255 named extant species, but members variety of diseases (Borkent, in press). occur virtually throughout the terrestrial In spite of their economic, medical and 362 veterinary importance, it is surprising that the species of Culicoides of many areas re- main poorly understood. The genus is so diverse that even in regions which have been studied for many years (e.g., United States), there are still quite a number of spe- cies represented in museums which remain to be described (and undoubtedly many more yet to be discovered!). Furthermore, there remains a great need to provide keys to species in some broader areas; there is still no key available to the species of either the United States or Canada! In this paper we describe 10 additional species of Culicoides from Central Ameri- ca. We have been fortunate to have at hand the excellent treatment of the Culicoides of Panama by Wirth and Blanton (1959) and the wing atlas of Neotropical species by Wirth et al. (1988). We have indicated where each new species terminates in their key, with additional comments when more recent revisions are available. Borkent and Spinelli (2000) recently catalogued all spe- cies of Ceratopogonidae south of the United States and provided references to the most recent revisions and keys. Although we have placed the new spe- cies in subgenera or species groups, readers should be aware that the current classifica- tion of Culicoides is in dire need of careful cladistic analysis. Some subgenera are “garbage can” groupings of species which cannot be placed elsewhere and every ma- jor region of the globe has species of Cu- licoides which cannot be placed even in those subgenera but are relegated to species groups or listed as “‘miscellaneous.”’ MATERIALS AND METHODS All specimens were mounted on micro- scope slides using the technique described by Borkent and Bissett (1990). Terms for structures follow those used in the Manual of Nearctic Diptera (McAlpine et al. 1981). Terms for wing veins follow the system of the Manual of Nearctic Diptera, with mod- ifications proposed by Szadziewski (1996) (Table 1). Names of veins are in upper case PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Terms for wing veins and cells as used here and by Wirth (and his coauthors) in his publica- tions. As Used in This Work Wirth and Coauthors Veins R, R, R, radial crossvein R, Rs or Ry; Rowe intercalary of Leptoconops M M,.> M, M, M, M, CuA, IME, CuA, Cu, CuP Cu, A A Cells 1“radial cell 1% radial cell (or I‘ anterior radial cell) 2™ radial cell 2° radial cell (or 2™ anterior radial cell) Tr; R; m, M, m, M, cua, M, anal and those of cells in lower case. Pale areas in cell r,; posterior to or immediately distal to the 2nd radial cell are called poststig- matic pale spots. More specific larval and pupal terms follow Lawson (1951). Ratios used are those explained in Spinelli et al. (P9993): Adults were collected by sweeping with an aerial net, with malaise traps or with light traps (a fluorescent bulb on a white sheet). Larvae and pupae were extracted through manual examination of substrates placed in shallow dishes or pans. Dates of collections of reared specimens refer to the date upon which the immatures were col- lected. Most of the specimens reported here have a “CD” number which refers to fur- ther collecting and habitat details in the ju- nior author’s personal collecting notes. Specimens are deposited in the following collections: VOLUME 106, NUMBER 2 CNCI—Canadian National Collection of Insects, Ottawa, Ontario, Canada. MLPA—Museo de la Plata, Facultad de Ciencias Naturales y Museo, Uni- versidad Nacional de La Plata, Argentina. INBC—Instituto Nacional de Biodiversi- dad, Santo Domingo de Heredia, Costa Rica. USNM—Smithsonian Institution, National Museum of Natural History, Washington, D.C., U.S.A. Wing photomicrographs were taken with a Canon PowerShot, S40, digital camera through a Leica, DMLB, compound re- search microscope. Wings were photo- graphed in either two or three sections at either 10X or 20 and the separate images assembled in Photoshop™, version 6. RESULTS Culicoides (Avaritia) hermani Spinelli and Borkent, new species (Figs. 1A, 2A—E, 10A) Diagnosis.—Small adults in the andicola species group, very similar to C. orjuelai Wirth and Lee, from which it may be dis- tinguished by the shorter distal flagello- meres, smaller number of mandibular teeth, tibiae with subbasal pale rings, the pale area centered on crossvein r-m broadly abutting costal wing margin, and the presence of a very faint, indistinct pale area in cell cua,. Description.—Male. Similar to female with usual sexual differences. Wing length 1.15 mm; width 0.52 mm; CR 0.61. Geni- talia (Fig. 2E): Tergite 9 subquadrangular, lacking apicolateral processes; sternite 9 short, with broad, deep posteromedial ex- cavation. Gonocoxite with slender, pointed dorsal, ventral roots, the latter contacting at midline; gonostylus shorter than gonocox- ite, moderately curved, tip pointed. Para- meres (Fig. 2D) separate, each with short, stout anterolateral process, midportion slen- der, nearly straight, tapering to slender, fil- amentous, recurved tip. Aedeagus triangu- lar; basal arch very low, extending 0.20 of 363 total length; lateral arms well sclerotized, tapering to short, posteromedial projection, tip blunt; with dorsal, subapical, sclerotized process present. Female. Head: Dark brown. Eyes with numerous interommatidial spicules, contig- uous by distance equal to diameter of 1.5 ommatidia. Flagellum (Fig. 2A) uniformly brown; flagellomeres 1—8 short, vasiform, 9-13 subcylindrical, stout to apices; AR 1.08 (1.00—1.16, n = 5); sensilla coeloco- nica on flagellomeres 1, 9—13. Palpus (Fig. 2B) brown; third segment with conspicuous irregular pit; PR 3.05 (2.90—3.40, n = 5); P/H ratio 1.03 (1.00—1.09, n = 5). Mandi- ble with 15 (n = 5) teeth. Thorax: Dark brown, scutum with sublateral, faint paler bands. Legs brown, fore and midfemora pale apically, tibiae with narrow, subbasal rings; hind tibial comb with five spines, one nearest spur longest. Wing (Fig. 1A), length 1.25 (1.16—-1.36, n = 5) mm; width 0.64 (0.60—0.68, n = 5) mm; CR 0.61 (0.59- 0.63, n = 5); with pale areas much reduced; faint pale area near basal arculus continuous with basal pale area in anal cell; conspicu- ous pale spot centered on crossvein r-m, broadly abutting costal wing margin; post- stigmatic pale spot in cell r, transverse, in- cluding distal half of second radial cell; very faint, indistinct pale area in cell cua,; anal cell with distal rounded pale area. Ma- crotrichia very sparse, scattered on distal half of wing. Halter pale. Abdomen: Dark brown. Two ovoid, unequal spermathecae, tapering to moderately long necks (Fig. 2C), measuring 47 (44-53, n = 5) by 39 (36—44, n = 5) um, and 38 (36—46, n = 5) by 34 (30-40, n = 5) wm: rudimentary third, ring present. Distribution.—Culicoides hermani is known only from high elevations in south- ern Costa Rica and northern Panama (Fig. 10A), from 1,600—2,750 meters. The adults from 15 km N San Isidro de el General were taken at Reserva Avalon, a few kilo- meters SW of Division. Bionomic information.—The holotype of C. hermani was swept from a flowering 364 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Wings of female Culicoides. (A) C. hermani; (B) C. annettae; (C) C. hondurensis; (D) C. monicae;: (E) C. chaverrii; (F) C. ronderosae; (G) C. cummingi; (H) C. picadoae; (1) C. zumbadoi; (J) C. trifidus. Scales = 0.2 mm. VOLUME 106, NUMBER 2 bush on the trail leading up to Volcan Bart in northern Panama. The allotype female and two of the paratypes were taken while biting humans and one paratype female was collected from a light. Adults were present in high elevation cloudforest habitat and have been collected in March, September and October. The female adult habit of biting humans and the presence of this species in higher altitudes is shared with the other Neotrop- ical species of the alticola group, all four of which are restricted to elevations above 3,000 meters in Colombia (Rodriguez and Wirth 1986). Taxonomic discussion.—Culicoides her- mani is a member of the andicola species group of the subgenus Avaritia. This new species keys out in Wirth and Blanton (1959) to couplet 13 where it may be dis- tinguished by the presence of a pale apical band on the fore and midfemora but with the hind femora entirely dark. As a member of the andicola species group, C. hermani will key in Rodriguez and Wirth (1986) to C. puracensis Wirth and Lee, a Colombian paramo species. Females of C. hermani can be distinguished from that species by the sensillar pattern (1, 11—13 in C. puracensis) and the flagellomere 9 without a distal con- striction. Furthermore, male C. hermani lack the apicolateral processes on tergite 9 and the stouter distal posteromedial projec- tion of aedeagus present in C. puracensis. Wirth and Lee (1967) mention an unde- scribed Costa Rican species that is almost certainly the same as that described here as C. hermani. The male and female were associated by the presence of shared wing pattern and body coloration. In addition, they represent the only species of the alticola species group in Costa Rica and Panama. Types.—Holotype ¢d, Panama, 8 km W Boquete, 21-IX-1993, 1,600 m, A. Borkent, CD1583 [CNCI]; allotype 2, Costa Rica, 4 km N Sacramento, 10-X-1993, 2,750 m, A. Borkent, biting human, CD1609 [CNCI]. Paratypes, 4 2, as follows: same data as 365 allotype, 1 2 [MLPA]; Costa Rica, 15 km N San Isidro de el General, 2,250 m, 12- IlI-1999, A. Borkent, biting human, CD5000, 2 2 [CNCI]; Costa Rica, 15 km N San Isidro de el General, 2250 m, 13/14- II-1999, A. Borkent, light trap, CD5002, 1 2 LINBC]. Derivation of specific epithet.—This spe- cies, the only one described herein which is known to bite humans, is named after the junior author’s brother and dear friend Her- man Borkent in recognition of his loving support through all the years. Culicoides (Hoffmania) annettae Spinelli and Borkent, new species (Figs. 1B, 2F—-J, 3A—F 10A) Diagnosis.—Medium sized adults in the guttatus species group very similar to C. ocumarensis Ortiz, from which it can be distinguished by the slender third palpal segment with capitate sensilla in a shallow pit as well as in 2—3 other shallow depres- sions, the shorter proboscis, the distal pale spot in cell r; barely abutting the wing mar- gin, the parameres broadly fused at their ba- ses, and the aedeagus with a truncated apex. Description.—Male. Similar to female with usual sexual differences. Wing length 0.94 mm; width 0.36 mm; CR 0.60. Geni- talia (Fig. 2J): Tergite 9 with apicolateral processes very close together, posterior margin rounded, distinctly notched poste- romedially; sternite 9 with moderately deep posteromedial excavation. Gonocoxite twice as long as broad: gonostylus slightly curved, apex pointed. Parameres (Fig. 21) stout, broadly fused at base, apices slender, with minute fringing hairs. Aedeagus with low basal arch, extending 0.20 of total length; lateral arms slightly convex; pos- teromedial projection elongate, apex trun- cated. Female. Head: Dark brown. Eyes bare, contiguous by distance equal to diameter of two ommatidia. Flagellum (Fig. 2F) brown, bases of flagellomeres 1—8 pale; AR 1.15 (1.10—1.18, n = 4); sensilla coeloconica on flagellomeres 1, 9-13. Palpus (Fig. 2G) 366 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1 0.05 mm 0.03 mm Fig. 2. Adult structures of C. hermani (A-E) and C. annettae (FJ): (A, F) female antenna; (B, G) female palpus; (C, H) female spermathecae, duct ring; (D, 1) parameres in ventral view; (E, J) male genitalia in ventral view. VOLUME 106, NUMBER 2 dark brown; third segment slender with capitate sensilla in shallow pit and 2—3 oth- er shallow depressions; long, slender por- tion beyond pit; PR 3.85 (3.70—4.00, n = 4); P/H ratio 0.92 (0.90—0.94, n = 4). Man- dible with 16-17 (n = 4) teeth. Thorax: Scutum dark brown, with sublateral yellow- ish brown patches; scutellum, postscutellum dark brown. Legs dark brown, fore, mid knees, broad apex of hind tibia yellowish; hind tibial comb with five spines, second from spur longest. Wing (Fig. 1B), length 1.02 (0.94-1.14, n = 4) mm; width 0.47 (0.44—0.52, n = 4) mm; CR 0.66 (0.64— 0.67, n = 4); with contrasting pattern; crossvein r-m pale; vein R, pale; transverse distal pale spot in cell r,; barely abutting wing margin; two distal pale spots in cell m, (apical one very faint, nearly impercep- tible in one paratype); distal pale spot in cell m, large, rounded, broadly abutting wing margin; pale spot in cell cua, con- nected with pale line bordering lower mar- gin of vein CuA,; anal cell with two distal pale spots; apices of veins M,, M, pale, api- ces of veins CuA, and CuA, dark. Macro- trichia scattered on distal fourth of wing, absent in cell cua,, anal cell. Halter brown. Abdomen: Dark brown. Two pyriform, un- equal spermathecae (Fig. 2H), measuring 49 (48-50, n = 3) by 41 (38-44, n = 3) wm, and 43 (42—44, n = 3) by 37 (34-40) jzm; rudimentary third, ring present. Pupa. Female exuvia pale yellowish brown. Length 2.30 mm. Body surface gen- erally smooth with spicules present on head, between abdominal segments, dorsal- ly on anal segment. Operculum (Fig. 3D) with OL 0.14 mm; OW 0.16 mm; OW/OL 0.87; with well-developed anteromarginal seta located on tubercle, with pore at tuber- cle base; well-developed medial projection situated near dorsal margin. One long, one shorter anterodorsal setae on short, rounded tubercle (Fig. 3C). One long, one short, slender dorsolateral setae (Fig. 3C). One slender dorsomedial seta (Fig. 3C). Six dor- sal sensilla, i a thick, moderately elongate 367 der seta, v, vi pores (Fig. 3B). Two ventro- median setae minute, slender with bases abutting. Two ventrolateral setae elongate, one longer than other, bases close. Respi- ratory organ (Fig. 3E) brown, pedicel stout; mid third with sparse scale-like pointed spicules, with 9-10 spiracles on apical half; P/H 0.21. Metathorax completely divided but abutting medially, with scutum not pro- truding beyond posterior margin of meta- thorax. Abdomen with setae separate from one another (none on common tubercle). Segment 4 setal pattern (Fig. 3A) with 2 d.a.s.m. with 1 short, 11 elongate; 5 d.p.m. v elongate, slender; l.a.s.m. short, thick spine; 3 |.p.m. with i, 111 moderately sized, thick, 11 elongate, slender; 3 v.p.m. with 1 short, slender, 11, elongate, slender, 111 very short peg. Anal segment (Fig. 3F) length 0.20 mm, width 0.15 mm; dorsal surface with sparse, posteriorly directed tubercles on mid portion, also present on posterolat- eral processes, latter directed posteriorly. Distribution.—Culicoides annettae is re- corded from lowland areas on both the east and west coasts of Costa Rica (Fig. 10A). Bionomic information.—The holotype, allotype and some paratypes were collected with a light trap near the Tarcoles River es- tuary. A single female reared from a pupa was collected from a freshwater spring sit- uated just west of Reserva Carara. The fe- male from 10 km NW Cahuita was collect- ed with a light trap at Aviarios del Caribe, a lodge situated on the delta of the Estrella River on the east coast. Adults were present in December and January. Taxonomic discussion.—Culicoides an- nettae is a member of the guffatus species group within the subgenus Hoffmania Fox. This new species keys out in Wirth and Blanton (1959) to couplet 7 and within that couplet would run to C. diabolicus Hoft- man except that C. annettae has a dark hal- ter knob. The male further differs from C. diabolicus in possessing an aedeagus with a truncated apex. Spinelli et al. (1993) re- viewed the Culicoides guttatius group for 368 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dorsal ventral Fig. 3. Pupal structures of C. annettae. (A) abdominal segment four; (B) right dorsal setae; (C) dorsolateral (dl), dorsomedial (dm), anterodorsal (ad) setae; (D) operculum; (E) respiratory organ, dorsomedial (dm), dor- solateral (dl) setae; (F) anal segment in dorsal view. VOLUME 106, NUMBER 2 the Neotropics and C. annettae keys out to C. ocumarensis in couplet 14 of that work. However, C. annettae may be distinguished from that species by the features mentioned above in the diagnosis. Culicoides paraignacioit Spinelli shares the slender third palpal segment and the truncate apex of aedeagus with C. annettae. However, the third palpal segment of C. pa- raignacioi bears an unique, definite sensory pit and crossvein r-m is clearly dark on its anterior half. Of the 38 Neotropical species of Culi- coides in the subgenus Hoffmania, only four have been described as pupae: C. cha- rruus Spinelli and Martinez, C. hylas Mac- fie, C. insignis Lutz and C. maruim Lutz. The pupa of C. annettae has a more spinose respiratory organ than C. hylas, a respira- tory organ with a row of spiracles distrib- uted along its apical half (apical in C. cha- rruus, C. insignis and C. maruim). The male and female were associated by similar pigmentation patterns and were col- lected together at Tarcoles. Types.—Holotype ¢, allotype 2, Costa Rica, Tarcoles, 1-I-1994, A. Borkent, CD1730 [CNCI]. Paratypes, 3 9, as fol- lows: same data as holotype, | 2 [MLPA]; Costa Rica, 2 km NE Tarcoles, 17-XII- 1993, A. Borkent, CD1714, 1 2 (with pu- pal exuvia) [CNCI]; Costa Rica, 10 km NW Cahuita, 9-XII-1993, A. Borkent, CD1700, ie? {INBC]. Derivation of specific epithet.—This spe- cies is named for the junior author’s wife, who shared the joys and many of the tasks of an expedition to Costa Rica in 1993— 1994 which resulted in the collection of all the type material. Culicoides (Culicoides) hondurensis Spinelli and Borkent, new species (Figs. 1C, 4A—-E 10B) Diagnosis.—Large adults in the subge- nus Culicoides very similar to C. luteove- nus Root and Hoffman, from which it can be distinguished by the irregular palpal pit, the scutum without a definite pattern, the 369 wing background white and the tip of ae- deagus truncate. Description.—Male. Similar to female with usual sexual differences. Flagellum as in Fig. 4A. Wing length 1.38 (1.26—1.52, n = 5) mm; width 0.47 (0.43-0.51, n = 5) mm; CR 0.58 (0.57—0.60, n = 5). Genitalia (Fig. 4F): Tergite 9 rounded posteriorly, apicolateral processes moderately devel- oped, slender, undivided posteromedial lobe well developed, with thickening on midline; sternite 9 short, with broad, shallow pos- teromedial excavation. Gonocoxite with short dorsal, ventral roots, mesal margin with heavy spinose setae; gonostylus lightly sclerotized on distal half, apex blunt, round- ed. Parameres (Fig. 4E): Separate, each with stout, abruptly bent base, stout and nearly straight basal portion, apically slen- der, bent, hairy. Aedeagus triangular; basal arch extending 0.25—0.33 of total length with faintly sclerotized membrane across distal half; lateral arms well sclerotized; apex slender, truncated. Female. Head: Dark brown. Eyes bare, contiguous by distance equal to diameter of 2-3 ommatidia. Flagellum (Fig. 4B) uni- formly dark brown; flagellomeres 1—8 va- siform, 9-13 subcylindrical; AR 0.98 (0.92—1.07, n = 10); sensilla coeloconica on flagellomeres 1, (2—3), (5—8), 9—13 (only one specimen with sensilla on flagellomeres 2-3). Palpus (Fig. 4C) brown; third seg- ment with conspicuous irregular pit; PR 2.82 (2.55—-3.10, n = 10); P/H ratio 0.76 (0.70—0.81, n = 10). Mandible with 14 (13-16, n = 10) teeth. Thorax: Dark brown, scutum without definite pattern. Legs dark brown, knees narrowly yellowish, hind tibia pale at extreme base; hind tibial comb with six spines, second from spur longest. Wing (Fig. 1C), length 1.36 (1.20—1.45, n = 10) mm: width 0.61 (0.58—0.65, n = 10) mm; CR 0.62 (0.60—0.65, n = 10); with white background, pattern roughly in form of three dark zig-zag bands, each more or less broken into separate dark spots centering on veins; distal pale spot in cell r,; occupying entire apex of cell to wing margin; distal 370 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0.1mm 0.1 mm H 0.05 mm | Fig. 4. Adult structures of C. hondurensis (A-F) and C. monicae (G—l): (A) male antenna; (B, G) female antenna; (C, H) female palpus; (E, I) female spermathecae, duct ring; (E) parameres in ventral view; (F) male genitalia in ventral view. VOLUME 106, NUMBER 2 pale spots in cells m, and m, broadly abut- ting wing margin; pale spot in cell cua, large. Macrotrichia numerous on distal % of wing. Halter yellowish. Abdomen: Dark brown. Two pyriform, subequal sperma- thecae (Fig. 4D), each measuring 62 (56— 70, n = 10) by 45 (42-48, n = 10) pm; rudimentary third present. Distribution.—This species is known from a number of localities El Salvador, Honduras (Fig. 1OB), ranging from 690— 991 meters. Bionomic information.—Adults have been collected throughout the year, from January to December. Taxonomic discussion.—This new spe- cies keys out in Wirth and Blanton (1959) to C. elutus in couplet 14. The male differs from that species by the presence in C. hon- durensis of a longer, tapering aedeagus and a more stoutly developed posteromedial projection on tergite 9. The female differs from C. elutus by its larger size, the pres- ence of sensilla coeloconica on flagello- meres 2—3 and 5—8 in many specimens, the third palpal segment with an irregular sen- sory pit, and apex of the hind tibia without a narrow yellowish band. Adults of this species also resemble C. neopulicaris Wirth, a species distributed from Louisiana and Texas in the USA to Costa Rica, by virtue of the irregular palpal pit and wing with white background. Adults of Culicoi- des neopulicaris, however, have a conspic- uous dark spot in the middle of cell cua,, the apicolateral processes of the male are practically absent, and the basal arch of the aedeagus is higher and its tip broadly rounded. The thick spines on the inner surface of the gonocoxite appear to be a synapo- morphy of at least some species in the sub- genus Culicoides. The four Neotropical species C. elutus, C. hondurensis, C. luteo- venus and C. neopulicaris bear these spines, as do the males of all Nearctic species we have examined. The male and female were associated by being collected these together at several lo- 37 cations and on several different dates. There are only three other species of the subgenus Culicoides in the area south of the United States and the fact that both males and fe- males were distinctive corroborates their as- sociation. Types.—Holotype ¢, allotype 2, Hon- duras, Comayagua, Siguatepeque, H-1965, FS. Blanton [MLPA]. Paratypes, 101 @, 13 3, as follows: same data as types, 17 2, 5 G [9 2,.3.¢. MEPA2 2. 2.6 USNs 2 2, INBC; 4 2, CNCI]; same data except IX-1966, 3 2 [MLPA]; same data except EH '967,. 39 be 6 2.027 hai Nie Ae sie CNCI]; same data except 3-VII-1966, J.E Matta, | 2 [MLPA]; same data except 6- VII-1966, | 2 [MLPA]; same data except 16-VII-1966, 3 2 [MLPA]; same data ex- cept 21-VIH-1966, 1 2, 1 6 [MLPA]; same data except 25-VII-1966, 1 2 [MLPA]; same data except 26-VII-1966, 1 @ [MLPA]; same data except VIII-1967, 1 3 [CNCI]; Copan, Santa Rosa, 26-V-1966, J.E Matta, 2 2 [MLPA, CNCI]; same data except 30-VI-1966, | 2 [CNCI]; same data except 7-VIH-1966, 1 2 [MLPA]; same data except 21-VII-1966, 2 2 [MLPA]; same data except 24-VII-1966, 1 2 [MLPA]; same data except X-1966, ES. Blanton, 18 Mee dy iSy 2) Fed sMEPAS 2°25 USNME 3 2, CNCI]; same data except XII-1966, 20 Pohl Ly MERA: 2 SS CNEL SeaNBeE same data except I-1967, 4 2° [MLPA]; Francisco Morazan, Zamorano, 8-VII-1966, J.E Matta, 1 d [MLPA]; same data except 12-VI-1966, 1 d [CNCI]; same data except 22-VII-1966, 1 2 [MLPA]; same data ex- cept 24-VII-1966, 1 2 [MLPA]; same data except X-1966, ES. Blanton, 17 2, 1 ¢ [13 2 MLPA: 2°29) DUSNM? 2.2, fioyCnch El Salvador, San Vicente, Santo Domingo, II-1967, ES. Blanton, 1 2 [MLPA]; Son- sonate, Armenia, III/IV-1967, ES. Blanton, POyrivd IMIEPAY. Derivation of specific epithet.—The name of this species reflects the presence of the majority of the collecting records from Honduras. 372 Culicoides (Anilomyia) monicae Spinelli and Borkent, new species (Figs. 1D, 4G—I, 10A) Diagnosis.—Large adults in the covagar- ciai species group very similar to C. marshi Wirth and Blanton, from which it can be distinguished by the narrowly contiguous eyes, the more slender third palpal segment and absence of a sensory pit, and a longer proboscis. Description.—Male. Unknown. Female. Head: Dark brown. Eyes bare, barely contiguous, V-shaped where eyes contact. Flagellum (Fig. 4G) pale brown, flagellomeres 5-8 bottle-shaped, 9-13 subcylindrical, moderately elongated; AR 0.93 (0.91—0.96, n = 3); sensilla coeloco- nica on flagellomeres 1, 9-13. Palpus (Fig. 4H) dark brown; third segment very slender without sensory pit, sensilla scattered on surface; PR 6.50 (6.00—7.00, n = 3); P/H ratio 1.56 (1.52—1.60, n = 3). Mandible with 28 (n = 3) teeth. Thorax: Specimens not in position to describe scutum pattern. Legs brown; broad yellowish bands apically on fore and mid femora, basally on tibiae; apex of hind tibia yellowish. Wing (Fig. 1D), length 1.57 (1.34-1.77, n = 3) mm; width 0.72 (0.64—-0.80, n = 3) mm; CR 0.64 (0.63—0.66, n = 3); predominantly pale, base broadly pale; crossvein r-m in- cluded in broad pale area abutting costal wing margin, which also includes % of first radial cell; second radial cell, except ex- treme base, included in large pale area; three narrow, transverse, dark bands across wing: one before level of crossvein r-m, one at level of vein R, to cubital fork and a sinuate band from just past apex of costa to tip of vein CuA,; cell r; with subapical, large pale spot; distal pale spot in cell m, separated from wing margin; distal pale spots in cells m,, cua, broadly abutting wing margin; two distal, rounded, narrowly connected pale spots in anal cell; pale spot straddling middle of vein M,. Macrotrichia scattered on distal half of wing, also present in anal cell. Halter pale. Abdomen: Brown. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Two pyriform, slightly unequal spermathe- cae (Fig. 41), measuring 46 (44—48, n = 3) by 38 (36—40, n = 3) pm, and 40 (36-44, n = 3) by 33 (30-36, n = 3) wm; rudi- mentary third present. Distribution.—Culicoides monicae is known from two localities in Costa Rica (Fig. 10A) at mid elevations (1,000—1,800 meters). Bionomic information.—This species is recorded from cloudforest habitat in the Ta- lamanca mountains of Costa Rica and has been collected in March and August. Taxonomic discussion.—Culicoides mo- nicae belongs in the covagarciai species group of the subgenus Anilomyia. It keys to C. marshi in couplet 16 in Wirth and Blan- ton (1959), with the exception that C. mo- nicae lacks the sensory pit on the third pal- pal segment. Wirth and Blanton (1956) re- viewed the Culicoides covagarciai species group for the Neotropics, and the species also keys out to C. marshi in couplet 2 in that work (again with the exception of the sensory pit feature). Of the nine species now recognized in the covagarciai species group, C. monicae and C. marshi have the most elongate third palpal segment. The two are easily distin- guished by examination of the broadly con- tiguous eyes of C. marshi, which are nar- rowly touching in C. monicae. Culicoides efferus Fox also has a somewhat elongate palpus with a small subapical pit and its legs are uniformly light yellow. Types.—Holotype 2, Costa Rica, Carta- go, PN. Tapanti, 1,800 m, VIII-1997, R. Delgado, CD5016 [INBC]. Paratypes, 3 °, as follows: same data as type, | 2 [CNCI]; Costa Rica, Cartago, P.N. Tapanti, 1,800 m, M. Alfaro, CD5043, 1 2 [INBC]; Costa Rica, Puntarenas, San Vito de Java, 22-III- 1964, ES. Blanton, 1 9 [MLPA]. Derivation of specific epithet.—The name monicae is proposed for the senior author’s wife, in recognition of her loving support throughout the past fifteen years. VOLUME 106, NUMBER 2 Culicoides (Anilomyia) chaverrii Spinelli and Borkent, new species (Figs. 1E, SA-I, 6A—H, 7A-B, 11A) Diagnosis.—Medium sized adults in the decor species group very similar to C. lu- tealaris Wirth and Blanton, from which it can be distinguished by the absence of sen- silla coeloconica on flagellomeres 2—5 and 7 in most specimens, the third palpal seg- ment not greatly swollen, the presence of a shallow sensory pit, the elongated gonoco- xite, and the aedeagus with a blunt tip. Description.—Male. Similar to female with usual sexual differences. Wing length 1.24 (1.16—-1.40, n = 5) mm; width 0.48 (0.44—0.56, n = 5) mm; CR 0.57 (n = 5). Genitalia (Fig. 5E): Tergite 9 long, with stout apicolateral processes, the postero- medial margin distinctly notched; sternite 9 short, with broad, shallow posteromedial excavation. Gonocoxite 2.5X as long as broad (basally), gradually tapering posteri- orly, dorsal, ventral roots slender, the later slightly bent apically; gonostylus distinctly shorter than gonocoxite, nearly straight, with pointed recurved tip. Parameres (Fig. 5D) separate, each with bent, knobbed base; basal portion slender, slightly sinuate, ta- pering to very slender, recurved, simple pointed apex. Aedeagus triangular; lateral arms strongly sclerotized; basal arch angu- lose, extending 0.50 of total length; pos- teromedial projection tapering to blunt tip. Female. Head: Dark brown. Eyes bare, narrowly separated by distance shorter than diameter of one ommatidium. Flagellum (Fig. 5A) pale brown, especially flagello- meres 1-8; flagellomeres 1—8 short, 9—13 subcylindrical; flagellomere 9 1.1 as long as 7-8 combined; AR 1.45 (1.23-1.56, n = 10); sensilla coeloconica on flagellomeres 1, (2-5), 6, (7), 8-13. Palpus (Fig. 5B) dark brown; third segment with large subapical deep pit; PR 2.19 (2.02—2.30, n = 10); P/ H ratio 0.69 (0.65—-0.72, n = 10). Mandible with 14 (13-16, n = 10) teeth. Thorax: Scu- tum light brown or yellow, with sublateral brown longitudinal bands, very small dark 373 patch just anterior to margin of scutellum; scutellum yellowish brown, postscutellum dark brown. Legs brown, knee spots black- ish; femora with subapical, tibiae with sub- basal broad pale bands; hind tibial comb with four spines, one nearest spur longest. Wing (Fig. 1E), length 1.37 (1.24—1.52, n = 10) mm; width 0.65 (0.60—0.70, n = 10) mm; CR 0.60 (0.59—0.63, n = 10); appear- ing yellowish with narrow, sometimes bro- ken, transverse dark bands, spots as figured; dark transverse band across midline of cell r, about a third as broad as yellow bands on each side; proximal pale spot in cell r, large, including distal portion of second ra- dial cell, produced beyond vein M.,; distal pale spot in cell r,; separated from wing tip by small dark area; distal pale spots in cells m,, m,, cua,, anal cell broadly abutting wing margin; basal transverse dark band in- terrupted in base of cell m,; rounded pale spot straddling middle of vein M, isolated. Macrotrichia abundant, extending nearly to base of anal cell. Halter pale. Abdomen: Tergites pale, sternites medium brown, pleura blackish. Two pyriform, subequal spermathecae (Fig. 5C), each measuring 54 (48-60, n = 10) by 45 (38-50, n = 10) pm; rudimentary third present. Pupa. Male, female exuvia yellowish brown. Length 2.36 (2.20—2.50, n = 5) mm. Body surface generally smooth with spicules or very small tubercles present on dorsal and ventral surface of cephalothorax, anterolateral margins of abdominal tergites, entire dorsal and anteroventral surface of anal segment. Operculum (Fig. 6A) with OL 0.170 (0.150—0.200, n = 5) mm; OW 0.200 (0.180—0.226, n = 5) mm; OW/OL 1.18 (1.07—1.25, n = 5); central disk cov- ered with abundant rounded tubercles; two well developed anteromarginal tubercles, bearing long, stout seta, with pore at tuber- cle base; well-developed medial projection situated near dorsal margin. Two antero- dorsal setae on short, rounded tubercle, one long, slender, one a short peg (Fig. 5G). Three moderately thick dorsolateral setae, one shorter, one moderate ‘in length, one 374 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dorsal ventral 0.05 mm Fig. 5. Structures of C. chaverrii, adult (A-E), pupa (F—I): (A) female antenna; (B) female palpus; (C) female spermathecae; (D) parameres in ventral view; (E) male genitalia in ventral view; (F) right dorsal setae; (G) anterodorsal (ad) setae; (H) dorsolateral (dl), dorsomedial (dm) setae; (I) abdominal segment four. VOLUME 106, NUMBER 2 elongate (Fig. 5H). One elongate, slender dorsomedial seta (Fig. 5H). Six dorsal sen- silla, i, ili very short setae; 11 elongate, slen- der, iv very elongate, slender seta, v, vi pores (Fig. 5F). Two ventromedian setae absent or perhaps a barely visible single pore. Two ventrolateral setae, one shorter, more slender than other, bases close. Re- spiratory organ (Fig. 6B) golden brown, pedicel very reduced; with scale-like point- ed spicules, along entire length but more abundant on basal half; with 2—3 lateral, 8— 9 apical spiracles; P/H 0.035 (0.020—0.047, n = 5). Metathorax completely divided me- dially, with scutum protruding to posterior margin of metathorax. Abdomen with setae separate from one another (none on com- mon tubercle). Segment 4 setal pattern (Fig. 51) with 2 d.a.s.m. with 1 short, ii very elon- gate; 5 d.p.m. with i very short peg, 11, i1 pores, iv short, thick, v elongate, slender; l.a.s.m. moderately elongate, thick spine; 3 l.p.m. very well developed, with 1, 111 mod- erately sized, thick, 1 very elongate, thick; 3 v.p.m. with i, 111 short, thick, ii, elongate, slender. Anal segment (Fig. 6C) length 0. 20 mm, width 0.15 mm; posterolateral pro- cesses directed posterolaterally. Fourth instar larva. Total length approx- imately 4.00 mm: Head capsule (Figs. 6D— E) yellowish brown, medium-sized, stout with rounded apex; HL 0.252 (0.240- 0.266, n = 6) mm; HW 0.187 (0.180— 0200. n = 6) mm; HR 1.35 (1.20-1.48) n = 6); SGW 0.113 (0.100—0.130, n = 6) mm; SGR 1.66 (1.48—1.80, n = 6); collar somewhat darker than rest of head capsule, well-developed ventrally with short anterior apodeme, separated medially; dorsolateral portion thick, not extending medially. Fron- tal suture extending to near anterior margin of labrum. Ventral suture well developed. Setae all simple, arrangement and sizes as in Figs. 6D-E. Antenna short, details not visible. Eye unknown. Labrum short, wide, further details not discernible. Premandibles small, triangular, apparently unarmed. Man- dible (Fig. 6G) strongly sclerotized; small, broad at base, single stout hooked tooth, 375 medial protuberance slender, subbasal sen- silla present but state uncertain. Epipharynx (Fig. 7A) with only two combs: dorsal comb sclerites short, with seven angular, lanceolate, unequal teeth/sclerite on each side; comb 4 short, with seven pointed teeth; curtains absent. Hypopharynx (Fig. 7B) with slender arms; anterior end short, triangular, with reduced minute teeth. Hypostoma (Fig. 6H) broad, without medial protuberance, with numerous small laterally placed, pointed teeth. Body pigmentation unknown. Caudal segment (Fig. 6F) greatly elongated, with six dorsal and six ventral long, stout, subequal caudal setae; CSL 0.49 (0.44-0.52, n = 3) mm; CSW 0.20 (0.16—0.24, n = 3) mm; CSR 2.46 (2.16— 2.75, n = 3); anal papillae slender, forked. Distribution.—Culicoides chaverrii is re- corded from the Tilaran and Talamanca mountains of Costa Rica (Fig. 11A). The species appears to be a mid to high eleva- tion species, recorded from 1,540—2,270 meters. Bionomic information.—Series of speci- mens have been reared from larvae and pu- pae collected from both arboreal and ground-level bromeliads and a treehole in the Talamanca mountains. Of the seven oth- er Neotropical species in the decor species group, the only two which are known as immatures have been also reared from bro- meliads (Wirth and Blanton 1970). Adults were collected with a light trap in the Ti- laran mountains (at Refugio Bioldgico Monteverde) and adults or reared adults have been collected in June and August. Taxonomic discussion.—Culicoides cha- verrii belongs in the decor species group of the subgenus Anilomyia. The adults of this new species key out to C. lutealaris in cou- plet 21 in Wirth and Blanton (1959), based on leg coloration; the wing length of C. chaverrii is somewhat shorter than that of C. lutealaris. The two species may be dis- tinguished using the features given in the diagnosis above. Culicoides nigrigenus Wirth and Blanton is another similar species which also occurs 376 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0.03 mm 0.01 mm Fig. 6. Structures of C. chaverrii, pupa (A—C), fourth instar larva (D-H): (A) operculum; (B) respiratory organ, dorsomedial seta (dm); (C) anal segment in ventral view; (D) head capsule in dorsal view; (E) head capsule in ventral view; (F) anal segment in ventrolateral view; (G) left mandible in ventral view; (H) hypostoma. VOLUME 106, NUMBER 2 in Costa Rica. However, the hind femur of C. nigrigenus is pale, with a conspicuous dark brown band only on its mid portion, the flagellomere 9 is longer, and the round- ed pale spot straddling the middle of vein M, is connected anteriorly to the poststig- matic pale spots in cell r; and posteriorly to the pale spot in cell cua,. Wirth and Blanton (1970) reviewed the Culicoides decor species group (as the ni- grigenus group) for the Neotropics. Although the immatures of two other species of the decor species group have been found in bromeliads, these have not been described and our description here is the first larval and pupal description of a species within the entire subgenus. The lar- vae have the elongate caudal setae typical of many species which occur in treeholes (Murphree and Mullen 1991). The male and female adults were asso- ciated by rearing these from similar larvae and pupae which were collected from the same habitat. Types.—Holotype d, allotype 2, (both with pupal exuvia), Costa Rica, Puntarenas, Buenos Aires, Potrero Grande, ACLA-P, P. Internac. La Amistad, Est. Altamira, Cerro Frantzius, LS 334150N/574450E, 2,134 m, 20-VI-1999, LGCh-119, G. Chaverri [INBC]. Paratypes 15 ¢ (11 with pupal ex- uviae), 26 2 (8 with pupal exuviae), 7 pu- pae, 7 larval exuviae, as follows: Costa Rica, 2 km E Santa Elena, 26-VIII-1993, 1540 m, A. Borkent, CD1559, 4 2 [CNCI]: Costa Rica, 2 km E Santa Elena, 26-VIII- 1993, 1,540 m, A. Borkent, CD1,560, 1 o, yond. ~ my — = e) Y ae ~ ia] vo 1-6. panamensis. Figs. tenuicornis. EF. schultzet. 6, |e 2; Figs. 7-12. occidentalis. 10, F. panamensis. 11, F. schultzei. 12, F. tenuicornis. from the Netherlands have been brachyp- terous, but more macropterous forms may be encountered in the future. In the U. S. it is a pest of cotton, peanuts, and tobacco. In brachypterous forms of F. fusca, ocelli are vestigial or absent (in brachypterous F. ni- griventris, they are present), and the micro- trichia on the posterior margin of tergite VIII are small and limited to the lateral margins (in brachypterous F. nigriventris, they are well developed across the entire PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Meso- and metanota of Frankliniella species, dorsal views. 7, F. fusca. 8, F. intonsa. 9, F. posterior margin). Macropterous F. fusca resembles F. tenuicornis; the head between the eyes is prolonged anteriorly in both spe- cies, although only weakly so in F. fusca. Macropterous F. fusca differs from F. ten- uicornis in having broad reticulations in the median sculpturation of the metanotum (median sculpturation in E tenuicornis 1s striated); also antennal segments [I-IV in F. fusca is nearly uniformly brown; in F. tenuicornis they are light yellow. VOLUME 106, NUMBER 2 Color.—Body usually brown to dark brown. Range.—(Europe) Netherlands; (North America) U.S.A. (widely distributed); Can- ada, Mexico. (Oceania) Hawaii. Hosts.—Wide variety of plants, including grasses. Frankliniella intonsa (Trybom, 1895) (Figs. 2, 8, 14, 19) Frankliniella intonsa is one of the four intercepted species that often reaches pest status in horticultural crops, especially in the flower industry. It is similar in mor- phology to F. occidentalis but can be dis- tinguished from that species by its shorter antennal segment VIII, its shorter fourth postocular setae, absence of campaniform sensilla on metanotum, and its usually dark- er uniform brown color. Color.—Body light to dark brown; wings clear; legs concolorous with body; head and pronotum light brown to brown. Antennal segments I and II brown; III—V yellow; VI— VIII brown. Range.—(Europe) Widespread through- out Europe. (Asia) China, India, Israel, Ja- pan, Korea, Mongolia, Pakistan, Taiwan, Thailand, Turkey. (North America) U.S.A. [WA only], Canada [British Columbia]. (Oceania) Australia, Philippines. Hosts.—Various species of plants. Inter- cepted from the Netherlands on: Allium sp. (cutflowers), Astilbe sp. (cutflowers), Cro- cosnia/Eremurus sp. (cutflowers), Delphini- um sp. (cutflowers), Gentiana sp. (cutflow- ers), Lisianthus sp. (cutflowers), Lysimachia sp. (cutflowers), Scabiana sp. (cutflowers), Solidago sp. (cutflowers), Viburnum sp. (cutflowers). Frankliniella occidentalis (Pergande, 1895) (Figs. 3, 9, 15, 20-22, 24) The western flower thrips, F. occidentalis is widespread throughout the world and fre- quently reaches pest status on a variety of crops. On the other hand, it also feeds on spider mites, and in California it has been 445 used as part of several insect pest manage- ment programs. It is the second most com- monly intercepted thrips species [Thrips ta- baci Lindeman is the most commonly in- tercepted species]. Although it probably originates in western North America, it has been transported throughout the world and is well established throughout Europe, Af- rica, and the Mediterranean. Frankliniella occidentalis can be distinguished from oth- er Frankliniella species by the following features: Antennal segment VIII approxi- mately twice as long as VII (Fig. 20); post- ocular setae IV significantly greater than one half as long as ocellar setae III (Fig. 3); metanotal campaniform sensilla present; and a complete complement of long micro- trichia evenly spaced on posterior margin of abdominal tergite VIII as in Fig. 15. Sculp- turation of metanotum as in Fig. 9. Color.—Variable, but usually as follows: body yellow with abdominal tergites me- dially brownish; wings clear to light yel- low; legs distally yellow, basally brown or yellow; head and pronotum yellow to light brown. Antennal segment I yellow; II brown; III yellow to light brown; IV—V partly yellow, VI-VIII brown. Range.—(Europe) Widely distributed, in- cluding Italy (It), Netherlands (Ne). (Afri- ca) Canary Islands, Kenya, South Africa, Swaziland. (Asia), Israel, Japan, Korea, Thailand. (Oceania) Australia, New Zea- land. (North America) U.S.A., Canada, Central America, Mexico. (South America) Widespread. Hosts.—Various species of plants. Inter- cepted from material from the Netherlands on: Achillea sp., Ageratum sp., Allium sp. (cutflowers), Alstroemeria sp., Ammi majus, Atriplex sp. (cutflowers), Bouvardia sp., Capsicum sp., Capsicum annuum, Celosia cristata, Chrysanthemum sp., Cystisus sp., Freesia sp., Genista sp. (cutflowers) (It), Ixia sp. (cutflowers), Lisianthus sp. (cut- flowers), Matthiola sp. (cutflowers), Men- tha sp., Ornithogallum sp., peonies, Phlox sp., Rosa sp., Scabiosa sp: (cutflowers), 446 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 13-18. Figs. 15, F. occidentalis. Solidago sp. (cutflowers), Trachelium sp. (cutflowers). Frankliniella pallida (Uzel, 1895) (Figs. 23, 30) Frankliniella pallida has seldom been in- tercepted at U.S. ports-of-entry. Color.—Body pale tan to yellow. Range.—(Europe) Albania, Austria, for- mer Czechoslovakia, Denmark, Germany, Finland, France, Hungary, Italy, Lithuania, Abdominal tergites VIII of Frankliniella species, dorsal views. 13, F. fusca. 14, F. intonsa. 16, F. panamensis. 17, F. schultzei. 18, F. tenuicornis. Netherlands, Poland, Romania, Russia, Swe- den, Switzerland. (Asia) Siberia, Turkistan. Hosts.—Wide variety of flowering plants. Intercepted between 1983 and 1999 on Salvia sp. from Greece [Louisville, KY]. Frankliniella panamensis Hood, 1925 (Figs. 4, 10, 16) Frankliniella panamensis is a Neotropi- cal species that as yet has not been estab- lished in other regions of the world, but VOLUME 106, NUMBER 2 Figs. 19-22. 447 Morphological features of Frankliniella species. 19-20, Antennal segments VII and VIII. 19, F. intonsa. 20, F. occidentalis. 21-22, F. occidentalis; male tip of abdomen. 21, Left dorsolateral. 22, Right dorsolateral. several specimens have been intercepted at U.S. ports-of-entry from the Netherlands. It is regularly intercepted in ports-of-entry in shipments of flowers (especially on Rosa species), fruits, and vegetables from Co- lombia and Ecuador, and it may eventually be inadvertently transported to tropical ar- eas of the Mediterranean and/or Africa. Al- though similar in appearance to F. occiden- talis, it is often darker in color, and the mi- crotrichia on the posterior margin of tergite VIII are greater in length and closer togeth- er than those of F. occidentalis. Color—Body usually brown, with head and pronotum, tibiae, and portion of anten- nae often yellow; wings may be weakly shaded. Range.—(North and South America) Panama, Colombia, Ecuador. Hosts.—Usually on flowers, but also re- cently intercepted on a variety of vegetables and fruits. 448 Frankliniella schultzei (Trybom, 1910) (Higs= 5; diay) Frankliniella schultzei is common throughout tropical regions of the world. Although originally not native to Africa, this species is now well established from Egypt and Morocco to South Africa and is frequently encountered in shipments arriv- ing from various countries in Africa. It is found in nature in two color forms: dark brown and pale yellow. The dark form is known to be a vector of TSWV disease, but in Australia this color form is also known to be beneficial as a predator of phytopha- gous mites (Mound and Kibby 1998). In ad- dition, it is a vector of the South African groundnut ringspot virus and tomato chlo- rotic spot virus (Wijkamp et al. 1995). It is an agricultural pest of a variety of crops, including tomatoes (Nakahara 1997). Frankliniella schultzei can be distinguished from other Frankliniella species by the po- sition of the ocellar setae HI between in- stead of in front of the posterior ocelli (Fig. 5), by the absence of microtrichia on ab- dominal tergite VIII (Fig. 17), and by the absence of metanotal campaniform sensilla. Sculpturation of metanotum as in Fig. 11. Color—Body yellow to light brown; wings clear to yellowish; legs yellow; head and pronotum yellow to light brown. An- tennal segments I and II light brown; III yellow; IV—VIII light brown. Range.—(Europe) Canary Islands, Eng- land, Italy, Netherlands (Africa) Wide- spread, including Cameroon, Chad, Congo, Egypt, Ethiopia, Gambia, Ghana, Kenya, Morocco, Namibia, Nigeria, Senegal, So- malia, South Africa (SA), Tanzania, Ugan- da, Zaire, Zimbabwe, Southwest Africa. (Asia) Widespread, including China, India, Indonesia, Israel, Japan, Korea, Southeast Asia. (Oceania) Australia, Hawaii, Philip- pines. (North and South America) Argen- tina, Brazil, Central America, Caribbean Is- lands, Colombia, U.S.A. [FL only]. (Other) Jamaica. Hosts.—Various species of plants. Inter- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cepted from: Delphinium sp. (cutflowers) (Ne), Lysimachia sp. (cutflowers) (Ne), Mo- raea sp. (cutflowers) (SA), Ornithogalum sp. (cutflowers) (Ne). Frankliniella tenuicornis (Uzel, 1895) (Figss6,02-"13)) Frankliniella tenuicornis is found pri- marily on grasses, although it is not a pest of cereal crops. It is easily distinguished from other commonly intercepted Frankli- niella species by the prolonged vertex of the head in front of the anterior margins of the compound eye (Fig. 6), only antennal segments III and IV yellow, all postocular setae short and often irregular in both num- bers and alignment, and abdominal tergite VIII with irregular number of short micro- trichia on posterior margin (Fig. 18). Sculp- turation of metanotum as in Fig. 12. Color.—Body brown; wings clear; fem- ora brown with bases and apices yellowish; head and pronotum brown. Antennal seg- ments I and II brown; III and IV yellow; V—VIII brown. Range.—(Europe) Austria, Balkans, Denmark, England, Germany, Finland, Ita- ly, Netherlands, Poland, Romania, Russia, Scotland, Sweden. (Asia) China, Israel, Ja- pan, Mongolia, Turkey. (Africa) Libya. (North America) Canada, U.S.A. (Austra- lasian) Australia. Hosts.—Cereals, various grasses, varie- ties of flowers, /ris sp. All interceptions from the Netherlands: Chicorum sp. (cut- flowers), Consolida sp. (cutflowers), [ris sp. (cutflowers), Ornithogallum sp. (cutflow- ers). Frankliniella tritici (Fitch, 1855) Frankliniella tritici is a New World spe- cies recently intercepted twice in material from the Netherlands. Because it is a com- mon species on a variety of North Ameri- can commodities, it may make its way to other countries in Europe, the Mediterra- nean, and Africa, where it may become es- tablished. VOLUME 106, NUMBER 2 449 Figs. 23-29. Morphological features of Frankliniella (24—25, 30—33) and Iridothrips species (23, 26-29). 23, Antennal segments III-IV, /. iridis, showing simple sense cones. 24, 2nd instar larva, F. occidentalis. 25— 27, Abdominal tergite VIII. 25, F. pallida. 26, I. iridis. 27, 1. mariae. 28-29, Head, dorsal view. 28, /. iridis. 29, I. mariae. 450 fe Figs. 30-33. tritici. 33, F. pallida. Range.—(North America) Canada, U.S.A. (South America) Argentina. Hosts.—Found on a wide variety of plants and often a pest species on many ag- ricultural crops. Intercepted from the Neth- erlands on Freesia sp. [Atlanta, GA] and Delphinium sp. (Dallas, TX]. Tridothrips Priesner, 1940 Diagnosis.—-Head as wide as long or longer than wide; antenna 8-segmented; an- tennal segments HII and IV each usually with simple sense cones (occasionally with fork sense cones) (Fig. 23); 3 ocelli present; 3 pairs of ocellar setae, with interocellar se- tae well developed and located laterad of anterior ocellus or only slightly posteriad of anterior ocellus; 4 postocular setae, all short and nearly equal in length; maxillary palpi 3-segmented; pronotum with | short pair of anteroangular, 1 long pair of anteromedial, and 2 long pairs of posteroangular setae and with 2—3 pairs of short anteromarginal and 5 pairs of posteromarginal setae; usually macropterous, with two complete rows of veinal setae regularly distributed on fore- wings; median pair of metanotal setae lo- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON weet et tec ee. 2: 33 Antennal segment II, showing basal pedicel. 30, Frankliniella fusca. 31, F. borinquen. 32, F. cated well posterior of anterior margin; in- tercoxal metathoracic process broad, trun- cate (at least for /. iridis); tarsi 2-segment- ed; ctenidia weakly to moderately defined on abdominal tergites V—VIII, on VII lo- cated anterolaterad of spiracle; abdominal segment X longer than segment IX. Tridothrips iridis (Watson, 1924) (Figs. 23,.26,,28) Although infrequently intercepted, /ri- dothrips iridis occasionally is found asso- ciated with /ris sp. flowers and leaves. It is distinguished from its sister species, /. mar- iae, by the shape of the head (cf Figs. 28 and 29) and body color: /. iridis is usually dark brown, while /. mariae is pale yellow). Color.—Body brown to dark brown with base of head, antennal segments III and IV, foretibiae, and all tarsi yellow; wings pale yellow; femora brown with bases and api- ces yellowish; head and pronotum brown. Range.—(Europe) Austria, former Cze- choslovakia, Denmark, England, Germany, Finland, France, Hungary, Netherlands, Po- land, Scotland, Ukraine. (North America) VOLUME 106, NUMBER 2 Canada [Montreal], U.S.A. [KS, IL, MA, MD, NH, NJ, NY, OH, OR, PA, VA, WA]. Hosts.—A pest of /ris spp.; Kniphofia uvaria; Tritonia uvaria. intercepted from Tris sp. (cutflowers) (United Kingdom). Tridothrips mariae Pelikan, 1961 (Piss: 27, 29) Tridothrips mariae as yet has not been intercepted at U.S. ports-of-entry. Color.—Body pale tan to yellow. Range.—(Europe) former Czechoslovak- ia; Netherlands; Romania; Ukraine. Hosts.—Typha latifolia; Typha angusti- folia. ACKNOWLEDGMENTS I express sincere appreciation to Joseph Cavey, USDA/APHIS, Hyattsville, MD, and Peter Touhey, Systematic Entomology Laboratory, USDA, Beltsville, MD, for providing information for the databases, Sueo Nakahara (retired), and Douglass R. Miller and Michael Pogue, SEL, USDA, for their comments in reviews of the manu- script. Specimens were prepared for SEMs by Suzanne Braden and Scott Whittacker, Scanning Electron Microscope Laboratory, National Museum of Natural History, Smithsonian Institution, Washington, DC. LITERATURE CITED Bagnall, R. S. 1926. Brief descriptions of new Thy- sanoptera. XV. Annals and Magazine of Natural History (9)18: 98-114. Fitch, A. 1855. The wheat thrips and three-banded thrips. Country Gentleman 88: 86-88. Hinds, W. E. 1902. Contribution to a monograph of the insects of the order Thysanoptera inhabiting North America. Proceedings of the United States National Museum 26: 79—242. Hood, J. D. 1925. New species of Frankliniella (Thy- sanoptera). Bulletin of the Brooklyn Entomologi- cal Society 22: 71-83. 1936. Frankliniella gossypiana new name. Proceedings of the Biological Society of Wash- ington 49: 68. . 1942. A century of new American Thysan- optera. III. Revista de Entomologia Rio de Janeiro 12: 547-678. Karny, H. 1910. Neue Thysanopteren der Wiener Ge- 45] gend. Mitteilungen des Naturwissenschaftlichen Vereins an der Universitat Wien 8: 41—5S7. Jacot-Guillarmod, C. E 1974. Catalogue of the Thy- sanoptera of the world. Annals of the Cape Prov- ince Museum 7(3): 517-976. Linnaeus, C. 1758. Systema naturae. Thrips 1. 10th ed. Stockholm, Sweden. p. 407. Moulton, D. 1928. Thysanoptera from Abyssinia. An- nals and Magazine of Natural History (10)7: 227— 248. Mound, L. A. and G. Kibby. 1998. Thysanoptera, An Identification Guide, 2nd Edition. CAB Interna- tional. Wallingford, U.K. 70 pp. Mound, L. A. and S. Nakahara. 1993. The genus Frankliniella (Thysanoptera, Thripidae): Charac- ter assessment at generic and specific levels. Ad- vances in Thysanopterology. Zoology (Journal of Pure and Applied Zoology) 4: 287-295. Nakahara, S. 1997. Annotated list of the Frankliniella species of the World (Thysanoptera: Thripidae). Contributions on Entomology, International 2: 355-389. Nakahara, S. and G. Vierbergen. 1998. Second instar larvae of Frankliniella species in Europe (Thy- sanoptera: Thripidae). Proceedings of the Sixth In- ternational Symposium on Thysanoptera. Antalya, Turkey. April 27—May 1, 1998, pp. 113-120. Nickle, D. A. 2003. A checklist of commonly inter- cepted thrips (Thysanoptera) from Europe, the Mediterranean, and Africa at U.S. ports-of-entry (1983-1999). Part 1. Key to genera. Proceedings of the Entomological Society of Washington 105: 80-99. Pelikan, J. 1961. New species of Thysanoptera from Czechoslovakia—lV. Acta Societatis Entomolo- gicae Czechosloveniae 58: 60—70. Pergande, T. 1895. Observations on certain Thripidae. Insect Life 7: 390-395. Priesner, H. 1920. Beitrag zur Kenntnis der Thysan- opteren Oberosterreichs. Jahresbericht des Muse- um Francisco Carolinum 78: 50—63. . 1938. Thysanopterologica, VI. Konowia 17: 29-35. 1940. Thysanopterologica, [X. Philippine Journal of Science 71: 403. Trybom, FE 1895. lakttagleser om vissa Blasfotingars (Physapoders) upptradande I grasens blomstall- ningar jémte nagra drag ur slaktet Phloeothrips utvecklingshistoria. 60. Entomologisk Tidschrift 16: 157-194. . 1910. Leonhard Schultze, zoologische und an- thropologische Ergebnisse einer Forschungreise im westlichen und zentralen Siidafrika (1903— 1905). Denkschriften medicinisch-naturwissen- schaftllichen Gesellschaft zu Jena 16: 147-174. Uzel, H. 1895. Monographie der Ordnung Thysanop- tera. K6niggratz, BhOmen. 472 pp. Vierbergen, G. 1995. The genus Frankliniella in the 452 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Netherlands, with a key to the species (Thysan- Wijkamp, I., N. Almarza, and D. Peters. 1995. Median optera: Thripidae). Entomologische Berichten, latent period and transmission of tospoviruses Amsterdam 55: 185-192. vectored by Thrips. /n Parker, B. L., M. Skinner, Watson, J. R. 1924. A new Bregmatothrips (Thysan- and T. Lewis, eds. Thrips Biology and Manage- optera) from England and Holland. Entomologist’s ment. Plenum Press, New York. 636 pp. Monthly Magazine 253-254. PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 453-459 A NEW SPECIES OF BRACHYCENTRUS CURTIS (TRICHOPTERA: BRACHYCENTRIDAE) FROM THE SOUTHERN APPALACHIAN MOUNTAINS AND VARIATION IN THE CADDISFLY B. SPINAE ROSS RICHARD C. HARRINGTON AND JOHN C. MORSE Department of Entomology, Soils, and Plant Sciences, Clemson University, Clemson, SC 29634-0315, U.S.A. (JCM e-mail: jmorse @clemson.edu) Abstract.—Description and illustrations for Brachycentrus lunatus, n. sp., from the southern Appalachian Mountains are provided. Intraspecific variation of larval color pat- tern previously has been noted among populations of the caddisfly Brachycentrus spinae Ross. We found a strong correlation between larval phenotype and geographical distri- bution. No significant adult variation was correlated with the larval B. spinae phenotypes. Key Words: Trichoptera, Brachycentridae, Brachycentrus, Brachycentrus spinae, intra- specific variation, southern Appalachian Mountains More than 11,000 species of Trichoptera have been described worldwide (Morse 2001). In North America, only about one- third of the continent’s nearly 1,500 de- scribed Trichoptera species have larvae as- sociated with their corresponding adults (Morse et al. 1997). Trichoptera larvae are known even more poorly in other major global regions (e.g., Morse 2002). Though numerous ecological studies have been con- ducted on Trichoptera species, most have focused on a single population or small number of populations, such that few spe- cies have been comprehensively investigat- ed throughout their range. Comprehensive investigations would more likely provide information concerning geographical asso- ciations and life-history patterns. The southern Appalachian caddisfly Bra- chycentrus spinae Ross exhibits consider- able intraspecific variation. The original de- scription was based entirely on adult males from one population (Ross 1948). When Flint (1984) associated larvae with adult B. spinae, he noted “perplexing” variation, but concluded that the different phenotypes represented a single species, owing to a lack of reared material and no apparent signifi- cant variation among adults. Larval head capsules vary considerably. Head capsules of larvae from the type locality, hereafter referred to as “typical,” have vivid yellow markings on the frontoclypeus and parietal sclerites (Fig. 6). Other larvae, hereafter re- ferred to as “‘dark-headed,”’ have uniformly dark brown or black head capsules that ap- pear more nearly rounded (Fig. 7). Larval variation in B. spinae has continued to per- plex benthologists and taxonomists, as in- traspecific variation to this extent is not fre- quently documented among larval Trichop- tera. The present study attempts to elucidate the perceived intraspecific variation of B. spinae and provides the description of a new Brachycentrus species. METHODS Material for this study was acquired from the Illinois Natural History Survey (INHS), Royal Ontario Museum (ROM), Virginia PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-5. view. 3-5, Male terminalia. 3, Genitalia, left lateral. 4, Tergum X, dorsal. 5, Left inferior appendage, postero- ventral. Brachycentrus lunatus. | Museum of Natural History (VMNH), and Virginia Tech Museum of Natural History (VTMNH) and fresh specimens were col- lected from streams throughout the southern Appalachian Mountains. Specimens were collected in 80% ethanol. Adult terminalia were cleared in a heated potassium hydrox- ide solution. Additional material and para- types are deposited in the Clemson Univer- sity Arthropod Collection, Clemson, SC (CUAC), and the National Museum of Nat- ural History, Smithsonian Institution, Washington, DC (NMNH). 5 \ , Larval head, dorsal. 2, Larval left metathoracic leg, posterior/lateral Brachycentrus (Sphinctogaster) lunatus Harrington and Morse, new species (Figs. 1-5) Adult.—Length of body, including head and folded wings: 11-12 mm. Color: Wings brown, each with pale stigmal spot between radius and subcosta at distal mar- gin. Sclerites fuscous to dark brown. Male genitalia: Preanal appendages elongate, ta- pered, and completely separate mesally in dorsal aspect. Tergum X divided into two lobes in dorsal aspect, each lobe rounded and narrowed apically, with one subapico- VOLUME 106, NUMBER 2 dorsal macrochaeta projecting laterad. In- ferior appendages broad basally, each pro- jecting into sharp point directed mesad. In posteroventral aspect, mesal margin pro- duced into subapical point, creating cres- cent-shaped margin between apical and subapical point. Larva.—Length to 12 mm. Sclerites dark brown and fuscous, marked with pale yel- lowish bands; frontoclypeus dark with well- defined yellow spots anterolaterally and not extending to anterior margin; parietal scler- ites dark, each with pale band posteriorly and between setae 16 and 17, parallel to coronal and frontoclypeal sutures; lateral pale band extending from eye to posterior margin. Femora of meso- and metathoracic legs dark dorsally, fading to yellow ven- trally, their tibiae dark basally and pale dis- tally; each tibia with three to four consec- utively larger setae along basoventral mar- gin. Type material—Holotype, ¢: Eastatoe Creek @ SR 100 (34.996733 N, 82.827783 W), Pickens Co., South Carolina. 20 April 2001, Coll: R.C. Harrington (INHS). Para- types: Same collection data as holotype, 2 3d (CUAC), 3 6 (NMNH), R.C. Harrington. Other material.—North Carolina, Hen- derson County, Green River at Bobs Creek Rd., 10 February 2002, 14 larvae, R. C. Harrington, D. R. Jones (CUAC); same lo- cality but 29 March 2002, 1 larva, 1 pupa, R. C. Harrington (CUAC); same locality but 16 April 2002, 2 pupae, | metamorpho- type, R. C. Harrington (CUAC); McDowell County, Old Fort, at bridge Northwest of town, 1 April 1971, 1 larva, J. C. Morse and F Sherberger (CUAC). South Carolina, Pickens County, Eastatoe Creek @ State Route 100, 14 March 2001, 12 larvae, R. C. Harrington (CUAC); same locality but 8 March 2002, 12 larvae, R. C. Harrington (CUAC); Eastatoe Creek near highway 143, 15 February 1995, 1 larva, 1 pupa, Y. J. Li (CUAC); “Rocky Bottoms Creek @ SR- 39-100" (actually Eastatoe Creek at SR 100), 1 April 76, 1 pupa, J. W. Chapin (CUAC). 455 Diagnosis.—Male genitalia of this spe- cies are very similar to those of Brachycen- trus spinae in the shape of the inferior ap- pendages and tergum X. Only one macro- chaeta is present, however, on each lobe of tergum X (as opposed to numerous macro- chaetae for B. spinae) and the subapical protuberance of each inferior appendage produces only a sharp point and lacks the more basal rounded portion present in B. spinae. Larvae of this species are also very similar to those of B. spinae but can be dis- tinguished by the clearly defined pale mark- ings on the frontoclypeal and parietal scler- ites (as opposed to the less clearly defined pale markings on the frontoclypeal and pa- rietal sclerites of B. spinae). Etymology.—lunatus (Latin) = crescent shaped, referring to the general appearance of the apical end of each inferior append- age. Notes.—This species is a member of the Brachycentrus numerosus species group (Flint 1984) and appears to be a sister spe- cies of Brachycentrus spinae Ross. It has been collected only from streams in upper Atlantic Coast drainages of the Appalachian Mountains. Brachycentrus spinae Ross (Figs. 6—12, 13) Populations of Brachycentrus spinae were encountered throughout the entire southern Appalachians, conforming almost completely to the southern Appalachian ecoregion (Fig. 13). “Typical” larvae (Fig. 6) were distributed along the western edge of the Appalachians, primarily along the Tennessee—North Carolina border, and in North Carolina in the vicinity of the Ten- nessee—Virginia border. Uniformly dark- headed individuals (Fig. 7) were found throughout the interior southern Appala- chians of North Carolina and northern Georgia. About 80% of dark-headed popu- lations had specimens with a mottled ap- pearance due to muscle scars along the pos- terior portion of the parietal sclerites. The degree of darkening also varied among pop- 456 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 12 Figs. 6-12. Brachycentrus spinae. 6-8, Larval heads, dorsal view. 6, “Typical” larva (from type locality— Camp Creek, Greene County, Tennessee). 7, “‘Dark-headed” larva from French Broad River (Transylvania County, North Carolina). 8, ‘‘Intermediate’’ larva from Rock Creek (Fannin County, Georgia). 9, Larval left metathoracic leg, lateral view (from type locality—Camp Creek, Greene County, Tennessee). 10-12, Male terminalia. 10, Genitalia, left lateral. 11, Tergum X, dorsal. 12, Left inferior appendage, posteroventral. VOLUME 106, NUMBER 2 Fig. 13. ulations, ranging from uniformly brown to nearly black. A cluster of populations from northern Georgia presented an intermediate larval color pattern. These intermediates possessed nearly completely dark head cap- sules with small pale markings on the fron- toclypeus and parietal sclerites (Fig. 8). Not all populations were sampled in both the larval and adult stages. However, for those population for which adults were as- sociated with larvae, adults that we consid- er B. spinae did not exhibit differences or trends distinct enough to warrant descrip- tion. In general, adults from both the dark- headed and typical populations possessed numerous macrochaetae on each lobe of tergum X (Fig. 1). Also, inferior append- rachycentrus spinae e phenotypes and allies B. spinae, typical B. spinae, dark-headed, uniformly brown B. spinae, dark-headed, nearly black B. spinae, "intermediates" B. lunatus, new species 50 Kilometers ee Distribution of Brachycentrus spinae larval phenotypes and Brachycentrus lunatus. ages from adults of both larval phenotypes had the same general appearance: a sharp apical point, a subapical point, and more basal “‘chin-like’’ protuberance (Fig. 12). On the other hand, although larvae of pop- ulations along the eastern Blue Ridge Es- carpment bore a strong resemblance to typ- ical populations of B. spinae, consistent dif- ferences in larval and adult characters pro- vide evidence that they represent a distinct species described above. Variation has been reported for several other Trichoptera, including Hesperophylax species (Parker and Wiggins 1985), Rhy- acophila nigrita Banks (Prather and Morse 2001), Arctopsyche grandis (Banks) (Hauer and Stanford 1981), Hydropsyche contuber- 458 nalis McLachlan (Guinand et al. 1997), Di- plectrona modesta Banks (Morse and Barr 1990), and Cheumatopsyche harwoodi enigma Ross, Morse, and Gordon (Gordon 1974). Despite the observed moderate lar- val or adult morphological variation in each situation, none of these studies concluded that multiple, distinct species exist. The un- usual contrast between dark-headed and typical larvae of Brachycentrus spinae strongly suggested a complex of at least two species. From among the larval phe- notypes that we eventually segregated, only B. lunatus had corresponding diagnostic characters in males. We found no corre- sponding adult characters to discriminate the other phenotypes in the adult form. Fur- thermore, the larval forms are allopatric, preventing inferences about natural repro- ductive isolation. The typical phenotype is restricted to the western edge of the south- ern Appalachians, the dark-headed pheno- type in the interior southern Appalachians (with the darkest larvae occurring in the southern and eastern parts of the range), and “intermediates” in the extreme south- em extent of the range (Bie, 13). The taxonomic significance of “‘intraspe- cific’? variation between dark and typical Brachycentrus spinae remains open to in- vestigation, particularly one that uses a mo- lecular population genetics approach. Other studies on organisms inhabiting the south- ern Appalachians have also found patterns of substantial variation that correlate with geography (e.g., Nalepa et al. 2002). The orientation and situation of the southern Appalachians might have once provided a refuge for northern species and served to fragment populations of less mobile south- ern species during peaks of Pleistocene gla- ciation (Hack 1969). We suspect that Bra- chycentrus spinae has low vagility in its adult phase due to its ephemeral adult life span and therefore may be a species that is particularly vulnerable to range fragmenta- tion. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ACKNOWLEDGMENTS Thanks are extended to Chuck Parker (USGS) for his valuable comments. We are grateful to Dan Jones (Clemson University) for assisting in the field collection of much of the new material. Thanks to Karl Kjer (Rutgers University), Karen Byrson, and Yayi Kusumah (Clemson University) for providing much guidance during the at- tempted molecular laboratory work, and Christy Geraci (Clemson University) for help with the map. Colin Favret (INHS), Brad Hubley (ROM), Reese Voshell (VTMNH), and Richard Hoffman (VMNH) and their institutions loaned specimens for this investigation. Funding was provided by an E. W. King Grant from the Department of Entomology, Clemson University. For providing critical review of this manuscript, we thank Peter Adler and Al Wheeler (Clemson University). This is Contribution Number 4859 of the South Carolina Agri- culture and Forestry Research System, Clemson University. LITERATURE CITED Flint, O. S., Jr. 1984. The genus Brachycentrus in North America, with a proposed phylogeny of the genera of Brachycentridae (Trichoptera). Smith- sonian Contributions to Zoology 398, 58 pp. Gordon, A. E. 1974. A synopsis and phylogenetic out- line of the Nearctic members of Cheumatopsyche. Proceedings of the Academy of Natural Sciences of Philadelphia 126: 117—160. Guinand, B., S. Champley, A. Clermidy, and H. Tach- et. 1997. Investigation of within and between sam- ple colour pattern differentiation in Hydropsyche contubernalis (Trichoptera). Biological Journal of the Linnean Society 61: 223-241. Hack, J. T. 1969. The area, its geology: Cenozoic de- velopment of the southern Appalachians, pp. 1— 17. In Holt, P. C., R. L. Hoffman, and C. W. Hart, Jr., eds. The Distributional History of the Biota of the Southern Appalachians. Part I: Invertebrates. Research Division Monograph 1, Virginia Poly- technic Institute, Blacksburg, Virginia. 295 pp. Hauer, E R. and J. A. Stanford. 1981. Larval special- ization and phenotypic variation in Arctopsyche grandis (Trichoptera: Hydropsychidae). Ecology 62: 645-653. Morse, J. C. 2001. Biodiversity, pp. 9-11. Jn Carswell, L. and P. Robinson, eds. Conserving South Car- olina: Sustaining and Protecting our Natural Re- VOLUME 106, NUMBER 2 sources. The Nature Conservancy, Columbia, South Carolina. 44 pp. . 2002. Following a dream, pp. 3—7. Jn Bae, Y. J., ed. The 21st Century and Aquatic Entomolog in East Asia: Proceedings of the Ist Symposium of the Aquatic Entomological Societies of East Asia. Korean Society of Aquatic Entomology, Seoul, Korea. 146 pp. Morse, J. C. and C. B. Barr. 1990. Unusual caddisfly (Trichoptera) fauna of Schoolhouse Springs, Lou- isiana, with description of a new species of Di- plectrona (Hydropsychidae). Proceedings of the Entomological Society of Washington 92: 58—65. Morse, J. C., B. P. Stark, W. P. McCafferty, and K. J. Tennessen. 1997. Southern Appalachian and other southeastern streams at risk: Implications for may- flies, dragonflies and damselflies, stoneflies, and caddisflies, pp. 17—42. In Benz, G. W. and D. E. Collins, eds. Aquatic Fauna in Peril: The South- 459 eastern Perspective. Special Publication 1, South- east Aquatic Research Institute, Lenz Design and Communications, Decatur, Georgia, 554 pp. Nalepa, C. A., P. Luykx, K. D. Klass, and L. L. Deitz. 2002. Distribution of karyotypes of the Crypto- cercus punctulatus species complex (Dictyoptera: Cryptocercidae) in the southern Appalachians: Relation to habitat and history. Annals of the En- tomological Society of America 95: 276-286. Parker, C. R. and G. B. Wiggins. 1985. The Nearctic caddisfly genus Hesperophylax Banks. Canadian Journal of Zoology 63: 2443-2472. Prather, A. L. and J. C. Morse. 2001. Eastern Nearctic Rhyacophila species, with revision of the Rhyaco- Phila invaria Group (Trichoptera: Rhyacophili- dae). Transactions of the American Entomological Society 127: 85-166. Ross, H. H. 1948. New species of sericostomatid Tri- choptera. Proceedings of the Entomological So- ciety of Washington 50: 151—157. PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 460-471 PRIMARY CONSUMER AND DETRITIVORE COMMUNITIES (DIPTERA: EPHYDRIDAE) IN NEWLY RESTORED AND CONSTRUCTED WETLANDS BRUCE A. STEINLY Miami University, Zoology Department, 212 Pearson Hall, Oxford, OH 45056, U.S.A. (e-mail: steinlba@muohio.edu) Abstract.—Although maximum diversity of higher Diptera has been reported in the interface between wetlands and other ecosystems, few studies have documented successful restoration of wetland insects within these ecotones. The consistent collection of shore flies (Diptera: Ephydridae) supports the hypothesis that shore flies rapidly colonize newly restored and constructed wetlands at Miami Trace and Winton Woods county Parks, Ham- ilton County, Ohio. Rapid colonization of shore flies exhibited a temporal shift that is associated with aquatic vegetation colonization, growth, and maturation. Species com- position of shore-fly communities suggests that a minimum of five general nutrient sources were exploited in newly established wetlands, and shore flies may be an essential part of the establishment of complex food webs. A comparison of Sorenson and diversity indices suggests that shore-fly communities in constructed and restored Ohio wetlands were dis- tinct species assemblages. Significant differences among shore-fly communities are attri- buted to variation in species abundance. Although restored and constructed wetlands were flooded during 1998, variation in species abundance and monthly species accumulation suggests that the development of ecosystems within each wetland was asynchronous. Key Words: food webs In many regions of the United States, the restoration of wetlands has been initiated to provide habitat for vertebrate species. The success of bird, amphibian, and mammal recolonization in wetlands is directly relat- ed to the type, quality, and abundance of food resources within wetland ecosystems (Wilson 1987). One of the major sources of food for waterfowl, shore birds, and fish species is insects (Martin and Uhler 1939, Zahl 1967, Clarke 1976, Murkin and Batt 1987). During egg laying and brood rear- ing, adult and juvenile ducks consume in- vertebrate food (including insects) (Krapu and Swanson 1975, Bataille and Baldassar- re 1993) to obtain the protein that is nec- restored wetlands, Diptera, Ephydridae, colonization, community diversity, essary for reproduction and rapid growth (Driver et al. 1974). Decreased availability of food has been associated with duckling mortality (Johnson et al. 1992, King and Brazner 1999), and duckling brood avoid- ance and abandonment of wetlands that have small numbers of invertebrates (Coo- per and Anderson 1996). Although the im- portance of insect resources to waterfowl development has been recognized, an un- derstanding of the contributions of insects to wetland food webs is limited (Rosenberg and Danks 1987, Batzer and Wissinger 1996, Hansen and Castelle 2000). As Batz- er and Wissinger (1996) stated, many of the previous experiments and assumptions con- VOLUME 106, NUMBER 2 cerning wetland insect ecology require re- evaluation to improve the management of insect resources as waterfowl food. Foun- dations for regulating insect and waterfowl communities are in the initial stages of de- velopment (Batzer and Wissinger 1996). Previously, a majority of wetland inves- tigations have focused on plant communi- ties, soils, hydrology, chemical processes, benthic invertebrates, and vertebrate popu- lations, such as waterfowl (Sharitz and Batzer 1999). Although maximum diversity of higher Diptera is at the interface between mature wetlands and other ecosystems (Deonier 1965, LaSalle and Rozas 1991, Marshall 1994, Scheiring and Foote 1973, Steinly 1986, Thier and Foote 1980, Keiper et al. 2002), and the greatest production of insect biomass is found in sparse emergent vegetation zones (Voigts 1976, Orains 1980, Kaminiski and Prince 1981, Mc- Lauglin and Harris 1990), only a few in- vestigations have focused on the successful restoration of wetland macroinvertebrates that are found in the water column and/or sediments (Danell and Sjoberg 1982, Flor- ida Department of Environmental Protec- tion 1994, Brown et al. 1997). The attrac- tion of shore flies to artificial pools and oth- er habitats in a constructed wastewater treatment wetland at 2 days post-flooding and the establishment of shore-fly popula- tions suggest that colonization by these spe- cies was not incidental (Keiper and Walton 2002, Keiper et al. 2002). Although two in- vestigations have focused on Diptera that inhabit soil and consume decaying vegeta- tion in restored Florida (Streever et al. 1996) and Washington (Hansen and Cas- telle 2000) wetlands, studies of primary consumer richness or abundance have not been reported. Layton and Voshell (1991) suggested that an increase in habitat diversity would most likely lead to increases in invertebrate di- versity within wetland ecosystems. It is common for detritus-feeding insects (1.e., insects that feed on decaying plant and an- imal tissues) to dominate areas that contain 461 newly flooded decaying organic materials (Layton and Voshell 1991). However, as an area matures, plant detritus decreases and production of unicellular and multi-cellular algae and macrophytes increases (Layton and Voshell 1991) resulting in increased primary consumer abundance and species richness. Although Batzer and Wissinger (1996) did not monitor primary consumer communities, their review suggests that de- tritivore communities are not affected by macrophyte and algal growth. They suggest that detritivore populations either remain constant or increase where multi-cellular plants were cut and debris removed. In this paper, evidence is presented to document the initial colonization of newly restored and constructed wetlands by shore- fly species that have been associated with the consumption of detritus and primary production. Temporal shifts in the richness and abundance of primary consumers and detritivores 1.e., shore flies are associated with the growth of macrophytes and matu- ration of newly established wetlands. Shore-fly diversity, relative abundance, richness, and similarity values from these wetlands are compared. Diversity values are compared with a f-test to identify signifi- cant differences in shore-fly communities. MATERIALS AND METHODS During 1997, the Hamilton County Park District completed the restoration of a sin- gle wetland at Miami Trace, and the con- struction of two wetlands at Winton Woods, Hamilton County, Ohio. At Miami Trace, the renewed flooding represents a restora- tion of a wetland that was surveyed and de- scribed before the setthkement of Ohio. A single wetland was leveed at Miami Trace County Park without basin alteration, and is located northwest of New Haven and ap- proximately, 0.55 km east (39°17.3'N, 84°44.8’W) of a larger restored wetland complex. Before wetland restoration, the area was covered with old-field vegetation and was well drained. Post-restoration veg- etation consisted of scattered stands of Ty- 462 pha latifolia L. and patches of filamentous algae that were surrounded by unvegetated mud shores. The Winton Woods County Park (Winton Woods, Ohio) sites consist of two basins on the northern edge of the park that are named Mallard (39°15.9’N, 84°31 W) Scand “Heron /2(39° 15:9" N; 84°30.9'W) Wetlands. These wetlands were constructed to control precipitation run-off from the surrounding watershed and to pro- vide habitat for vertebrate species. Mallard Wetland was bordered on the south by a steep grass shore while the wetland interior contained scattered stands of 7. latifolia mixed with Sparganium americanum Nut- tall, Alisma subcordatum Raf., and Carex. Heron Wetland had a similar grass shore on the south side that was well drained. The shallow slope of the northern shore of Her- on Wetland promoted the retention of soil water that sustained the dense growth of sedges and scattered Sagittaria latifolia Willd. on the shoreline. Emergent Ranun- culus flabellaris Raf. occupied two thirds of the wetland pool. Although trace amounts of precipitation fell during July—September, the Winton Woods sites held water at a fair- ly constant level until mid August. During late August through September, areas of limestone sediment in both Winton Woods wetlands were exposed that was covered with sparse plant debris. As the season pro- gressed, larger areas of sediment were ex- posed in Mallard Wetland and these mud shores were colonized by widely scattered clumps of Setaria glauca Kuntze (yellow foxtail), grass (-es), and Sp. americanum, while Carex sp. and Sa. latifolia grew on newly exposed mud shore at Heron Wet- land. Visual evidence of leaf mining con- firmed that pioneering grasses were colo- nized by dipteran primary consumers. Bi-weekly collections of shore flies were initiated at Miami Trace and Winton Woods in May and continued through October 1998. Wetlands were subdivided into habi- tats that were characterized by substrate and vegetation types and sampled with a mod- ified aerial sweep net (150 net sweeps per PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON habitat) (Scheiring and Foote 1973; Re- gensburg 1976; Steinly 1978, 1986, 1990, 2001). One sweep sample is the combina- tion of a back and forth movement of the net through a habitat. Sweep net sampling is a quick and inexpensive means of sam- pling diverse communities of invertebrates (Murkin et al. 1983, Cheal et al. 1993). The size of sampling areas and the number of net sweeps were equal among _ habitats. Plant debris was removed immediately from the samples and stored temporarily in petri dishes. Mounted and unpinned speci- mens were identified, labeled and counted. Mounted and vialed voucher specimens are deposited in the Miami University Insect Collection. The percentage relative abundance is cal- culated with the formula R. A. = A,/N X 100, where R. A. is the percentage relative abundance, A, is equal to the abundance of each species, and N is the total number of shore flies within the wetland. The percent- age ranges are characterized as follows: 1— 2% rare (r), >2—8% occasional (occ), >8— 14% common (c), >14—25% abundant (a), and >25—100% very abundant (va) (Deon- ier 1965; Scheiring and Foote 1973; Deon- ier and Regensburg 1978; Steinly and Deonier 1980; Steinly 1984, 1986, 1990). The Shannon-Wiener diversity index (H’) was calculated for all wetlands because it incorporates both species richness and abundance (Scheiring 1974). Shore-fly di- versity (H’) values for each wetland were compared with a f-test to identify differenc- es in shore-fly communities (Zar 1984). Di- versity is calculated by: H’ = — p; logyop; where p,; is n/N, n, is the number of indi- viduals of the ith species of the habitat be- ing considered, and N is the total number of individuals per habitat. H’ is essentially dimensionless and usually not affected by sample size (N) (Olive and Dambach 1973, Scheiring 1974). Newly restored wetland habitats of comparable area were sampled for approximately the same amount of time and probable differences in richness and abundance of shore-fly species reflect bio- 463 VOLUME 106, NUMBER 2 ‘soroadg aIOANLNAG UMOUY x. 8187= COS 66 LOEE = [P10], =— a aa J (691) OI I (L9'°0) ELE (UOISI[[IMA) Sadan sos1az I (LOT) 9 9 (Line B) STI J (80) oI 4 M207T DAD vdojIsds ] Jd (81°O) I = ae = —— a —_— (M207) SUAMIAOAID DILIIDIAS J (er 1) 8 B (6r SI) Lvl I (€€'7) fifi (ug|[e4) SIVUsYIS D]]aIDIS J (9¢°0) G — —_ — a (STO) (s (UasIBI) WnpnyDpd Y]]aAIvIg J (FIT) cl = = = J (170) fl MQO'T VJAJOSgO YIJAIVIS . 990 (Opt) (Sie == = == = = — M20'7T DaIvD]]IAD{ D]]aIVDIS — — — I (ZE'0) € I (730) LZ NopiInbog vusiua vidwojud 1 (PSO) c Say = == J (770) Q uossaig vidnp vdojisg I (29'T) CI 990 (6L'€) O¢ 990 (80'S) gg x (M207) SIig4o vAoydoyojAjog 990 (C79) cE J (110) I = = = (M20°J) Sijigap DiUSA]IYd i (S10) I = > = = == MOOT DINJDIUNA] DUILAd i (ILO) 1 J (TE'O) € I (O€ 1) Pp M207] VIDINIAAGnILpONh vApKivd == => = J (YAS ]0) el I GED) jis M207 SdadiAasg DAPAIDG = a = J (€¢'0) ¢ I (O¢'O) Ol (ug|[e4) vpinbv vapKivg J (96'T) Il 990 (TES) 6L I (6€ 7) 6L (UUPLUSpel AA) SIUUadyound puuijppiévd I (ILO) 14 = a ie J (90'0) G UaASNR|D soylolwuY vAIYIYIO — — = I (117) OT I (-7 1) Iv 4 Sle, sisdoapyd pjiydyonyy — — — I GS) TZ J (cO'1) Pe 4SIWeY, Vinosnnd vj1ydiyoyy == — == J (€9'(Q) 9 J (90'0) G 4¥UOSSIIZD 1Ma0] DjIYdIONy J (PSO) € I (C60) 6 I (90'0) G ,SIWIRIY Visnpy vpiydyoNy ra ae aa J (780) 8 I (60'0) € UOSSAID S1D]JAINIS DUISON == = = J (+780) g I (€Q'O) | (A9TSOU AA PUR JURADLINIG) VIVAVDIXA AAISDSOIAT —~ = a a 5 F I (€0'O) I y(M207J) DudUIAD DdojIsdojdaT J (SO) € a ad = I (€L°0) 1 £6 y(M2OT) WnjI0AdOINa] DUSDYIOAPAH] ) (9t'6) €S 990 (06'r) LY 990 (iSES) OII UOSSAID SyVIgU VIYJaAPAH rc ag = a a a d (€O'O) I M207T VIVIYISI DIJAAPAH 2 (LOT) o9 S) (CL:O1) TOI 990 (prs) 6LT (Ug|[V4) VjOasl4s vIjaipAHY a Zz — J (PL'0) Mk =F = = M207] VsOWsof D1I]AAPAH] a ae ae I (110) I I (€0'0) I (uOSSaID) VsouInsd DUIpDAYY d (7S'0) € Jd (971) Cl J (€O'O) | (UOSSOID) VIDJOUIG DUIPKAY x 7 ea 990 (69°¢) cE = oe = yapinbogd vsouaaogy vuipyay J (9€°0) (6 =e oe 3 J (€0'0) | yUOSSA pnsixa vaoydoyriq 990 (LS) (OE B (S907) 961 BA (O1'S9) €SIZ y (UP) DIJAANISGO VUIAAIOISIG a = = I (110) I I (O€'1) 'Shvg yUOSSAID SUAJIUOAUUNAG DUIAIIOISIG = =e = I (TE'O) € d (900) G x (US][BR4) VDIDPNDI VJavYyIIG BA (67 6b) jute 990 (79'p) tr 990 (Sv'7) 18 4(MQO7J) Xajduus DWUOYILMIO]] Y (6) VU Jaquinyy (™Z) Vou JOquUINN (%) ‘VU JOquUInN aoRly, TUR purple pasoisay SPOOAA UOIUIAY purples uo1dH SPOOAA UOIUT AA puBpoA PsP] “‘SpueTIOM OLYO pPo1oj}sol pue pajyonysuos ATMauU UI ddURpUNQeR dATLIeI puke soURpUNGe saloads AY-a10Y4S “| 2IqGeL 464 logical differences among wetlands (Scheir- ing 1974). The community composition of each wetland was compared by means of the So- renson Index of Similarity (I). Similarity is calculated with the formula I = 2 C/(A + B), where I is the index of similarity, C is the number of species shared, A is the num- ber of species in habitat A, and B is number of species in habitat B (Scheiring and Deonier 1979, Steinly 1984). The Sorenson index ranges from 0 when there is no sim- ilarity (no species shared) between habitats to 1 when there is complete similarity. RESULTS During May through October of 1998, 4,818 shore flies were collected from the newly restored and/or constructed wetlands within Miami Trace and Winton Woods County Parks, respectively (Table 1). Col- lections from Mallard and Heron wetlands at Winton Woods, and the restored wetlands at Miami Trace contained 31, 26, and 21 (Table 1) species of shore fly within sam- ples of 3,307, 949, and 562 specimens, re- spectively (Tables 1—4). Of the total num- ber of shore flies, detritivores accounted for approximately 78, 49, and 60% of the ephy- drid communities within Mallard, Heron, and Miami Trace wetlands, respectively (Table 1). Approximately 86.7%, 1.e., 4,177 of the total number of shore flies collected during the season from all wetlands con- sisted of Allotrichoma simplex (Loew) (7.7%), Discocerina obscurella (Fallén) (48.8%), Hydrellia griseola (Fallén) (9.2%), Hydrellia tibialis Cresson (4.5%), Paralimna_ punctipennis (Wiedmann) (3.5%), Polytrichophora orbitalis (Loew) (4.6%), Scatella stagnalis (Fallén) (4.82%) and Typopsilopa atra Loew (3.1%). The abundance of these species ranged from oc- casional (occ) to very abundant (va) (Table 1). All other species were rare (r) and were represented by fewer than 100 individuals (Tables 1—4). May samples from the Heron and Mallard wetlands (Tables 2 and 3) and October collections from Miami Trace (Ta- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ble 4) did not contain shore-fly specimens (Fig. 1). Although initial collection of shore flies from Mallard and Heron wetlands started in June and July, respectively, spe- cies accumulation patterns are similar (Fig. 1). Shore-fly species accumulation in the Miami Trace wetland started in May and approached asymptote during September (Fig. 1). In the Mallard and Heron wetlands, the abundance of Allotrichoma simplex, Poly- trichophora orbitalis, and Discocerina ob- scurella, increased during August and Sep- tember (Table 2 and 3), respectively, while A. simplex and Hydrellia griseola appeared in May collections and abundance contin- ued to increase through July in the restored wetlands at Miami Trace (Table 4). AIl- though a single Po. orbitalis adult was col- lected in May in the restored wetland, ad- ditional specimens were not captured until July (Table 4). A. simplex and Po. orbitalis consume decaying organic matter (Foote and Eastin 1971, Foote 1995). The abundance of two leaf-mining spe- cies, Hydrellia griseola and Hydrellia tibi- alis, and T. atra increased dramatically dur- ing July through August, and September in Miami Trace and Winton Woods wetlands, respectively (Tables 2—4). Also, appreciable numbers of Paralimna punctipennis and Scatella stagnalis were found in Mallard and Heron wetlands (Tables 2 and 3) during September. Sc. stagnalis, and Pa. puncti- pennis feed on cyanobacteria and diatoms, respectively (Foote 1995). Hyadina albov- enosa Coquillett, Hyadina binotata (Cres- son), Hyadina pruinosa (Cresson), and Hy- drellia formosa Loew were rare (r) species (Table 1) and collected early in the season from grass shore habitat at Heron Wetland (Table 3). Although relatively large num- bers of Parydra (Table 5) and Notiphila species were routinely collected in aquatic habitats and mature wetlands in Ohio (Steinly 1978, Todd and Foote 1987, Lar- son and Foote 1997), respectively, Notiphi- la adusta Mathis, Notiphila loewi Cresson, Notiphila pauroura’ Mathis, Notiphila VOLUME 106, NUMBER 2 Table 2. Shore-fly community of Mallard Wetland at Winton Woods County Park. SS June July Allotrichoma simplex Dichaeta caudata Discocerina brunneonitens Discocerina obscurella 3 Ditrichophora exigua Hyadina binotata Hyadina pruinosa Hydrellia griseola 1 Hydrellia ischiaca Hydrellia tibialis Hydrochasma leucoproctum Leptopsilopa atrimana Lytogaster excavata Nostima scutellaris Notiphila adusta Notiphila loewi Notiphila pauroura 2; Notiphila phaeopsis Ochthera anatolikos Paralimna punctipennis Parydra aquila Parydra breviceps Parydra quadrituberculata Polytrichophpora orbitalis Psilopa dupla Ptilomyia enigma Scatella obsoleta Scatella paludum Scatella stagnalis Typsilopa atra Zeros flavipes Total = 12 4 Nn £& phaeopsis Mathis, Parydra aquila (Fallén), Parydra breviceps Loew, and Parydra quadrituberculata Loew were rare (r) in all newly established wetlands. Additionally, Scatella favillacea Loew, Scatella obsoleta Loew, Scatella paludum (Meigen), and Sc. stagnalis were found on filamentous algal mats in the spring at Miami Trace, but the habitat was submerged by a mid-summer torrential rainfall and Scatella species and Setacera atrovirens (Loew) were not found after 18 August. In contrast, Sc. obsoleta and Sc. paludum were collected from Mal- lard Wetland at Winton Woods in late sum- mer (Table 2). A comparison of Mallard and Heron shore-fly communities with the Sorenson Aug. Sept. Oct. Total N7/ 43 21 81 = — 2 = 2 5) 29 9 43 eS 1116 839 PASS: a= 1 _ = 1 _ 1 aa = I = = _ 1 27 149 94 = 279 1 aS _ = 1 _ 99 17 = 116 an _ 24 24 ss as = | = Le = - 1 = 1 2 = 3 2 a = 2 = 1 = = 2 -_ 23 3 34 8 33 — = 41 = 1 = 2 6 51 22 = 79 4 3 — 10 6 43 = = 57 39 = — - 43 —_ 2 16 168 2 2 _- = 8 1] 16 = DT = 7 = 7 a= 5 = 5) 4 63 10 do 1 11 — = 16 3 15 4 = 22 Ol 1848 1092 = 3307 index suggests that the constructed wet- lands contained similar assemblages (Table 6) while comparison of Miami Trace with both Winton Woods localities yielded lower similarity values (Table 6). Community di- versity values (H’) of 1.10 (Heron), 0.83 (Miami Trace), and 0.67 (Mallard) com- pared with a f-test suggest that shore-fly communities within each wetland were sig- nificantly different. DISCUSSION Monthly increases in A. simplex, D. ob- scurella, and Po. orbitalis abundance (Ta- bles 2—4) suggest that detritivore coloniza- tion, continuous immigration, and/or the growth of newly established’ communities 466 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Shore-fly community of Heron Wetland at Winton Woods County Park. July Aug. Sept. Oct. Total Allotrichoma simplex l — 41 2 44 Diachaeta caudata — — oo 3 = 3 Discocerina brunneonitens — = 1 — = 1 Discocerina obscurella — 2 39 155 = 196 Hyadina albovenosa 34 — 1 — = 35 Hyadina binotata 10 — 2 — = 12 Hyadina pruinosa i — — — = | Hydrellia formosa 6 1 — — = yf Hydrellia griseola a 4 82 16 = 102 Hydrellia tibialis — — 47 a = 47 Lytogaster excavata 3 — 4 1 = 8 Nostima scutellaris 2 ] > — = 8 Notiphila adusta — 1 8 — = 9 Notiphila loewi — — 5 1 = 6 Notiphila pauroura — _ 2D, — = 22 Notiphila phaeopsis — 2 18 = 20 Paralimna punctipennis a 3 70 6 = 79 Parydra aquila — — 5) — = 5 Parydra breviceps == 3 10 a = 13 Parydra quadrituberculata --- — 3 — = 3 Philygria debilis | — — = = 1 Polytrichophora orbitalis — - 30 6 = 36 Ptilomyia enigma — _ 3 — = 3} Scatella stagnalis — — 145 Dy 147 Typsilopa atra 13 9 58 45 = 125 Zeros flavipes — — 14 2 = 16 Total = 71 26 613 23 = 949 were not adversely affected by low quan- tities of decaying organic matter. Although shore-fly primary consumers were collected from Miami Trace and Mallard wetlands, approximately 60 to 78% of the specimens collected were detritivores, respectively (Table 1). This investigation confirms that shore-fly primary consumers and detriti- vores rapidly colonize restored and con- structed wetlands in Ohio. Although Scheir- ing and Deonier (1979) proposed that re- source quantity increased the richness and abundance of well-adapted ephydrid spe- cies in transient habitats, the low detritus quantities, and abundance of detritivores in newly restored wetlands supports Batzer and Wissinger’s (1996) contention that de- tritivore colonization was not affected by the growth of macrophytes and algae, and insect abundance remained constant or in- creased when macrophytes were cut and de- bris removed. The early colonization of restored wet- land ecosystems by large numbers of A. simplex, D. obscurella, and Po. orbitalis suggests these species are important com- ponents in the early development of viable food chains that sustain diverse predaceous invertebrate and/or vertebrate communities. During late summer, increases in the pop- ulations of HAydrellia griseola, Hydrellia tibialis (1.e., leaf miners), 7. atra, and a di- atom feeder, Paralimna punctipennis are as- sociated with increased density and growth of macrophyte and diatom food resources. Typopsilopa spp. have been associated with the consumption of decaying tissue on dam- aged monocot stems (Keiper et al. 2001). The increase of micro- and macrophyte pro- duction represents a significant augmenta- VOLUME 106, NUMBER 2 467 Table 4. Shore-fly community of a new wetland at Miami Trace County Park. May June July Aug. Sept. Total Allotrichoma simplex 23 Te 102 80 — = PAG Discocerina obscurella = == > 24 3 = 32 Ditrichophora exigua — = = 2 an = 2 Hyadina binotata == —_ 3 = —_ = B Hydrellia griseola 4 20 34 4 — = 62 Hydrellia tibialis 9 11 33 — = 53 Hydrochasma leucoproctum — — — —_ 3 - 3 Notiphila adusta — l == I 1 = 3 Ochthera anatolikos —- 2 2 = = = 4 Paralimna punctipennis os ] 2) 3 5 = 11 Parydra quadrituberculata 4 — = at = = 4 Pelina truncatula — — l aii = = l Philygria debilis 33 ce 2 — = B35 Polytrichophora orbitalis l “= 3 4 7 15 Psilopa dupla — — 3 _ == = 3 Scatella favillacea 25 — = — = = DS Scatella obsoleta at — ] == = = 12 Scatella paludum l a 1 — = = 2 Scatella stagnalis a 2 5 l — = 8 Setacera atrovirens l — = == = | Typsilopa atra — | 5 — — = 6 Total = 103 108 180 152 19 562 tion of nutrient resources in newly restored Deonier 1985). Only a few individuals wetlands and a probable increase in food wed complexity. Early colonization, species accumulation (Fig. 1), and temporal abun- dance increases of shore-fly species (Tables 2—4) may be characteristic of initial stages of food chain development and the estab- lishment of complex food web interactions. Although Parydra aquila, Parydra brev- iceps, and Parydra quadrituberculata were collected in semi-aquatic grass, sand and mud shore habitats in Ohio (Table 5) (Stein- ly 1978), the rarity of these species in new- ly restored wetlands (Tables 1—4) suggests that obligate diatom species were not abun- dant. Species of Parydra are specialist on particular diatoms species (Bischof and within the genus Parydra were collected at Miami Trace and suggests that Parydra spe- cies are late colonizers and/or diatom spp. were not abundant. Hydrellia formosa and four Notiphila species colonized the Winton Woods sites, while one species of Notiphila was collect- ed at the Miami Trace wetland (Table 1). In all probability, the richness and abundance of N. adusta, N. loewi, N. pauroura, and N. phaeopsis is dependent on detritus accu- mulation on the root surfaces of different species of emergent vegetation (Larson and Foote 1997) at Heron Wetland. Although eight Notiphila species were collected dur- ing 1995 through 1998 from the older Mi- Table 5. Relative abundance of shore flies in Ohio aquatic habitats. Grass Shore Mud Shore Sand Shore Parydra aquila occ I r Parydra breviceps occ c occ Parydra quadrituberculata oce occ oce Abbreviations: c = common: occ = occasional; r = rare. 468 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 6. Sorenson Index of Similarity values for a newly created and two constructed wetlands in Ohio. Heron Wetland Winton Woods Mallard Wetland Winton Woods Mallard Wetland Winton Woods New Wetland Miami Trace 0.81% 0.51* = 0.65% * Comparison of community diversity values P < 0.001. ami Trace complex of restored wetlands (Steinly, unpub. data), only a few adults were collected from the restored wetland (Table 1) and the low richness and abun- dance of Notiphila spp. are attributed to the paucity of emergent vegetation in the re- stored wetland. The abundance of detritivores and leaf- miners, diatom consumers, 1.e., Pa. puncti- pennis, and Sc. stagnalis, a possible sec- ondary consumer of damaged monocots stems, 7. atra, and a cyanobacteria feeder suggests that at least five main nutrient sources were exploited by shore flies during the early stages of wetland restoration. Dur- ing the initial stages of wetland restoration, the abundance of detritivores, leafminers, a secondary consumer, an algivorous species, and consumers of cyanobacteria (e.g., Nos- tima and Hyadina species) suggest that the rudiments of viable food chains were pre- sent in late spring and early summer within the Mallard and Heron wetlands. Early col- lection, the increase of shore-fly richness and abundance (Tables 2—4) during the summer months, and the utilization of five different nutrient resources, i.e., algae, mac- rophytes, damaged monocot stems, detritus, and cyanobacteria suggest that shore-fly colonization provides a foundation for the development of diverse food chains and food webs. The successful restoration and maturation of wetland food chains is de- pendent on establishment and growth of aquatic micro- and macrophyte and microbe communities, accumulation of detritus, and concurrent colonization and establishment of resident primary consumers and detriti- vores, 1.e., shore flies that move nutrients into food webs. Once detritus, bacterial, and plant nutrients are assimilated by shore flies, wetland production is accessible to predaceous invertebrates (e.g., Ochthera anatolikos Clausen) and vertebrates. The abundance of insect consumer populations has been linked to waterfowl reproductive success that is dependent on the quantity and quality of insect protein (Driver et al. 1974). In some instances early emergence of the Chironomidae (Diptera) provides a large quantity of protein (McLaughlin and Harris 1990), while waterfowl utilize shore flies and other families of Diptera for food later in the season. Superficially, the significant difference in H’ values and the high Sorenson similarity value for the Mallard and Heron richness comparison are contra indicators. Although a Sorenson index of 0.81 and monthly spe- cies accumulation (Fig. 1) suggest that shore-fly community richness values in Mallard and Heron wetlands are compara- ble, the significant difference in H’ values (P < .0001, t = —22.77, df = 2540) sug- gests that these wetlands harbored unique species assemblages. The difference in shore-fly assemblages is attributed to ex- treme variation of individual species abun- dance within each wetland (Table 1). Al- though, the relatively high index of simi- larity (Table 6) suggests that biological and physical conditions were comparable in Mallard and Heron wetlands, the variation in shore-fly species abundance and the dif- ference in the abundance of detritivores and primary consumers suggest that ecosystem development and/or maturation within each wetland was asynchronous. Comparison of Miami Trace to Mallard and Heron Wetlands yielded low Sorenson indices (Table 6), differences in species ac- cumulation patterns, and H’ values that VOLUME 106, NUMBER 2 35 —i—— Miami Trace Wetland Mallard Wetland 30 -=-«-=- Heron Wetland Shore-fly Species Accumulation June July lense dle were significantly different (P < 0.001, t = 9.38, df = 922 and t = —5.68, df = 863, respectively). These differences suggest that the Miami Trace species assemblage was unique. The difference in shore-fly communities is attributed to local precipi- tation frequency and quantity, and disparate physical and biological conditions that are unique to restored and/or constructed wet- lands. Further, the difference in shore-fly communities at Miami Trace as compared to the Winton Woods wetlands is confirmed by early colonization of shore flies in May vs. June and July and early development of species accumulation asymptote (Fig. 1) in the restored wetlands. The importance of insect primary con- sumers and detritivores within food chains has been acknowledged (Batzer and Wis- singer 1996, Hansen and Castelle 2000), but a comprehensive understanding of in- sect herbivore, detritivore, and predator 469 August September October Month Shore-fly species accumulation in newly constructed and restored wetlands. communities in restored wetlands is want- ing. The scarcity of information is attribut- ed to the daunting task of identifying large numbers of invertebrate species, and spe- cies interactions within and between food webs, and the lack of sampling in shoreline habitats (Keiper et al. 2002). Additionally, the number of food chain interactions and food web dynamics may vary from one geographic region to another. The Ephydridae are a trophically diverse family that provides a unique opportunity to study the movement of wetland produc- tion into food webs because shore-fly spe- cies are routinely collected and abundant in wetlands. Without the colonization of wet- lands by shore flies and/or other insect fam- ilies that contain primary consumers and detritivores, the movement of primary pro- duction and detritus into food chains and development of food webs may be limited. Design and management of wetlands that 470 promotes the colonization and growth of in- sect communities to expedite the cycling of plant resources (1.e., primary production) and energy transfer will provide a founda- tion for diverse food chains and complex food webs that sustain an abundance and variety of invertebrate and vertebrate spe- cies. ACKNOWLEDGMENTS We thank the Cinergy Corporation and Hamilton County Park District for their support that made this project possible. The cooperation of John Kline and Robert Ma- son was invaluable. Nancy Solomon pro- vided statistical advice and counsel. The as- sistance and dedication of Kevin Hill, Chris Yeager, Brit Farrar, Jeff Eakin, and Susan Ytsma in the preparation of specimens is greatly appreciated. LITERATURE CITED Bataille, K. J. and G. A. Baldassarre. 1993. Distribu- tion and abundance of aquatic marcro-inverte- brates following drought in three prairie pothole wetlands. Wetlands 13: 260—269. Batzer, D. P. and S. A. Wissinger. 1996. Ecology of insect Communities in nontidal wetlands. Annual Review of Entomology 41: 75—100. Bischof, M. R. and D. L. Deonier. 1985. Life history and immature stages of Parydra breviceps (Dip- tera: Ephydridae). Proceedings of the Entomolog- ical Society of Washington 87: 805-820. Brown, S. C., K. Smith, and D. Batzer. 1997. Macro- invertebrate response to wetland restoration in northern New York. Environmental Entomology 26: 1016-1024. Cheal, F, J. A. Davis, J. E. Growns, J. S. Bradley, and F H. Whittles. 1993. 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H. Prince. 1981. Dabbling duck and aquatic macroinvertebrate responses to manipulated wetland habitat. Journal of Wildlife Management 45: 1-15. Keiper, J. B. and W. E. Walton. 2002. Effects of three vegetation management strategies on shore-flies (Diptera: Ephydridae) in newly constructed treat- ment wetlands. Annals of the Entomological So- ciety of America 95: 570-576. Keiper, J. B., J. Jiannino, M. Sanford, and W. E. Wal- ton. 2001. Biology and immature stages of Ty- popsilopa nigra (Wirth) (Diptera: Ephydridae), a secondary consumer of damaged stems of wetland monocots. Proceedings of the Entomological So- ciety of Washington 103: 89-97. Keiper, J. B., W. E. Walton, and B. A. Foote. 2002. Biology and ecology of higher Diptera from fresh- water wetlands. Annual Review of Entomology 47: 207-232. King, R. S. and J. C. Brazner. 1999. Coastal wetland insect communities along a trophic gradient in Green Bay, Lake Michigan. Wetlands 19: 426— 437. Krapu, G, L. and G. A. Swanson. 1975. Some nutri- tional aspects of reproduction in prairie nesting pintails. Journal of Wildlife Management 39: 156— 162. Larson, L. and B. A. Foote. 1997. Biology of four species of Notiphila Fallén (Diptera: Ephydridae) associated with the yellow water lily, Nuphar leu- VOLUME 106, NUMBER 2 tum (Nymphaeaceae). Proceedings of the Ento- mological Society of Washington 99: 541-559. LaSalle, M. W. and L. P. Rozas. 1991. Comparing the benthic macrofaunal assemblages of creekbank beds of the spikerush Eleocharis parvula (R&S) Link and adjacent unvegetated areas in a Missis- sippi brackish marsh. Wetlands 11: 229-244. Layton, R. J. and J. R. Voshell. 1991. Colonization of new experimental ponds by benthic macroinver- tebrates. Environmental Entomology 20: 110— LG Marshall, S. A. 1994. Peatland Sphaeroceridae (Dip- tera) of Canada. Memoires of the Entomological Society of Canada 169: 173-179. Martin, A. C. and E M. Uhler. 1939. Food of game ducks in United States and Canada. USDA Tech- nical Bulletin 634. 156 pp., 153 pl. McLaughlin, D. B. and H. J. Harris. 1990. Aquatic insect emergence in two Great Lakes marshes. Wetland Ecology and Management 1: 111-121. Murkin, H. R., PB. G. Abbott, and J. A. Kadlec. 1983. A comparison of activity traps and sweep nets for sampling nektonic invertebrates in wetlands. Ca- nadian Journal of Fisheries and Aquatic Sciences 48: 2355-2364. Murkin, H. R. and B. D. J. Batt. 1987. The interactions of vertebrates and invertebrates in peatlands and marshes. Memoires of the Entomological Society of Canada 140: 15-30. Olive, J. H. and C. A. Dambach. 1973. Benthic macro- invertebrates as indexes of water quality in Whet- stone Creek, Morrow County, Ohio (Sciota River Basin). Ohio Journal of Science 73: 129-149. Orians, G. H. 1980. Some adaptations of marsh nesting birds, Caribou Parklands, British Columbia. Monographs in Population Biology, Vol. 14. Princeton University Press, Princeton, New Jer- sey. Regensburg, J. T. 1976. The shore flies of southern Ohio (Diptera: Ephydridae). M.S. Thesis, Miami University. Oxford, Ohio. 192 pp. Rosenberg, D. M. and H. V. Danks. 1987. Aquatic in- sects of peatlands and marshes in Canada: Intro- duction. Memoirs of the Entomological Society of Canada 140: 1-4. Scheiring, J. T. 1974. Diversity of shore flies (Diptera: Ephydridae) in inland freshwater habitats. Journal of the Kansas Entomological Society 47: 485— 491. Scheiring, J. T. and D. L. Deonier. 1979. Spatial pat- terns in shore fly community structure, pp. 73-80. In Deonier, D. L., ed. First Symposium on the 471 Systematics and Ecology of Ephydridae (Diptera). North American Benthological Society. Scheiring, J. T. and B. A. Foote. 1973. Habitat distri- bution of the shore flies of northeastern Ohio. Ohio Journal of Science 73: 152—166. Sharitz, R. R. and D. P. Batzer. 1999. An introduction to freshwater wetlands in North America and their invertebrates, pp. 1-22. In Batzer, D. P, R. B. Rader, and S. A. Wissinger, eds. Invertebrates in Freshwater Wetlands of North America: Ecology and Management. John Wiley & Sons, Inc., New York, New York, 1100 pp. Steinly, B. A. 1978. The shore flies of northern Ohio (Diptera: Ephydridae). M.S. Thesis, Miami Uni- versity. Oxford, Ohio. 249 pp. . 1984. Shore fly (Diptera: Ephydridae) com- munity structure in a xeric grass habitat. Proceed- ings of the Entomological Society of Washington 84: 749-759. . 1986. Violent wave action and the exclusion of Ephydridae (Diptera) from marine temperate intertidal and freshwater beach habitats. Proceed- ings of the Entomological Society of Washington 88: 427-437. . 1990. Shore-fly (Diptera: Ephydridae) com- munity structure in selected terrestrial grass hab- itats of Ohio and Illinois. Proceedings of the En- tomological Society of Washington 92: 188-193. . 2001. Shore-fly (Diptera: Ephydridae) colo- nization of saline habitats in Ohio. Entomological News 113: 121-124. Steinly, B. A. and D. L. Deonier. 1980. New records of Ohio shore flies (Dipera: Ephydridae). Ohio Journal of Science 80: 41—42. Streever, W. J., K. M. Portier, and T. L. Crisman. 1996. A comparison of dipterans from ten created and ten natural wetlands. Wetlands 16: 416—428. Thier, R. W. and B. A. Foote. 1980. Biology of mud- shore Ephydridae (Diptera). Proceedings of the Entomological Society of Washington 82: 517— 535: Todd, J. L. and B. A. Foote. 1987. Spatial and temporal distribution of shore flies in a freshwater marsh (Diptera: Ephydridae). Proceedings of the Ento- mological Society of Washington 89: 448—457. Voigts, D. K. 1976. Aquatic invertebrate abundance in relation to changing marsh vegetation. American Midland Naturalist 95: 313-322. Wilson, E. O. 1987. The little things that run the world (the importance and conservation of inverte- brates). Conservation Biology |: 344—346. Zahl, P. 1967. Life in a ‘““‘Dead’’ Sea—Great Salt Lake. National Geographic Magazine 132: 252—263. Zar, J. 1984. Biostatistical Analysis. Prentice Hall, En- glewood, New Jersey, 718 pp. PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 472-474 NOTE First Report of Ixodes kopsteini Oudemans (Acari: [xodida: Ixodidae) from the Kingdom of Cambodia, with a Summary of Known Hosts of this Tick in Continental Southeastern Asia The uniquely ovoviviparous tick [xodes kopsteinit Oudemans is a widespread para- site of emballonurid, molossid, pteropodid and vespertilionid bats in the Oriental Zoo- geographic Region and, to a lesser extent, the Afrotropical and Australian Regions (Hoogstraal and Aeschlimann 1982, Petney and Keirans 1994). In the archipelagic states of southeastern Asia, collections of this tick have been made as far southward as Bogor in western Java, Indonesia (An- astos et al. 1973), and as far eastward as Luzon in the Philippines (Kohls and Clif- ford 1968) and Ambon and Seram in In- donesia’s Moluccan Islands (Maluku) (Kohls and Clifford 1961, Durden et al. 1990). However, outside peninsular Malay- sia (Kohls and Clifford 1968) and Thailand (Gould et al. 1970, Tanskul et al. 1983), no collections of /. kopsteini have heretofore been reported from continental southeastern Asia, perhaps because of the human turmoil that prevailed there throughout the second half of the twentieth century. During the summer of 2001, teams from U.S. Naval Medical Research Unit No. 2 (NAMRU-2), Jakarta, Indonesia, and Cambodia’s Nation- al Institute of Public Health (NIPH), Phnom Penh, conducted surveys for natural reser- voirs of Nipah virus and lyssaviruses (ra- bies and close relatives) in Cambodia. On 11 July, while sampling a large population of the wrinkle-lipped bat, Chaerephon pli- cata (Buchanan), which has long inhabited the enormous attic space of the Royal Cam- bodian National Museum (Fig. 1), one of us (MJB) collected an adult male C. plicata (total body length 35 mm, mass 22 g) that harbored a single female /xodes just above the right ear canal (not on the tragus or an- titragus). The tick’s location presumably shielded it from host grooming. Subsequent examination revealed this specimen to be the first Cambodian example of J. kopsteini, which has been deposited (as RML 123173) in the U.S. National Tick Collection, Insti- tute of Arthropodology and Parasitology, Georgia Southern University, Statesboro. The bat was accessioned (as voucher no. 6218) at the NAMRU-2/NIPH Laboratory, Phnom Penh. The Cambodian National Mu- seum attic, which also harbored colonies of Taphozous melanopogon Temminck and T. theobaldi Dobson (the latter was not en- countered during the 2001 surveys), is re- puted to be among the largest single aggre- gations of bats found in an artificial struc- ture anywhere in the world (C. E. Rup- precht, personal communication). With one exception (Gould et al. 1970), all known collections of /. kopsteini from mainland Southeast Asia (Table 1) have been made from members of the so-called Mastiff bat family Molossidae, which is only modestly represented in tropical Asia (of 80 molossid species accepted by Wilson and Reeder (1993), just 8 occur in the Ori- ental Zoogeographic Region). However, some Southeast Asian molossids are widely distributed. Chaerephon plicata, in partic- ular, has a vast range, extending from India and Sri Lanka eastward through southern China to Vietnam, Hainan Island and the Philippines, then southward through the Malay Peninsula, Sumatra, Borneo, Java, the Lesser Sunda Islands, and, southwest of Indonesia, the Cocos (Keeling) Islands in the Indian Ocean (Wilson and Reeder 1993, Nowak 1999). The range of this bat alone all but guarantees that /. kopsteini will eventually be found in the four Southeast Asian countries that have yet to yield col- VOLUME 106, NUMBER 2 ioe de lections: Laos, Myanmar (Burma), Singa- pore, and Vietnam. For technical assistance during this study, we warmly thank the staff of the National Institute of Public Health, Government of Cambodia. Thanks also to Charles E. Rup- precht, Rabies Section, Viral and Rickettsial Zoonoses Branch, Division of Viral and Rickettsial Diseases, National Center for In- Royal Cambodian National Museum, Phnom Penh, Cambodia; photo by M.J. Bangs, July 2001. fectious Diseases, Centers for Disease Con- trol and Prevention, Atlanta, Georgia, and to Karen N. Wolf, formerly Virginia-Mary- land Regional College of Veterinary Med- icine, Blacksburg, Virginia. Portions of this research were funded by National Institute of Allergy and Infectious Diseases grant Al 40729 to JEK. The opinions and assertions advanced herein are those of the authors Table 1. Bat species known to be hosts of /xodes kopsteini in continental Southeast Asia. Bat Species Country and State/Province Reference/Collection No. Emballonuridae: Taphozous theobaldi Dobson Molossidae: Chaerephon johorensis (Dobson) Chaerephon plicata (Buchanan) Cheiromeles torquatus Horsfield Mops mops (de Blainville) Thailand: Saraburi Malaysia: Selangor Cambodia: Phnom Penh Malaysia: Melaka Thailand: Kanchanaburi and Saraburi Malaysia: Selangor Malaysia: Selangor Gould et al. 1970 Kohls and Clifford 1968 RML 123173 RML 112146 Tanskul et al. 1983 Kohls and Clifford 1961 Kohls and Clifford 1968 a 474 and are not to be construed as official or reflecting the views of the U.S. Depart- ments of the Army, Navy, or Defense. LITERATURE CITED Anastos, G., T. S. Kaufman, and S. Kadarsan. 1973. An unusual reproductive process in /xodes kop- steini (Acarina: Ixodidae). Annals of the Ento- mological Society of America 66: 483—484. Durden, L. A., B. V. Peterson, N. Wilson, and B. Christiansen. 1990. Some ectoparasites of bats from Seram Island, Indonesia. Entomological News 101: 48-56. Gould, D. J., C. M. Clifford, P. Lakshana, K. C. Em- erson, H. Hoogstraal, J. T. Marshall, H. E. Stark, M. Nadchatram, E. W. Davis, I. Inlao, N. Manee- chai, N. Klaimanee, and P. Punthusiri. 1970. Dis- tribution and ecology of ectoparasites of verte- brates in Southeast Asia, pp. 97-105. /n Annual Progress Report of the SEATO Medical Research Laboratory, April 1969—March 1970. United States Army Medical Component, Bangkok, Thai- land. Hoogstraal, H. and A. Aeschlimann. 1982. Tick-host specificity. Mitteilungen der Schweizerischen En- tomologischen Gesellschaft/Bulletin de la Société Entomologique Suisse 55: 5—32. Kohls, G. M. and C. M. Clifford. 1961. A new species of Ixodes (Lepidixodes) from bats in Malaya, North Borneo, and the Congo (Acarina: Ixodidae). Acarologia 3: 285-290. . 1968. lvodes (Lepidixodes) kopsteini Oude- mans, 1925, valid name for /. (L.) paradoxus PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Kohls and Clifford, 1961, with notes on distribu- tion and hosts (Acarina: Ixodidae). Journal of Medical Entomology 5: 503-506. Nowak, R. M. 1999. Walker’s Mammals of the World, 6th ed., Vol. I. Johns Hopkins University Press, Baltimore. Ixx + 836 pp. Petney, T. N. and J. E. Keirans. 1994. Ticks of the genus /xodes in South-east Asia. Tropical Bio- medicine 11: 123-134. Tanskul, P. (L.), H. E. Stark, and I. Inlao. 1983. A checklist of ticks of Thailand (Acari: Metastig- mata: Ixodoidea). Journal of Medical Entomology 20: 330-341. Wilson, D. E. and D. M. Reeder. 1993. Mammal Spe- cies of the World: A Taxonomic and Geographic Reference, 2nd ed. Smithsonian Institution Press, Washington, DC. xviii + 1206 pp. Richard G. Robbins, Armed Forces Pest Management Board, Walter Reed Army Med- ical Center, Washington, DC 20307-5001, U.S.A. (e-mail: richard.robbins @ osd.mil), Michael J. Bangs, American Embassy Jakar- ta, U.S. Naval Medical Research Unit No. 2, Department of Entomology, Parasitic Diseas- es Program, FPO AP 96520-8132 (e-mail: bangsmj @namru2.med.navy.mil), and James E. Keirans, Institute of Arthropodology and Parasitology, Georgia Southern University, P.O. Box 8056, Statesboro, GA 30460-8056, U.S.A. (e-mail: jkeirans @ gasou.edu) PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 475-477 NOTE Mealybug Species (Hemiptera: Pseudococcidae) Found on Ornamental Crops in California Nursery Production Mealybugs (Hemiptera: Pseudococcidae) are troublesome pests of ornamental crops throughout temperate regions of the world, including California. However, it is not cer- tain which mealybug species are most com- mon in California greenhouses and nurser- ies, and it is not known if recent, invasive species such as the pink hibiscus mealybug, Maconellicoccus hirsutus (Green), are pre- sent. Surveys and collections were made be- tween November 16, 2001 and March 7, 2003 to nurseries that reported mealybugs. Additionally, mealybugs were also sent di- rectly to the author by growers and advisors in University of California Cooperative Ex- tension. Mealybugs were collected or re- ceived from a total of 19 nurseries in five counties. The mealybugs were slide-mounted ac- cording to methods adopted from Kozar- zhevskaya (1968) and Williams and Gran- ara de Willink (1992). Slide-mounted mealybugs were identified using keys by McKenzie (1967) and an unpublished man- uscript by R. J. Gill. The mealybug species most frequently found in this survey of California green- houses were the longtailed mealybug, Pseu- dococcus longispinus (Targioni Tozzetti), and the citrus mealybug, Planococcus citri (Risso) (Table 1). With the exception of ro- ses, P. citri fed on many short-term crops such as coleus, whereas P. longispinus of- ten fed upon perennial crops such as cycad and Phormium tenax. The lilybulb mealy- bug, Vryburgia amaryllidis (Bouché), and the obscure mealybug, Pseudococcus vibur- ni (Signoret), were commonly found as well. Vryburgia amaryllidis is limited to a few plant families (especially Liliaceae and Iridaceae). It occurs on the bulb and on the basal portion of the leaves. Pseudococcus viburni was found both on the roots and the aerial portion of the plants, most commonly on short-term crops. Seven other mealybug species were found to occur less frequently. Many of these mealybugs were specialists found only on one species of plant. For example, the phormium mealybug, Balanococcus diminutus (Leonardi), is found only on Phormium tenax, and the noxious bamboo mealybug, Antonina pretiosa Ferris, is found only on bamboo species (Ben-Dov 1994). This survey allowed us to obtain a record of the mealybug species present in green- houses or nurseries in predominant growing areas of California. As growers are increas- ingly interested in incorporating natural en- emies into their insect control programs, in- formation on species composition is helpful for directing future biological control pro- jects. Thanks to K. Robb, S. Tjosvold and J. Newman (U.C. Cooperative Extension) for assisting with mealybug collections. We would also like to thank the University of California at Davis (arboretum and conser- vatory), Humbolt State University, Dramm and Echter Inc., Robert Hall Nursery, Gold- en State Bulb Co., Anderson Seed Co., Weidners’ Gardens Inc., Aedes and Gish Nurseries, Browns’ Plant Inc., Sea Coast Greenhouses, Suncrest Nurseries Inc., Ca- vanaugh Color, Kitayama Bros. Inc., Boe- thing Treeland Farms Inc., Sunshine Floral, Keeline Wilcox Nursery, McLellan Botan- icals, and Pajaro Valley Greenhouses Inc. for allowing the collection of mealybug specimens at their nurseries. Thank you to Ray Gill for allowing the use the manu- script; The Scale Insects of California: Part 476 Table 1. Records of mealybugs found during the California nursery survey. i Mealybug Species um Antonina pretosa Ferris Balanococcus diminutus (Leonardi) Phenacoccus defectus Ferris Phenacoccus solani Ferris Planococcus citri (Risso) Pseudococcus calceolariae (Maskell) Pseudococcus longispinus (Targioni Tozzetti) Pseudococcus viburni (Signoret) Rhizoecus falcifer Kunckel d’Herculais Spilococcus mamillariae (Bouché) Vryburgia amaryllidis Bouché Vryburgia trionymoides (De Lotto) i Counties Santa Cruz Santa Cruz Yolo Yolo San Diego Santa Cruz Ventura Yolo Yolo Humbolt San Diego Santa Cruz Yolo San Diego Santa Cruz Ventura Yolo San Diego Santa Cruz Yolo Yolo Number of Nurseries 1 10 Host Plants Bambusa sp. Phormium tenax Graptoveria var. ‘spirit of 76° Verbena tenuisecta ‘alba’ Oxalis corniculata Rosa sp. Rosa sp. Rosa sp. Rosa sp. Rosa sp. mandevilla (=Dipladenia sp.) Coleus blumei Peperomia sp. Aphelandra squarrosa Aeschynanthus radicans Cyrtomium sp. Calathea sp. Stehanotis sp. Dendranthema grandiflora Philodendron sp. Limonium sp. Nerium oleander Achillea ‘moonwalker’ Rosa sp. cycad cycad Phormium tenax Phormium tenax orchid orchid Lotus var. “Amazon sunset” Dracaena sp. Schefflera arboricola Abutilon hybridum Heuchera sanguinea Philodendron *‘Anderson’s Red’ Fuchsia sp. Howea forsterana Gymnocalycium stenopleurum Agapanthus africanus Eucomis sp. Cryptanthus purpureus Cryptanthus purpureus Hippeastrum sp. Nerine sp. Echeveria chihuahuaensis PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Collection Dates 7/31/02 7/31/02 8/5/02 9/16/02 7/30/02 11/16/01 7/16/02 7/31/02 7/17/02 2/1/03 7/16/02 7/16/02 7/16/02 7/16/02 7/16/02 7/16/02 7/16/02 7/16/02 7/16/02 7/16/02 7/31/02 7/17/02 4/29/02 11/16/01 3/26/02 2/5/03 7/16/02 7/17/02 7/31/02 2/1/03 7/31/02 11/16/01 11/16/01 6/12/02 4/29/02 7/17/02 7/31/02 10/15/02 3/7/03 6/27/02 7/7102 7/17/02 7/31/02 7/31/02 7/31/02 6/12/02 VOLUME 106, NUMBER 2 4. The Mealybugs (Coccoidea: Pseudococ- cidae). Funding was provided by The Cal- ifornia Association of Nurserymen and the USDA Nursery and Floriculture Research Initiative. LITERATURE CITED Ben-Dovy, Y. 1994. A Systematic Catalogue of Mealy- bugs of the World. Intercept Limited, Andover U.K., 686 pp. Kozarzhevskaya, E. F 1968. Techniques for preparing slides for coccid (Homoptera: Coccoidea) deter- mination. Entomologicheskoye Obozreniye 47: 477 248-253 (English translation, Entomological Re- view 47: 146-149). McKenzie, H. L. 1967. Mealybugs of California. Uni- versity of California Press, Los Angeles, 526 pp. Williams, D. J. and M. C. Granara de Willink. 1992. Mealybugs of Central and South America. C.A.B. International, Wallingford, 635 pp. H. M. Laflin, P. J. Gullan and M. P. Par- rella, Department of Entomology, One Shields Avenue, University of California, Davis, CA 95616, U.S.A. (e-mail: hlaflin@ hotmail.com; pjgullan@ ucdavis.edu; mpparrella@ ucdavis.edu) PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 478-480 NOTE Larval Development of Cylindrotoma distinctissima americana Osten Sacken, 1865 (Tipulidae: Cylindrotominae) on Maianthemum dilatatum (Liliaceae) in California The genus Cylindrotoma Macquart is unique among North American crane flies because its members feed on the developing leaves of higher plants. Other members of the nearly worldwide subfamily Cylindro- tominae feed on moss as well as higher plants (Alexander and Byers 1981). One of us (GP) had the opportunity to collect the larvae of Cylindrotoma distinc- tissima americana Osten Sacken on leaves of the false-lily-of-the-valley (or mayber- ry), Maianthemum dilatatum (Wood) Nels. & Macbr. (Liliaceae) in northern California. This report discusses this find and briefly describes the larvae and pupae. This is the first record of this crane fly species, and of the subfamily Cylindrotominae, in Califor- nia. The previously known distribution of Cylindrotoma distinctissima americana Was from British Columbia to Oregon (Alex- ander 1967, Oosterbroek 2002). This is also the first record of this crane fly species feeding on Maianthemum dilatatum. Pre- viously recorded host plants in North America are Allium, Anemone, Trautvetter- ia, Stellaria, and Viola (Brodo 1967). The host plant, M. dilatatum, is native to western North America from Alaska to Cal- ifornia (Hitchcock and Cronquist 1973). It is especially abundant along the Pacific Northwest coast, where it normally grows in the shade of large trees, but can be found completely exposed, even almost adjacent to the high tide level. It is a perennial nor- mally not reaching over 40 cm tall, that dies back in the fall and grows anew in the spring from slender rhizomes (Hitchcock and Cronquist 1973). Populations of M. dilatatum harboring Cylindrotoma larvae were discovered in a moist forest of Sitka spruce (Picea sitch- ensis (Bong.) Carr. (Pinaceae) near Lake Earl in Humboldt County, northwestern California, on April 13, 2001. The larval and pupal stages (n = 8) were observed and photographed at the site. The mature larvae were unlike any other leaf- feeding insects found on this plant. They were green and relatively smooth except for a pair of small protuberances on the dorsum of each abdominal segment except the last (Fig. 1). The larvae fed on the dorsal and ventral sides of the leaves, leaving large holes very similar to damage caused by sawfly larvae feeding on the same plant species in Oregon. When mature (length = 12 mm), the larvae pupated on the upper surface of the leaves in a very peculiar manner. The last molt was incompletely shed, with the posterior portion of the pupa remaining within the anterior part of the last larval skin (Fig. 2). The skin served as a holdfast to the leaf surface and could not be removed without tearing it apart. This was the only portion of the pupa attached to the leaf surface. The pupae were smooth, green, 13 mm in length, with a pair of tho- racic respiratory horns as the only body projections (Fig. 2). Both the larvae and pu- pae were well camouflaged on the leaves. The adults (length 13 mm) emerged within two weeks after pupation (Fig. 3). Observations reported here are similar to those reported by Cameron (1918), who studied the same species of Cylindrotomi- nae (under the name of C. splendens Doane, 1900) developing on the leaves of Traut- vetteria caroliniensis (Walt.) Vail (= T. grandis Nutt.) (Ranunculaceae) in British Columbia. It is interesting that this species can develop on both monocots and dicots. In both cases (California and British Co- lumbia) the host plants were understory herbs in damp, well shaded, forested areas. VOLUME 106, NUMBER 2 479 Figs. 1-3. Cylindrotoma distinctissima americana. 1, Mature larva (L = 12 mm) feeding on a leaf of Maianthemum dilatatum. 2, Pupa (L = 13 mm) attached to a leaf of Maianthemum dilatatum by the dried last larval skin. 3, Adult that emerged from the pupa in Fig. 2. 480 Cameron (1918) reported mature larvae measuring 17 mm, which is larger than the California specimens. However the sizes of the pupae (11.7—13.3 mm) in British Co- lumbia were similar to those in California (13 mm). According to Cameron (1918), the larvae overwinter and resume develop- ment the following spring. This could be quite possible with the California popula- tions, although this aspect of the life cycle was not investigated. LITERATURE CITED Alexander, C. P. 1967. The crane flies of California. Bulletin of the California Insect Survey 8: 1—269. Alexander, C. P. and G. W. Byers. 1981. Tipulidae, pp. 153-190. In Manual of Nearctic Diptera, Vol. 1, Monograph No. 27, Research Branch, Agriculture Canada, Ottawa. Brodo, E 1967. A review of the subfamily Cylindro- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tominae in North America (Diptera: Tipulidae). University of Kansas Science Bulletin 47: 71-115. Cameron, A. E. 1918. Life-history of the leaf-eating crane fly Cylindrotoma splendens, Doane. Annals of the Entomological Society of America 11: 67— 89. Hitchcock, C. L. and A. Cronquist. 1973. Flora of the Pacific Northwest. University of Washington Press, Seattle, 730 pp. Oosterbroek, P. 2002. Catalog of the Crane Flies of the World (Tipuloidea: Families Pediciidae, Limoni- idae, Cylindrotomidae, and Tipulidae). Digital cat- alog, version October 2002, available from author. George Poinar, Jr., Department of Zoology, Oregon State University, Corvallis, OR 97331, U.S.A. (e-mail: poinarg@casco.net), and Jon Gelhaus, Department of Entomology, The Academy of Natural Sciences, Philadel- phia, PA 19103-1195, U.S.A. (e-mail: gelhaus @discovery.acnatsci.org) PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 481—482 NOTE Beetles (Coleoptera) Associated with the External Debris of Atta mexicana (FE. Smith) (Hymenoptera: Formicidae) from Ojuelos, Jalisco, México Beetles are closely associated with sev- eral species of social insects, especially with ants of the genera Afta Fabricius, 1804, and Acromyrmex Mayr, 1865 (Navar- rete-Heredia 2001 recorded 411 beetle spe- cies of 25 families). In México, there are a few faunistic studies on Aftta-beetle associ- ations. These were done in localities from Estado de México, Morelos, and Querétaro (Hinton and Ancona 1934, 1935; Deloya 1988; Rojas 1989; Marquez-Luna 1994), In this paper, we provide additional in- formation on beetles associated with the ex- ternal debris of Atta mexicana (FE Smith, 1858) in a “matorral xerofilo”’ plant asso- ciation (““chaparral” sensu Leopold 1950), from Presa el Cuarenta, Ojuelos, Jalisco. Specimens were collected from debris of a single ant mound and are deposited in the Entomological Collection of the Centro de Estudios en Zoologia with the following la- bel data: MEXICO: JAL, Ojuelos, Presa El Cuarenta, matorral xer6ofilo, 27.VII.2003, J. Cortés-Aguilar y C. I. Hermosillo cols., ex detrito de Atta mexicana. In this locality, the following species were collected: Morion lafertei Guérin- Meéneville, 1844, and one unidentified ca- rabid (Carabidae); Oosternum attacomis Spangler, 1962 (Hydrophilidae); Epiglyptus costatus (LeConte, 1852), Pseudister rufu- lus (Lewis, 1888), Xestipyge multistriatum Lewis, 1888, Hister sp., and two unidenti- fied histerid species (Histeridae); one un- identified Aleocharinae; Philonthus alius Bernhauer and Schubert, 1914, Platydracus fulvomaculatus (Normann, 1837) (Staphy- linidae); Ptichopus angulatus (Percheron, 1835) (Passalidae); Onthophagus rufescens Bates, 1887, Ataenius holopubescens Hin- ton, 1938, Diapterna dugesi (Bates, 1887), Euparixia mexicana Gordon and McCleve, 2003 (Fig. 1) (Scarabaeidae); one uniden- tified scydmenid (Scydmaenidae); Lapethus sp. (Cerylonidae); one unidentified anthicid (Anthicidae); and Bycrea villosa Pascoe, 1868 (Tenebrionidae). Euparixia mexicana was recently de- scribed as a cryptic species similar to E. duncani Brown. Actually some specimens were confused with E. duncani by Wood- ruff and Cartwright (1967). It is now known from the Mexican states of Sonora, Sinaloa, and Jalisco. This species is recognized eas- ily by the shape of the pronotum that is con- Fig. 1. Dorsal view of Euparixia mexicana. 482 stricted only in the posterior one-third, with the posterior angles prominent and acute, and by the pronotal disc with shallow punc- tures separated by a flat pronotal surface (Fig. 1). Label data lacks specific host re- cords, but A. mexicana was presumed to be the host. In this note, we record for the first time an accurate host record for this spe- CIES: One question remains on the identity of the specimens cited as E. duncani by Wood- ruff and Cartwright (1967) from México. Gordon and McCleve (2003) stated that E. duncani is not known from México, so do the specimens cited from Durango and Puebla belong to E. mexicana or to a new cryptic species? Acknowledgments.—I thank my family (JLNH): G. A. Quiroz-Rocha, J. Quiroz Ya- mada, L. A. Navarrete-Quiroz, and E. M. Navarrete-Quiroz (her first fieldwork at the age of 18 months), and to our friends C. I. Hermosillo Pena, Ma. F Nunez-Yépez, and M. Vasquez-Bolanos for their company during the fieldwork. LITERATURE CITED Deloya, C. 1988. Coledpteros lamelicornios asociados a depositos de detritos de Atta mexicana (Smith) (Hymenoptera: Formicidae) en el sur del estado de Morelos, México. Folia Entomologica Mexi- cana (75): 77-91. Gordon, R. D. and S. McCleve. 2003. Five new spe- cies of Euparixia Brown (Coleoptera: Aphodiidae: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Eupariinae), with a revised key to species. Pro- ceedings of the Entomological Society of Wash- ington 105(3): 685—697. Hinton, H. E. and L. Ancona H. 1934. Fauna de co- le6pteros en nidos de hormigas (Atta), en México y Centro América. Anales del Instituto de Biolo- gia 5(3): 243-248. . 1935. Fauna de cole6pteros en nidos de hor- migas (Atta), en México y Centro América. II. Anales del Instituto de Biologia 6(3—4): 307-316. Leopold, A. S. 1950. Vegetation zones of Mexico. Ecology 31: 507-518. Marquez-Luna, J. 1994. Coleopterofauna asociada a detritos de Atta mexicana (FE Smith) (Hymenop- tera: Formicidae) en dos localidades del norte de Morelos, México. Tesis Profesional, Facultad de Ciencias, UNAM, México. Navarrete-Heredia, J. L. 2001. Beetles associated with Atta and Acromyrmex ants (Hymenoptera: For- micidae: Attini). Transactions of the American Entomological Society 127(3): 381—429. Rojas, P. 1989. Entomofauna asociada a los detritos de Atta mexicana (FE Smith) (Hymenoptera: Formi- cidae) en una zona darida del centro de México. Acta Zoolégica Mexicana (nueva serie) (33): 1— Sill. Woodruff, R. E. and O. L. Cartwright. 1967. A review of the genus Euparixia with description of a new species from nest of leaf-cutting ants in Louisiana (Coleoptera: Scarabaeidae). Proceedings of the United States National Museum 123(3616): 1—21. José Luis Navarrete-Heredia and Jestis Cor- tés-Aguilar, Centro de Estudios en Zoolo- gta, CUCBA, Universidad de Guadalajara, Apdo. Postal 234, 45100, Zapopan, Jalisco, México (e-mail: snavarre@cuchba.udg.mx, jcortesaguilar@ hotmail.com) PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 483-484 NOTE Homonymy in the Coccinelidae (Coleoptera), or Something Fishy About Pseudoscymnus Chapin Edward Chapin (1962) discovered some anomalous characters in certain species of Scymnus from Japan while preparing a re- port on the lady beetles (Coccinellidae) of Micronesia. The species in question possess true trimerous tarsi and reduced antennae (only nine antennomeres) terminating in ex- traordinarily long setae. Chapin proposed the new genus Pseudoscymnus for these un- usual lady beetles, based on Scymnus ha- reja Weise (type species) and five other species—S. kurohime Miyatake, S. seboshii Ohta, S. sylvaticus Lewis, S. pilicrepus Lewis and S. quinquepunctatus Weise—the last four included tentatively. In the decades that followed Chapin’s contribution, numerous additional species have been added to the genus, including the delightfully sibilant Pseudoscymnus tsugae Sasaji and McClure 1997, a Japanese spe- cies which has been introduced for control of woolly hemlock adelgid, Adelges tsugae Annand, in the eastern United States (Mc- Clure and Cheah, in press). Unfortunately, the name Pseudoscymnus Chapin is a junior homonym of Pseudoscymnus Herre 1935 (Vandenberg 2002). Herre’s (1935) usage of the name for a genus of kitefin shark (Squaliformes: Dalatiidae) predates Chap- in’s by more than a quarter century (Mould 1999) and therefore has priority according to article 52.3 of the International Code of Zoological Nomenclature (ICZN) (Interna- tional Commission on Zoological Nomen- clature 1999). Sasajiscymnus Vandenberg, new name Pseudoscymnus Chapin 1962: 50 (Type species: Scymnus hareja Weise 1879) (not Herre 1935, type P. boshuensis Her- ite) 4 Type species.—Scymnus hareja Weise 1879 (according to article 72.7 of the ICZN £999): Etymology.—The replacement name is composed of Sasaji + Scymnus (= a young animal, cub, whelp) masculine, and is ded- icated to Hiroyuki Sasaji, a leading spe- cialist in lady beetle systematics and an in- spiration to all who wish to understand the evolution and diversity of this important group. Remarks.—In accordance with articles 60.3 and 72.7 of the ICZN (1999), I pro- pose Sasajiscymnus as a replacement name and objective synonym of Pseudoscymnus Chapin. The more than 50 named species (Yu and Montgomery 2000) currently con- sidered to be in the genus Pseudoscymnus Chapin are hereby transfered to Sasajiscym- nus. To maintain brevity, only the following new combinations are proposed herein: Sasajiscymnus tsugae (Sasaji and Mc- Clure), S. kurohime (Miyatake), S. seboshii (Ohta), S. sylvaticus (Lewis), S. pilicrepus (Lewis) and S. guingquepunctatus (Weise). It is interesting to note that the name Scymnus also has entered into homonymy. It has been proposed at least twice as a ge- neric name for a shark (Scymnus Cuvier 1816, type Sgualus americanus Gmelin 1789, and Scymnus Quoy and Gaimard 1824, type Scymnus brasiliensis Quoy and Gaimard 1824), but senior status is held by the coccinellid genus Scymnus Kugelann 1794 (type Scymnus nigrinus Kugelann 1794, by subsequent designation of Kor- schefsky (1931)). Likewise, the bionomial Scymnus brasiliensis appears as both a shark (Scymnus brasiliensis Quoy and Gai- mard 1824, above, presently classified in /s- istius) and as a lady beetle (Scymnus bras- iliensis Weise 1929, presently regarded as a 484 junior synonym of Diomus seminulus (Mul- sant 1850) (Gordon 1999). According to Article 57.8.1 of the ICZN (1999), such homonymy between identical species-group names in combination with homonymous generic names established for different nominal genera is to be disregarded. LITERATURE CITED Chapin, E. A. 1962. Pseudoscymnus, a new genus of Asiatic Scymnini (Coleoptera: Coccinellidae). Psyche 69: 50-51. Cuvier, G. 1816. Le Régne Animal distribué d’apres son organisation pour servir de base a l/histoire naturelle des animaux et d’introduction 4a l’anatomie comparée. Les reptiles, les poissons, les mollusques et les annélides. Edition |. Regne Animal, Paris. 2: i—xviii + 1-532. Gmelin J. EF 1789. Pisces. Jn Caroli a Linné, Systema Naturae per regna tria naturae, secundum classes, ordines, gerera, species, cum characteribus, differ- entiis, synonymis, locis, Lipsiae. Edition 13. 1: 1126-1516. Gordon, R. D. 1999. South American Coccinellidae Part IV: A systematic revision of the South Amer- ican Diomini, new tribe. Annales Zoologici (War- saw) 49 (Supplement 1): 1-219. Herre, A. W. C. T. 1935. Notes on fishes in the Zoo- logical Museum of Stanford University. II. Two new genera and species of Japanese sharks and a Japanese species of Narcetes. Copeia 1935 (no. 3): 122-127. International Commission on Zoological Nomencla- ture. 1999. International Code of Zoological No- menclature, Fourth Edition. International Com- mission on Zoological Nomenclature, The Natural History Museum, London. 306 pp. Korchefsky, R. 1931. Coleopterorum Catalogus, pars 118, Coccinellidae I, Berlin. 224 pp. Kugelann, J. G. 1794. Verzeichniss der in einigen Ge- genden Preussens bis jetzt entdeckten Kiéfer-Ar- ten, nebst kurzen Nachrichten von denselben. Fortsetzung, Neuestes Magazin fiir die Liebhaber der Entomologie 1(5): 513-582. McClure, M. S. and C. A. S-J. Cheah. In press. Re- leased Japanese ladybugs are multiplying and kill- ing hemlock woolly adelgids. Frontiers of Plant Science 50(2). Mould, B. 1999. Classification of the recent Elasmo- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON branchii, a classification of the living sharks and rays of the world, third edition. Copyright Brian Mould. 178 pp. (available on-line as a pdf file at: http://ibis.nott.ac.uk/elasmobranch.html) Mulsant, M. E. 1850. Species des Coléopteéres triméres sécuripalpes. Annales des Sciences Physiques et Naturelles, d’Agriculture et d’Industrie, Lyon 2: 1—1104. Quoy, J. R. and P. Gaimard. 1824. Description des poi- sons. Chapter 9, pp. 192—401, plates 43-65. In L. de Freycinet, Voyage autour du monde, entrepris par ordre du Roi, exécuté sur les corvettes de S.M. l’Uranie et la Physicienne, pendant les années 1817, 1818, 1819 et 1820. Pillet Ainé, Paris, Vol. 3. Sasaji, H. and M. S. McClure. 1997. Description and distribution of Pseudoscymnus tsugae sp. nov. (Coleoptera: Coccinellidae), an important predator of hemlock woolly adelgid in Japan. Annals of the Entomological Society of America 90: 563-568. Vandenberg, N. J. 2002. Family 93. Coccinellidae La- treille, 1807, pp 371-389. Jn Arnett, R. H., Jr. and M. C. Thomas, eds. American beetles, Polyphaga: Scarabaeoidea through Curculionoidea, vol. 2: 95-97. Weise, J. 1879. Beitrage zur Kaferfauna von Japan. Deutsche Entomologische Zeitschrift 23: 149— ils. Weise, J. 1929. Ergebnisse einer zoologischen For- schungsreise nach Westindien von Prof. W. Ku- kental und Dr. R. Hartmeyer im Jahre 197. Wes- tindische Chrysomeliden und Coccinelliden, 2, Coccinelliden. Zoologischen Jahrbucher, Abtei- lung fiir Systematik Oekologie und Geographie der Tiere, supplement, 16: 11—34. Yu, G., M. E. Montgomery, and D. Yao. 2000. Lady beetles (Coleoptera: Coccinellidae) from Chinese hemlocks infested with the hemlock woolly adel- gid, Adelges tsugae Annand (Homoptera: Adel- gidae). The Coleopterists Bulletin 54: 154-199. Natalia J. Vandenberg, Systematic Ento- mology Laboratory, PSI, Agricultural Re- search Service, U.S. Department of Agri- culture, c/o National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC-168, Washington, DC 20013- 7012, U.S.A. (e-mail: nvandenb@ sel.barc. usda. gov) PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 485-487 NOTE First Report of the 13-Year Periodical Cicada, Magicicada tredecim (Walsh and Riley) (Hemiptera: Cicadidae) in Maryland Periodical cicadas (Magicicada spp.) are restricted to the eastern United States and attract attention from the general public be- cause of the amazing numbers of individ- uals that typically emerge. Marlatt (1907) designated specific Roman numerals for the possible 17-year periodical cicada broods (I-X VII) and the possible 13-year periodi- cal cicada broods (XVIII-XXX). The au- thor has been engaged in determining the extent of the range of each periodical cicada brood found in the mid-Atlantic area since Brood IIT emerged in 1996. Historically, only two broods of period- ical cicadas have been recorded from Mary- land. Brood II occurred in 1996 and was restricted to the southern Maryland counties of Calvert, Charles, and St. Mary’s, and was unexpectedly found to be absent just south of the region below Mattapany Road be- tween St. Mary’s City and St. James, St. Mary’s County, Maryland, south to Point Lookout, an area that appeared otherwise suitable for periodical cicadas. Brood X, the most widespread and abundant brood in Maryland occurring in the central and west- ern sections of the state, emerged in 1987 and will soon emerge again in 2004. Brood V has been recently documented in the westernmost county in Maryland, Garrett County, in 1999 (J. Zyla, in litt.). Although no 13-year cicadas had been reported from Maryland, and no emergence of any peri- odical cicada brood was expected in Mary- land during 1998, scattered reports of pe- riodical cicadas during May 1998 led to a survey of the magnitude and geographical extent of this emergence. In May 1998, the emergence year for 13-year Brood XIX far- ther south, a single live Magicicada male was presented to the author by a local res- idence living at the end of Demko Road in Dameron, Maryland. At first it was thought this specimen may be a two-year delayed 17-year cicada from Brood II. During the next week however, more reports of peri- odical cicadas surfaced. Subsequent visits to the neighborhood yielded hundreds of emerging individuals. Because the Brood II emergence was well documented in Maryland in 1996 and its distribution mapped out, the area south of the Brood II emergence was selected for study. The same survey system used for the Brood II study was employed. Beginning with the area where the initial specimen was found, the study area was crisscrossed by vehicle and each woodlot in the study area was surveyed for the presence of Mag- icicada (represented by either specimens or sound recordings). Voucher specimens were collected at various sites and sent to the University of Michigan, Museum of Zool- ogy, along with sound recordings for con- firmation. These specimens were confirmed to be M. tredecim (Walsh and Riley), a 13- year periodical cicada species and not M. septendecim (L.), the 17-year periodical ci- cada from Brood II, found just north of this area. This is the first report of Magicicada tredecim in Maryland. Although specimens of M. tredecim and M. septendecim are dif- ficult to distinguish, M. tredecim has a dom- inant call pitch (or frequency) of 1.0—1.1 kHz, while M. septendecim has one of 1.3 to 1.45 kHz (Marshall and Cooley 2000). The call recordings from the southern Maryland emergence had a dominant pitch of 1.1 kHz (D. Marshall, personal commu- nication) confirming that the emerging ci- cadas were M. tredecim and not M. septen- decim. In addition, the M. tredecim speci- mens were characterized by mostly orange abdominal sternites, rather than the darker coloration of M. septendecim. 486 Fig. 1. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ~ } ¥ Ant ty ot Amst f { fa Ai ce | c u > TS di a if Y abay 5 Zi; ths *f ve Aes The 1998 emergence of the 13-year periodical cicada (Magicicada tredecim) in St. Mary’s County, Maryland. This represents the northernmost locality of this species in eastern North America. The 1998 emergence of Brood XIX in Maryland was restricted to lower St. Mary’s County, in and around the small town of Dameron, 8.0 miles south of Lexington Park and 1.8 miles north of Ridge. Only Magicicada tredecim were collected and re- corded. The first occurrence was the lone individual provided to the author on May 9. The overall emergence began and _ be- came noticeable on May 15, 1998, and in- dividuals were collected as late as June 9. The emergence density appeared similar to that observed during emergences of other Magicicada broods in Maryland. No chorus was heard after June 9, possibly because the weather turned unseasonably wet and cold. Brood XIX periodical cicadas emerged in 1998 in large numbers in a very small area in St. Mary’s County, Maryland. The coun- ty is isolated by water, the Potomac River on the west and the Chesapeake Bay to the east. Just north of this area, Brood II emerged in great numbers in 1996. The contact area between these two broods was Mattapany Road, which runs in a west to east direction. To the north of this road, Brood II species of M. septendecim, M. sep- tendecula Alexander and Moore and M. cassinii (Fisher) occurred in 1996. South of this road, M. tredecim of Brood XIX was present in 1998, while M. tredecula Alex- ander and Moore and M. trecassini Alex- ander and Moore were noticeably absent. There is no obvious difference between the forest types found on either side of Matta- pany Road. Why a relict population of M. tredecim has survived in such a small area just south of and in contact with a 17-year periodical cicada brood is unknown. The nearest known location of M. tredecim is 60 miles due south of Dameron, Maryland, in St. James City and Charles City counties, Virginia (Sahli and Ware 1998; previously the northernmost reported localities of M. tredecim). This new site constitutes the northernmost known locality of Brood XIX Magicicada tredecim occurrence in the eastern United States (Cooley et al. 2003). The Maryland emergence is now the north- ernmost known occurrence of Brood XIX Magicicada tredecim in the eastern United States (Cooley et al. 2003) and is a consid- erable extension of the known geographic range of M. tredecim. Acknowledgments.—I thank Patty Craig for supplying the first specimen. Thanks are also due to David Marshall, who confirmed the identity of Magicicada tredecim from Maryland by analysis of the specimens and song recordings sent to the University of Michigan, Museum of Zoology. I also VOLUME 106, NUMBER 2 thank the anonymous reviewers for their suggestions and time. LITERATURE CITED Cooley, J. R., C. Simon, and D. C. Marshall. 2003. Temporal Separation and Speciation in Periodical Cicadas. BioScience 53(2): 151—157. Marlatt, C. L. 1907. The Periodical Cicada. United States Department of Agriculture, Bureau of En- tomology. Bulletin no. 71. Marshall, D. C. and J. R. Cooley. 2000. Reproductive 487 character displacement and speciation in periodi- cal cicadas, with description of a new species, 13- year Magicicada neotredecim. Evolution 54: 1313-1325. Sahli, H. F and S. Ware. 1999. Oviposition Sites and Habitats of 13-Year Periodical Cicadas (Brood XIX) in Eastern Virginia. Virginia Journal of Sci- ence 50(2): 168. John D. Zyla, J.F. Taylor, Inc., Lexington Park, MD 20680, U.S.A. (e-mail: jzyla@ Jfti.com) PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 488-490 NOTE Reports of Four Year Accelerated Occurrences of the 2004 Emergence of Periodical Cicadas, Magicicada Spp. (Hemiptera: Cicadidae) Brood X in Maryland, Virginia, and the District of Columbia Areas of the mid-Atlantic region of the United States (Maryland, Virginia and the District of Columbia) experienced an un- expected sporadic emergence of periodical cicadas (Magicicada spp.) in 2000. Peri- odical cicada emergences in this area attract public attention due to their abundant num- bers, with population densities approaching 1 million individuals per acre (Dybas and Davis 1962). Each regional Magicicada emergence in a specific year is designated as a brood. Broods have been differentiated by an assigned Roman numeral reflecting their order of emergence (Marlatt 1907). There are twelve known 17-year broods (I— X, XIII, and XIV) and three 13-year broods (XIX, XXII, and XXIII). Since Brood X is expected to emerge in 2004 in this same area, it is believed that the 2000 emergences were 4-year accelerations of Brood X. Oth- er 4-year accelerations of Brood X in 2000 were reported from Ohio (Kritsky and Si- mon 1996), Indiana, and Illinois (Cooley, Richards, Marshall, personal communica- tion). In the past, Brood VI has been reported sporadically throughout the mid-Atlantic region. However, recent studies by Cooley and Marshall (personal communication) suggest that Brood VI occurs further west, in western portions of North and South Car- olina, and northern Georgia. Because the 2000 mid-Atlantic emergence was outside the known area of Brood VI, researchers have theorized that this emergence was a 4- year acceleration of Brood X. If this hy- pothesis is true, then all the reports should be contained in the known range of Brood X. Since 1996, the author has documented the presence or absence of periodical cicada broods (II, V, VIII, and IX) in the mid-At- lantic region both in the field and through reviewing historical records in the litera- ture. To date, there has not been a single record of two different broods occurring in the same woodlot. No known populations of the four species of periodical cicadas (3 species of the 17-year cicada, M. septen- decim (L.), M. septendecula Alexander and Moore, and M. cassinii (Fisher), and one species of the 13-year cicada, M. tredecim (Walsh and Riley), found in the mid-Atlan- tic region (Pennsylvania, West Virginia, Virginia, Maryland, District of Columbia, and Delaware) overlap. In fact, many con- tact areas where two different broods meet are characterized by a gap where no peri- odical cicadas are known to occur. Based on this information, it is the author’s opin- ion that any populations that occur in off- years within a known well-established brood area are most likely accelerations or decelerations of that well-established brood and not a separate brood. Brood X, in the mid-Atlantic, is found in northern Virginia, western and central Maryland, northern Delaware and southern Pennsylvania, as well as, the District of Co- lumbia. During the unexpected 2000 emer- gence, reports of stragglers were solicited throughout the area. All the information ob- tained was organized by county and placed on a map of the known range of Brood X. Following is a list of the recorded popula- tions that were reported to the author during May and June of 2000. All the sites re- ported in 2000 were within the historical boundaries of Brood X. A total of 124 independent reports were obtained from the Maryland, Virginia, and District of Columbia area. Six were re- ceived from the District of Columbia, 90 VOLUME 106, NUMBER 2 Frome reports were received from Maryland (7 counties), and 28 were recorded from Vir- ginia (4 counties). No reports were obtained from outside the known historic area of Brood X. All the reported 2000 sightings were of sporadic occurrences and although they may have contained dense local emer- gences of periodical cicadas, they were not the wide-ranging, regional emergences typ- ical of these species. An entomologist residing in Bowie, Prince George’s County, Maryland, record- ed the daily numbers of periodical cicada exuviae collected in her yard. She reported that mud turrets began to appear on May 4. On May 9, 62 nymphs were found walking around from 10 pm to midnight. Almost 3,000 (2902) exuviae were collected from the period May 8th to June Ist, 2000. How- 489 oe ees Four-year accelerations reported in 2000 by county within the historic range of Brood X. ever, throughout the remainder of the neigh- borhood, most yards had no periodical ci- cadas emerging at all. This same neighbor- hood was the site of an abundant emergence of Brood X in 1987. There are similar re- ports throughout the 2000 emergence area. The following reported emergences of periodical cicadas in 2000. DISTRICT OF COLUMBIA: Georgetown, Northwest. VIRGINIA: Alexandria, Arlington, Fairfax, and Prince William counties. MARY- LAND: Allegany, Anne Arundel, Balti- more, Cecil, Howard, Montgomery, and Prince George’s counties. Acknowledgments.—I thank Gaye Wil- liams for her encouragement and soliciting and/or supplying many of the reports ob- tained. I also thank the anonymous review- ers for their suggestions and time. 490 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED cadidae: Magicicada spp.) in Ohio. Ohio Journal Science 96(1): 27-28. Marlatt, C. L. 1907. The Periodical Cicada. United Dybas, H. S. and D. D. Davis, 1962. A population States Department of Agriculture, Bureau of En- census of seventeen-year periodical cicadas (Ho- tomology. Bulletin no. 71. moptera: Cicadidae: Magicicada). Ecology 43: 432-444. John D. Zyla, J.F. Taylor, Inc., Lexington Kritsky, G. and S. Simon. 1996. The unexpected 1995 Park, MD 20680, U.S.A. (e-mail: jzyla@ emergence of periodical cicadas (Homoptera: Ci- Jfti.com) PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 491-492 Book REVIEW Encyclopedia of Insects. Vincent H. Resh and Ring T. Cardé, editors. 2003. Aca- demic Press, San Diego. ISBN-O-12- 586990-8. 1266 pp. $99.95 (hardcover). Web site at http://www.apnet.com/ insects/ This weighty tome is a compendium of all things insect, with a few things arachnid thrown in for good measure. True to its title, all of the information is presented alpha- betically in a series of articles, beginning with “Acari”? and ending with “‘Zygento- ma’ Sorry, no K’s, Q’s, or X’s! But those looking for essays covering katydids, killer bees, and queens need not despair! They are directed to “‘Orthoptera,”’ “Neotropical Af- rican bees,”’ and “‘Caste.”’ The Encyclopedia is handsomely and sturdily bound. Its colorful jacket bears what is now almost obligatory for any in- sect book of caliber, praise from Edward O. Wilson and a somewhat tongue-in-check foreword by May Berenbaum. The Ency- clopedia is printed on heavy acid-free paper and gives every appearance of being able to survive a long life of use. The book features 271 self-contained ar- ticles written by some of the world’s lead- ing authorities. The list of 164 contributors reads like an international ““who’s who” in entomology. Each contributor listed is fur- ther identified by their institution and a list of their titles that appear in the work. Edi- tors Resh and Cardé are to be commended for assembling such a fine group of experts. A guide to the Encyclopedia lays out the format of the work, including subject areas and article format. Each article is fully cross-referenced and followed by a brief bibliography that serves as an extension of the article itself rather than an exhaustive accounting of the articles’ subject. Nearly all of the articles are peppered with black and white or color illustrations and photo- graphs. Most of the color photos used are bright, sharp images that seem to jump off the page. Curiously, the sources of many of the images used, other than letter-opening photos, are not credited in their captions, nor are those credits to be found anywhere in the book. Readers can locate topic articles either al- phabetically in the Table of Contents or find them listed in the Contents by Subject Area. In this section, twelve subject areas are identified, including anatomy, physiology, behavior, evolution, reproduction, develop- ment and metamorphosis, major groups and notable forms, interactions with other or- ganisms, interactions with humans, habitats, ecology, and history and methodology. The content of the subject areas “interactions with humans” and “history and methodol- ogy” might have been more finely tuned and divided into three categories: medical and veterinary entomology, cultural ento- mology, and applied entomology. Stil, the interested reader should have no trouble in locating pertinent articles of interest. Thirty-three orders of insects are covered in the book, including the recently discoy- ered Mantophasmatodea, the first new in- sect order to have been described in more than 80 years. Some readers may quibble with the usage of some ordinal names, such as Archaeognatha for Microcoryphia, Em- bidiina for Embioptera, and Zygentoma for Thysanura. Readers seeking information on the order Hemiptera will have to work a little harder. Listed in the table of contents, but not the contents by subjects, readers are directed to hop, skip and jump through the entire work to find separate essays on the suborders Auchenorrhyncha, Prosorrhynca (Heteroptera and Coleorrhynea for the un- initiated, including myself!) and Sternor- rhynca. Of particular interest to this reviewer, and what I think sets this book apart from other comprehensive works on insects thus far, 492 are the detailed articles on cultural ento- mology, folk beliefs and superstitions, mu- seums and display collections, biotechnol- ogy and insects, insect zoos, photography of insects, and teaching resources. These benchmark articles are important resources for not only entomologists, but also for ed- ucators and naturalists and go a long way to increasing the utility of the Encyclopedia to an audience well beyond the entomolog- ical community. Another useful feature is a glossary de- signed specifically to define approximately 800 terms within the context that they are used in the text. I think non-specialist read- ers in particular will find the glossary very accessible. The Encyclopedia also includes a subject index that includes taxa cited in the text. In the Preface, the editors’ stated goal was “‘to convey the exciting, dynamic story of what entomology is today. It is intended to be a concise, integrated summary of cur- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rent knowledge and historical background on each of the nearly 300 entries presented. Our intention has been to make the ency- clopedia scientifically uncompromising; it is to be comprehensive but not exhaustive.” They also wanted to “cover the diverse in- terests” of not only entomological special- ists and generalists, but also educators, stu- dents, hobbyists, and naturalists. To these ends the editors and contributing writers have clearly succeeded in creating a rich, spectacular volume that is a must for those who demand libraries with thorough, up-to- date, and superbly accessible information on insects. Arthur V. Evans, Research Associate, Department of Entomology, National Mu- seum of Natural History, Smithsonian In- stitution, and Department of Recent Inver- tebrates, Virginia Museum of Natural His- tory. Home address: 1600 Nottoway Ave., Richmond, WA 23227, U.S.A. (e-mail: arthurevans @ earthlink.net) PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 493-494 Book REVIEW Water Beetles of South Carolina (Cole- optera: Gyrinidae, Haliplidae, Noteridae, Dytiscidae, Hydrophilidae, Hydraenidae, Scirtidae, Elmidae, Dryopidae, Limnichi- dae, Heteroceridae, Psephenidae, Ptilo- dactylidae, and Chelonariidae). Janet C. Ciegler. South Carolina Agriculture and Forestry Research System. 2003. 210 pp. ISBN 0-9712527-7-7. U.S. $40.00. As an identification manual, Janet Cie- gler’s ““‘Water Beetles of South Carolina” goes far beyond any other state-wide treat- ment of the aquatic beetle families. It is “Volume 3” of the “Biota of South Caro- lina”’ series which has thus far been devot- ed to highly speciose groups of Coleoptera. Florida is the only other southeastern state for which the entire water beetle fauna has been comparably covered at the species lev- el in a single volume (Epler 1996, Young 1954). This book additionally includes the terrestrial Hydrophilidae (Sphaeridiinae) and several families that are semi-aquatic. While it does not describe larval stages, it offers illustrations of adult examples of each genus known to be represented in the state, and those expected to occur there. De- scriptions of 374 species in 96 genera are given, with notes and citations of the most recent monographic works for each group. General distributional data are given for each species in addition to the specific lo- calities known for the state, and often with notes on habitats and seasonal occurrence. The eye-catching color cover and photo- graphs will get the attention of any ento- mologist. Along with the keys and comparative di- agnoses, the various illustrations make this volume more useful and attractive. Minor shortcomings with some of the reproduced photographs of the smaller species are ev- ident, e.g., being out of focus, or the spec- imen chosen does not depict the body out- line accurately (elytra separated or head de- flexed). However, these are probably the first published images of many of these spe- cies, and with the associated life-size sil- houettes or scale lines, are nonetheless in- formative. The drawings among the keys and descriptions are especially helpful in il- lustrating some features that are difficult to describe or interpret, along with compara- tive ones for and among closely related spe- cies, and the book includes figures of many of these not available in other identification manuals. The family key includes terrestrial groups as well, helpful in sorting out the frequent drowned contaminants in aquatic samples, and instructs the user to “‘see gen- eral reference, When a large assemblage of unrelated beetle families is treated with this level of detail by a single author, occasional errors or omissions can be expected. A few minor items are noted here to serve as construc- tive clarifications for users of this fine vol- ume. In two of the anatomical diagrams, Fig- ures 5.1 and 6.1, the posterior part of the prosternum is labeled as “‘mesosternum”’; the actual mesosternum is mostly hidden beneath the midlegs of these beetles and seen as a small bridge between them, or is part of a ventral keel. In the diagram of Suphis inflatus (Figure 4.1) the numbering of the visible abdominal sterna is mislead- ing, and not “homologous” to the number- ing used for the Acilius fraternus (Figure 5.1) because the basal (divided) segment is unnumbered; sterna 2 and 3 (labeled | and 2 in the figure) are fused as a single large sclerite in all Noteridae and the suture is generally not discernable (Roughley 2001). One probable misidentification should be pointed out for Figure 4.4, Notomicrus nan- ulus (Noteridae). This species is much more elongate-oval and of a paler color than the beetle shown, which may be.a specimen of Desmopachria (Dytiscidae). A likely omis- 494 sion in the book is the family Lutrochidae, represented in the region by Lutrochus la- ticeps Casey (Spangler et al. 2001); no re- cords are known for South Carolina and it does not quite fit the author’s criteria for inclusion (page 3), but the beetle would be expected to occur there as it is known from a broad area including Alabama, Tennessee and Maryland. In the coverage of Hydrophilidae, a re- cent review of the distribution, habitats and literature for Hydrobiomorpha casta (Say), including a northern range extension for Virginia (Steiner 1996) would have been worthwhile to cite under that species. Users of this volume should be aware that the hy- drophilid subfamilies Hydrochinae and Hel- ophorinae are elevated to family level by many hydrophiloid systematists, but with continuing disagreement, as reviewed by Van Tassell (2001); the author uses the trib- al classification of Hansen (1991) but chose not to use his elevated familial scheme. The book offers the user full citations of the related literature, a glossary of anatom- ical and other terms, appendices listing lo- calities with physiographic regions and spe- cies newly reported for the state, and an in- dex of taxa including synonyms. While re- searchers using macroinvertebrates for water quality assessment may need to con- sult other references in order to identify beetle larvae, this is nevertheless the most comprehensive coverage of the species for the southeastern region. It will be useful in South Carolina’s neighboring states for PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON identification at the species level, and a good guide to genera and families of east- ern North America. The book is a tool for the regional biogeographer, teacher of nat- ural history, heritage program biologist, and museum curator, and may inspire other state-wide works of a similar caliber. LITERATURE CITED Epler, J. H. 1996. Identification Manual for the Water Beetles of Florida (Coleoptera: Dryopidae, Dytis- cidae, Elmidae, Gyrinidae, Haliplidae, Hydraeni- dae, Hydrophilidae, Noteridae, Psephenidae, Ptil- odactylidae, Scirtidae). Bureau of Water Resource Protection, Florida Department of Environmental Protection, Tallahassee. 259 pp. Hansen, M. 1991. The Hydrophiloid Beetles. Phylog- eny, classification and a revision of the genera (Coleoptera: Hydrophiloidea). Biologiske Skrifter 40: 1-367. Roughley, R. E. 2001. Noteridae C. G. Thompson, 1857, pp. 147-152. Jn Arnett, R. H. Jr. and M. C. Thomas, eds. American Beetles. Volume 1. CRC Press, Boca Raton, Florida. Spangler, P. J., C. L. Staines, P. M. Spangler, & S. L. Staines. 2001. A checklist of the Limnichidae and Lutrochidae (Coleoptera) of the world. Insecta Mundi 15(3): 151-165. Steiner, W. E., Jr. 1996. Hydrobiomorpha casta (Say) in Virginia (Coleoptera: Hydrophilidae). Banister- ia 7: 53-55. Van Tassell, E. R. 2001. Hydrophilidae Latreille, 1802, pp. 187-208. Jn Arnett, R. H. Jr. and M. C. Thom- as, eds. American Beetles. Volume 1. CRC Press, Boca Raton, Florida. Young, FE N. 1954. The water beetles of Florida. Uni- versity of Florida Press, Gainesville. 238 pp. Warren E. Steiner, Jr., Department of En- tomology, NHB-187, Smithsonian Institu- tion, Washington, DC~ 20560, U.S.A: (e-mail: steiner.warren@nmnuh.si.edu) PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 495-496 REPORTS OF OFFICERS EDITOR Volume 105 of the Proceedings included four issues with a total of 1,104 pages. Ninety-seven regular papers, 23 notes, 4 book reviews, 3 obituaries, the membership list, minutes of Society meetings, reports of officers, instructions for authors, and the ta- ble of contents for volume 105 were pub- lished. About 115 manuscripts (regular pa- pers and notes) were submitted for consid- eration for publication from November 1, 2002, to October 31, 2003. Memoir No. 24, ““Holcocerini of Costa Rica (Lepidoptera: Gelechioidea: Coleo- phoridae: Blastobasinae)”” by David Adam- ski, was published in December 2002. Memoir No. 25, “A Catalog of the Ceci- domylidae (Diptera) of the World” by Ray- mond J. Gagné, is in press and will be pub- lished in February 2004. I extend thanks to Ray Gagné, book re- view editor, for his excellent work in ob- taining book reviews, and to Ray, Tom Henry, and Wayne Mathis of the Publica- tions Committee for their encouragement and support. Also, I am grateful to the many reviewers for their time-consuming efforts and constructive reviews of manu- scripts. Their contributions are essential to help increase the quality of papers pub- lished in the Proceedings. Respectfully submitted, David R. Smith, Editor TREASURER SUMMARY FINANCIAL STATEMENT FOR 2003 Special General Publication Total Fund Fund Assets Assets: November 1, 2002 $43,979.32 $196,717.44 $179,896.87 Total Receipts for 2003 $76,441.84 $13,060.61 $89,502.45 Total Disbursements for 2003 $66,853.72 $73,029.74 $139,883.46 Assets: October 31, 2003 $53,567.44 $136,748.31 $190,315.75 Net Changes in Funds $9,588.12 ($59,969.13) ($50,381.01) Audited by the Auditing Committee, November 22, 2002, consisting of Allen L. Norrbom, Chairman, Steven W. Lingafelter, and Michael W. Gates. Presented to the membership at the meeting of December 4, 2002. Respectfully submitted, Michael G. Pogue, Treasurer 496 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MEMBERSHIP SECRETARY In 2003, the Society received applications for membership from 23 people: Patrick B. Beauzay Susan Broda Ebru Gil Cilbiroglu Andrew Richard Cline Jane Earle Colin Favret Dennis Heimdal Ronald A. Hellenthal Harold W. Ikerd Kevin Kirchner Heather Laflin Chris Looney John W. McCreadie Mark A. Muegge Jose Luis Navarette-Heredia Cervantes Peredo John Dennis Plakidas Valerie Schawaroch Doug Strom James Bolling Sullivan Lu Sun Nick Wiersema James R. Wiker Each applicant was sent a letter of welcome to the Society and his/her name was read at a regular monthly meeting. The number of applications decreased 8% from 2002. Other letters from the Membership Secretary included five letters to guest speakers, 11 letters to special fund contributors, two replies to requests for information, one thank you letter to an outgoing officer, and one letter of welcome to an incoming officer. Respectfully submitted, Holly B. Williams, Membership Secretary PRESIDENT-ELECT The 1,075th meeting of the Entomologi- cal Society of Washington was held on 11 June 2003. It consisted of the Annual Ban- quet, held this year at the Uniformed Ser- vices University of Health Sciences in Be- thesda, Maryland. A cash bar opened at 6:00 p.m., with the formal banquet follow- ing at 7:00 p.m. Approximately 65 mem- bers and guests were in attendance. At 8:00 p.m. the speaker, Carll Goodpasture, an en- tomologist, geneticist and freelance photog- rapher, was introduced by President-Elect Eric Grissell. The title of Carll’s talk was “Entomological Crossingover: Where Art and Science Meet.” The talk was illustrated with Carll’s photographs and two DVD pre- sentations. The meeting ended at about 9:30 p.m. Respectfully submitted, E. Eric Grissell, President-Elect PROC. ENTOMOL. SOC. WASH. 106(2), 2004, pp. 497-499 SOCIETY MEETINGS 1,076th Regular Meeting—October 2, 2003 The 1,076th regular meeting of the En- tomological Society of Washington (ESW) was called to order in the Cathy Kerby room of the National Museum of Natural History, Washington, D.C., by President- elect Eric Grissell, standing in for President Jon Mawdsley, at 7:06 p.m. The meeting was attended by 18 members and 10 guests. The minutes of the 1,074th and 1,075th meetings were approved with minor modi- fication. There were no new applicants for mem- bership or new members present. Four guests were introduced. In miscellaneous business, Dave Furth announced that the ESW website has been recently updated. For exhibits, Art Evans gave a rapid-fire slide show of impressive new shots of ar- thropods from Virginia, North Carolina, and Tennessee. Dave Furth displayed the books “Results of the Zoological Missions to Australia of the Regional Museum of Natural Sciences of Turin, Italy” by M. Daccordi and P. M. Giachino (eds.) and “Chrysomelidae: The Leafbeetles of Eu- rope and the Mediterranean Area” by A. Warchalowski. Dave Furth passed around a sticky trap that had captured an invasive close relative of the venomous Brown Re- cluse spider in the Natural History Building basement. Dave Furth introduced the speaker, Dr. Chris Desjardins, a Ph.D. student of the University of Maryland and Maryland Cen- ter for Systematic Entomology. His presen- tation was entitled “‘Diparine Wasp Diver- sity and Collecting in South Africa and Australia.’” The pteromalids are generally the waste-basket family of Chalcidoidea, with some 20 to 30 subfamilies. One of them is Diparinae, a group comprising Over 100 species in 31 genera, with lengths often in the 1-2 mm range. Sexual dimorphism is widespread, with many females wingless, though little is known of their habits. The subfamily is cosmopolitan, with highest ge- neric richness in Australia and South and East Africa. Off he went, with his sights set on fresh specimens for his molecular anal- ysis. Desperately trying to follow the un- cooperative rains with his yellow pan-traps, Chris was getting pretty dejected until he was side-tracked by some harrowing trials of the field, which were eventually com- pensated for in chalcidoids. There were many rhinos but it was a tick that nailed him. The meeting was adjourned at 8:25 p.m. Refreshments were provided by the Society. Respectfully submitted, Stuart H. McKamey Recording Secretary 1,077th Regular Meeting— November 6, 2003 The 1,077th regular meeting of the En- tomological Society of Washington (ESW) was called to order in the Cathy Kerby room of the National Museum of Natural History, Washington, D.C., by President Jon Mawdsley, at 7:12 p.m. The meeting was attended by 18 members and 13 guests. The minutes of the 1,076th meeting were approved as read. There were 17 new applicants for mem- bership, which will be re-read at the Janu- ary meeting. One new member was present. Four guests were introduced. The ESW nominating committee, Dave Furth and Dave Smith, reported that all but one of the current officers have offered to continue in their positions. Jason Hall was nominated for President-Elect. Voting will take place at the next meeting. For exhibits, Jon Mawdsley had the new book, “A Field Guide and Identification Manual for Florida and Eastern U.S. Tiger Beetles,” by Paul M. Choate, Jr. Edd Bar- 498 rows welcomed feedback on his arthropod checklist of Dyke Marsh, which can be found at _ http://biodiversity.georgetown. edu/index.cfm. Dave Furth had on hand the coffee-table book, *‘Wildlife Spectacles,” of the CEMEX publication series, by R. A. Mittermeier et al. Dave Furth introduced the speaker, Dr. Stefan Cover, of the Museum of Compara- tive Zoology, Harvard University, who spoke about “‘Ant Systematics in the 21st Century: North America, a Case Study.” Dr. Cover asserted that a third directional advance, after phylogenetics and molecular analysis, is emerging among ant workers. The catalyst was the key to world genera published in 1990 and development of ad- ditional collecting techniques, which has re- sulted in a flood of new specimens and a push for biodiversity inventorying. The au- tomontage imaging system is invaluable in this endeavor, as have been the 10-day ant courses being offered for the last three years, which have trained 74 students and served as the nerve center of the colony of busy ant workers. Dr. Cover argued that the world ant fauna will never be known unless ‘“‘business-as-usual”’ is scrapped for weav- ing together smaller pieces, such as regional faunal lists, making more information avail- able on the WWW, and reliance on digital images accompanied by simplified descrip- tions. The meeting was adjourned at 8:37 p.m. Refreshments were provided by the Society. Respectfully submitted, Stuart H. McKamey Recording Secretary 1,078th Regular Meeting— December 4, 2003 Despite the forecast for snow and the threat of no electrical power, and in the face of stiff competition from the annual tree- lighting gala at George W.’s place, the stal- wart attendees (18 members and 9 guests) of the 1,078th regular meeting of the En- tomological Society of Washington were all PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON settled in for a long winter’s nap by Presi- dent Jonathan Mawdsley at 7:05 p.m. I se- riously doubt that there were visions of sug- ar-plums dancing in anyone’s head ... but I could be wrong. Minutes of the 1,077th meeting were read by John Brown, pinch- hitting for Recording Secretary Stu Mc- Kamey, and were approved following triv- ial comments. As the December meeting functions as the Annual Meeting for the Society, there were brief reports from the officers. Dave Smith (editor) reported that volume 105 of the Proceedings included a whopping 1,104 pages—our largest volume ever. Jon Mawdsley reported on the chores handled by Holly Williams, our corresponding sec- retary/membership chair (a new amalgam- ation of duties). Dave Furth (program co- chair) commented on the status of our pro- gram for the upcoming year. Furth also pro- vided a brief overview of our financial state compiled by Mike Pogue, our Treasurer. With total assets of $190,315 and change, the Society is in good financial shape. The reports were followed by the annual election of officers, with all but the presi- dent coerced into continuing to perform in their present capacity: Buck Lewis as Cus- todian, Dave Furth and John Brown as Pro- gram Co-chairs, Holly Williams as Corre- sponding Secretary, Stu McKamey as Re- cording Secretary, Dave Smith as Editor, Mike Pogue as Treasurer, and newly elected President-Elect Jason Hall. Jon Mawdsley will serve as Immediate Past President when President Eric Grissell takes the reins of the Society in January. The notes and exhibitions portions of the program was mercifully brief, unlike this report, with Mawdsley displaying a new book, “The Pollinator Conservation Hand- book” (published by the Xerces Society); Warren Steiner sharing a new book, “Water Beetles of South Carolina’; and Furth aw- ing the audience with two new ‘tees’ pur- chased at the recent ESA meeting. Finally, Program Co-chair Dave Furth in- troduced the evening’s speaker, John Stra- VOLUME 106, NUMBER 2 zanac, from the Department of Plant and Soil Sciences/Entomology at West Virginia University, Morgantown, whose talk was ti- tled ““The Agony and the Ecstasy: Results of Seven Years of Sampling Central Ap- palachian Arthropod Diversity.”’ Dr. Stra- zanac presented an interesting array of fac- toids, methodologies, and species accumu- lation curves resulting from this intensive survey in West Virginia and Virginia. The primary focal groups of the sampling effort were macrolepidoptera, Symphyta, Carabi- dae, Araneae, and Tachinidae. Sampling techniques included canvas bands, foliage pruning, light traps, malaise traps, and pit- fall traps. Dr. Strazanac estimated that well over a million specimens were handled dur- ing the course of the work. 499 In his final action as President of the So- ciety, Mawdsley handed off the gavel to in- coming President Grissell. It was a touching moment, as usual—yawn. In his first action as President, Grissell boldly and confident- ly, but in a gentler and kinder manner, ad- journed the meeting at about 8:30 p.m. Re- freshments following the meeting were pro- vided by the Society, amply augmented by Ralph Eckerling, whose nog, hard stuff, and Christmas cookies added to the festive mood of the post-meeting chatter. The room was vacated by a little after 9:00 p.m. to avoid being left in the dark, something most of us are used to anyway. Respectfully submitted, John W. Brown Pinch-hitting Recording Secretary NOTICE OF NEW PUBLICATION A Catalog of the Cecidomyiidae (Diptera) of the World Entomological Society of Washington Memoir No. 25 This catalog, the first complete catalog of the family since 1913, lists the 5,451 species and 598 genera of living and fossil Cecidomyiidae or gall midges of the world. It provides information on species distribution, hosts, and types, and original and subsequent helpful references. Within subfamilies, genera are listed in alphabetical order but are each cross referenced in an appendix where they are arranged in an annotated classification. One new species is named, many new names, new synonyms, and new combinations are proposed, and several type species are designated. A single index lists all generic and specific names of Cecidomyiidae with their authors, as well as hosts, host family for plants, host order and family for arthropods, and order for fungi. ENTOMOLOGICAL SOCIETY OF WASHINGTON nttp://entomology.si.edu/ES W/ES W Menus. lasso Information and Officers Bylaws History First 100 Years Past Presidents History of the ESW Seal Information for Authors Available Publications Meetings Members Database Membership Application Subscription PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael E. SOU cS 210 aT Ee MON La Vn ACN FN Pe PPE PRR RL EON eae $10.00 A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. Eric CrerewanniuNienack ie (Scaauit ey PPylGOy yl a a 15.00 Revision of the Oriental Species of Aphthona Chevrolat (Coleoptera: Chrysomelidae), by Alexander S. MonsianiunOviancesteveniw wleinoatelter S49pp..2002) (so oo ee es i ee 40.00 Revision of the Genus Anoplophora (Coleoptera: Cerambycidae), by Steven W. Lingafelter and E. oS SATPSMI SIG eee: 00a REAR OE LE ed I Sg TO RTL NT 1g 30.00 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 __ $15.00 No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 15.00 No. 5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 ____ 15.00 No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi Bite kit ap wea pee tem kek SS hd AO eet On 15.00 No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato- porontdac);.by W. L. Grogan, Jr. and W. W. Wirth. {25 pp..1979) 0 12.00 No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. Re eetpre tee meme ee eee epee ee ee A a le ee 5.00 No. 14. Biology and Phylogeny of Curculionoidea, edited by R;.S. Anderson and C. H. C. Lyal. 174 jou Ua) a AE 3s he TAG De SIE ee 2 eae ee ee eee ee SRO ere REE CES) 25.00 No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: aga Seawhee by A. Borkent FHETO TLE Fest valed atl cc aia] 9h AS 12 PE ia eee Se eR, CO Rae ee Caine) Ab 25.09 No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 __ 25° ®) No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Kroinbein, edited byw Nomenand Ay SsMenkes 21Gipp: 1996), 2k et ee ee Ne MD No. 18. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and Vy CUM en UD Gr ro iee iu PBN PAS 0) cI AS 10) OU Ru ERLE ES ee ac, Ce AOA LAI! 25.00 No. 19. Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by PUTO nIASH ME yamEL Oop Yabo There team LM ry a ue Ne ee ee 18.00 No. 20. The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- POLC ete Sa DMMleomeemens tion west Mee ema iy. Po la ee | Ne Ne ee 12.00 No. 21. New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. FMP VTIG Teype EUS bf earn OE DSS Eas d 2S SEE SSO SSNS ce Oe Met) RS CL 12.00 No. 22. Systematics of the North American Species of Trichogramma Westwood (Hymenoptera: richorraminanadse) by Jonni Pmto. 287, pp. L990. 2 28.00 No. 23. Revision of the Net-Winged Midges of the Genus Blepharicera Macquart (Diptera: Blepha- riceridae) of Eastern North America, by Gregory W. Courtney. 99 pp. 2000 ___-.----------------------- 14.00 No. 24 Holcocerini of Costa Rica (Lepidoptera: Gelechioidea: Coleophoridae: Blastobasinae), by David PNGLALNAS lem LE ant T| Hee) ne ene Ng ees ie ane gee Me ee een 18.00 No. 25 A Catalog of the Cecidomyiidae (Diptera) of the World, by Raymond J. Gagné. 408 pp. 2004 50.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume to non-members and $25.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra on all orders. Dealers are allowed a discount of 10 percent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Treasurer, Entomological Society of Washington, “% Department of Entomology, Smithsonian Institution, P.O. Box 37012, MRC 168, Washington, D.C. 20013-7012. CONTENTS (Continued from front cover) PACHECO, JOSE, WILLIAM MACKAY, and CYNTHIA MORGAN—A new species of Gnamptogenys Roger of the sulcata group (Hymenoptera: Formicidae) from Bolivia ...... 434 PEREDO, LUIS CERVANTES—A4lloeorhynchus trimacula (Stein) (Heteroptera: Nabidae: Prostemmatinae), a predator of Rhyparochromidae (Lygaeoidea) associated with figs in IMPS STOOME BOE AME bat amb a ae a SAAR 5 a Ren Kn Men non Smear ciycis Mer Cnet 346 PINTO, JOHN D.—A review of the genus Doirania Waterston (Hymenoptera: Trichogrammatidae), with a description of a new species from North America .............. 352) POINAR, GEORGE, JR. and ALEX E. BROWN—A new genus of primitive crane flies (Diptera: Tanyderidae) in Cretaceous Burmese amber, with a summary of fossil tanyderids .......... 339 SCHAEFER, CARL W.—Key to the genera of New World Alydidae (Hemiptera: Heteroptera) .. 280 SPINELLI, GUSTAVO and ART BORKENT—New species of Central American Culicoides Latreille (Diptera: Ceratopogonidae) with a synopsis of species from Costa Rica ............ 361 STEINLY, BRUCE A.—Primary consumer and detritivore communities (Diptera: Ephydridae) in newly restored and constructed wetlands ...............ceeeeee cece ee eee eee tenner eect eee tees 460 TOGASHI, ICHIJI—Description of a new species of the genus Eriocampopsis Takeuchi (Hymenoptera: Tenthredinidae) from Japan ........... 0.6.22 eee eee e eee cnet eee e ee een eens 421 WHEELER, A. G., JR. and E. RICHARD HOEBEKE—New records of Palearctic Hemiptera (Sternorrhyncha, Cicadomorpha, Heteroptera) in the Canadian maritime provinces ........ 298 YASUNAGA, TOMOHIDE—A new genus and new species of mirine plant bug (Heteroptera: Miridae:) Marini) strom the Rey kena 0a eins ye hic Sh chs hooters tantra ets oleh tose oie ele ae tele 407 ZHOU, CHANG-FA, LU SUN, and W. P. McCAFFERTY—A new species of Brachycercus Curtis (Ephemeroptera: Caenidae) from China | ..............cseb scene cece eee enone cee rntanteesene ress 312 NOTES LAFLIN, H. M., P. J. GULLAN, and M. P. PARRELLA—Mealybug species (Hemiptera: Pseudococcidae) found on ornamental crops in California nursery production .............. 475 NAVARRETE-HEREDIA, JOSE LUIS and JESUS CORTES-AGUILAR—Beetles (Coleoptera) associated with the external debris of Atta mexicana (F. Smith) (Hymenoptera: Formicidae) from Ojuelos, Jaliscas Mexico yi fo.) Po) Babe. 11202 oe Senn dia he eae ee ne ar a 481 POINAR, GEORGE, JR. and JON GELHAUS—Larval development of Cylindrotoma distinctis- sima americana Osten Sacken, 1865. (Tipulidae: Cylindrotominae) on Maianthemum dilata- tum (Miliaceac) iam Califo cia’ Wes oe acs 2a rie alate Gi) le ele ovation ares Fe eT ae ee 478 ROBBINS, RICHARD G., MICHAEL J. BANGS, and JAMES E. KEIRANS—First report of Ixodes kopsteini Oudemans (Acari: Ixodida: Ixodidae) from the Kingdom of Cambodia, with a summary of known hosts of this tick in continental southeastern Asia ...............++++++- 472 VANDENBERG, NATALIA J.—Homonymy in the Coccinelidae (Coleoptera), or something fishy about Pseudoscymnus' Chapin .4 06). 6 see alee Sek sates dle Cae eo 2 oe eine we ane ee ee 483 ZYLA, JOHN D.—First report of the 13-year periodical cicada, Magicicada tredecim (Walsh and Riley) (Hemiptera: Cicadidae)}in Maryland 0.202. oe, re ieee ooh em ee lens oe aed aee 485 ZYLA, JOHN D.—Reports of four year accelerated occurrences of the 2004 emergence of peri- odical cicadas, Magicicada spp. (Hemiptera: Cicadidae) Brood X in Maryland, Virginia, and the District: of Columbian sy Ve ses COE ee ete ara et ae eh a 488 BOOK REVIEWS EVANS, ARTHUR V.—Encyclopedia of Insects, Vincent H. Resh and Ring T. Cardé, editors .... 491 STEINER, WARREN E., JR.—Water Beetles of South Carolina (Coleoptera: Gyrinidae, Haliplidae, Noteridae, Dytiscidae, Hydrophilidae, Hydraenidae, Scirtidae, Elmidae, Dryopidae, Limnichidae, Heteroceridae, Psephenidae, Ptilodactylidae, and Chelonariidae), by Jatiet: Cy. Cieglee yo ten Rbeusc cues deere tare tek site hela Miy = Wiehe aie Sime meta tie see a 493 MISCELLANEOUS Reports'ot Officers) .y2y gor hOkees sea et as bielanlak name's cles ow gas ee Oey lok Galatea ies once 495 Society. Mietings 30) 12 1) semen acids aiele said ving pce and lala tur ied area MIR aha 2 ae ee 497 Notice of New Publication: A Catalog of the Cecidomyiidae (Diptera) of the World .............. 500 VOL. 106 JULY 2004 NO. 3 Oo. fe (ISSN 0013-8797) “ bv ain eames 649 BRAILOVSKY, HARRY—tThe tribe Colpurini (Hemiptera: Heteroptera: Coreidae) of the Philippine Republic, with descriptions of four new species and a synonymical note.......... 606 BUENO-SORIA, JOAQUIN and ALICIA ROJAS-ASCENSIO—New species and distribution of the genus Marilia Miller (Trichoptera: Odontoceridae) in Mexico and Central America .... 679 CALMASUR, ONDER and HIKMET OZBEK—Heterarthrus ochropoda (Klug) (Hymenoptera: Tenthredinidae), a new record and new pest of Populus spp. (Salicaceae) in Turkey .......... 717 DEITZ, LEWIS L. and CHARLES R. BARTLETT—Publications of Thomas Kenneth Wood (CNEMEBAU DIY 62 Se RBG te ieee eae Silas) Re ASR. et Sr RTT a 0S EC Rnee oA oe AY 586 GAGNE, RAYMOND J., FRANCISCO POSADA, and ZULMA NANCY GIL—A new species of Bruggmanniella (Diptera: Cecidomyiidae) aborting young fruit of avocado, Persea americana (ent ACcAL) Sins OOO MmOlaraaa | COStaRIGA et ia. tue Sali. oleltnodat enable ssa eee eo etea acest on 547 GAIMARI, STEPHEN D., LYNN S. ADLER, and SONJA J. SCHEFFER—Plant host affiliation and redescription of Phytomyza subtenella Frost (Diptera: Agromyzidae) .................... S01 GE, SI-QIN and XING-KE YANG—Two new Chinese species of Tenomerga Neboiss (Coleoptera: Cupedidae), wit aworldicatalop of the\genus. \.)... 8920 ).5 oss. eo eee de dan deece send ah ue 631 HENRY, THOMAS J.—Raglius alboacuminatus (Goeze) and Rhyparochromus vulgaris (Schilling) (Lygaeoidea: Rhyparochromidae): Two Palearctic bugs newly discovered in North America... 513 KIMSEY, LYNN S.—Taxonomic changes and new generic synonymies in the tiphiid wasp subfamily ay ninAc. uN Meno tera, LapaiUAac)) sien ys. Geecain suas oe eee twat cate ies Se einene Ska odes 508 KIMSEY, LYNN S.—Illustrated keys to genera of the male wasps in the subfamily Thynninae AREA TIO ILE LA SUSI ANIIAG pipe ah he AR STE ial ee ay slote An ahi tis bore he ER Cela e wb cle habe bee owas oa coer 571 MATHIS, WAYNE N. and TADEUSZ ZATWARNICKI—A review of two Nearctic shore-fly species in the genus Psi/opa Fallén that were included in the genus Cres sie fae Arnaud eR EEMPUNIMCIMICN Sree. Cele SE EAE RE Be Ai a le cok Cala aie Me ete edn wee edes tel Leneue ns 639 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2004 E. E. GrisseELL, President Jon A. Lewis, Custodian Jason P. W. Hatt, President-Elect MicHaEL G. PoGuE, 7reasurer Stuart H. McKamey, Recording Secretary | JOHN W. BRown AND Davin G. Furtu, Program Chairs Ho .is B. WitLiAMs, Membership Secretary JONATHAN R. Mawps_ey, Past President Davip R. Smitu, Editor Publications Committee RAYMOND J. GAGNE Tuomas J. HENRY Wayne N. Maruis Honorary President DoNALD M. ANDERSON Honorary Members WILLIAM E. BICKLEY Kar V. KROMBEIN RONALD W. HopDGEs PAuL J. SPANGLER Manya B. STOETZEL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168. MEETINGS.— Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu- tion, on the first Thursday of each month from October to June, inclusive, at 7:00 PM. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $25.00 (U.S. currency). PROCEEDINGS.— The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub- lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send address changes to the Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomo- logical Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE PP. 731-732 OF THIS ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 29 June 2004 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 106(3), 2004, pp. 501-507 PLANT HOST AFFILIATION AND REDESCRIPTION OF PHYTOMYZA SUBTENELLA FROST (DIPTERA: AGROMYZIDAE) STEPHEN D. GAIMARI, LYNN S. ADLER, AND SONJA J. SCHEFFER (SDG) California State Collection of Arthropods, California Department of Food and Agriculture—Plant Pest Diagnostics, 3294 Meadowview Road, Sacramento, CA 95832- 1448, USA, (e-mail: sgaimari @cdfa.ca.gov); (LSA) Department of Biology, 2119 Derring Hall, Virginia Polytechnic Institute and State University, Blacksburg, VA 24061, USA, (e-mail: Isadler@vt.edu); (SJS) Systematic Entomology Laboratory, PSI, Agricultural Re- search Service, U.S. Department of Agriculture, Building 005, BARC-West, 10300 Bal- timore Ave., Beltsville, MD 20705, USA, (e-mail: sscheffe @sel.barc.usda.gov) Abstract.—The biology of Phytomyza subtenella Frost (Diptera: Agromyzidae) was observed for the first time. This species was found to be a seed feeder on the hemiparasitic Indian paintbrush, Castilleja miniata Douglas (Orobanchaceae), in Colorado. Biological observations are provided, along with a redescription of the species and illustrations of the male genitalia. Additionally, the 17 orobanch-feeding agromyzids are discussed, along with comments on several possible lineages, based on similarities of male genitalia, within the 13 orobanch-feeding Phytomyza. Key Words: host plant The agromyzid genus Phytomyza Fallén contains more than 450 described species, all of which have internally feeding, plant- parasitic larvae (Spencer and Steyskal 1986). Most species of Phytomyza are leaf- miners, but some are known to feed in other plant parts, including stems, roots, flowers, and seeds. Members of this genus attack species in nearly 30 plant families, although most individual Phytomyza species are highly restricted in diet, usually monopha- gous or feeding on few closely related plant species (Spencer 1990, Scheffer and Wieg- mann 2000). Within Phytomyza, morpho- logically similar species tend to feed on closely related plants (those in the same ge- nus or family) in a pattern suggestive of host-associated radiations Scheffer and Wiegmann 2000). This type of pattern has been discussed for a number of other (e.g., Agromyzidae, Phytomyza subtenella, Orobanchaceae, Castilleja miniata, groups of highly specialized insects (Eh- rilch and Raven 1964, Becerra 1997, Farrell 1998, Becerra and Venable 1999), although the processes giving rise to such patterns are not fully understood. A critical first step in investigating either the patterns or pro- cesses involved with insect diversification is gaining a clear understanding of the host affiliations of the species of interest. Agromyzid species attacking leaves gen- erally form an externally visible tunnel or ‘“‘leafmine” as they feed, making them con- spicuous to entomologists. In contrast, those species that feed deeply within tissues such as stems, roots, flowers, or fruits often provide no external symptom of their pres- ence. These species are not readily discov- ered without targeted collection followed by dissection of plants. Not surprisingly, our knowledge of such species and their PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. h = hypandrium, p = paramere, sd = sperm duct, ss = surstylus. 1, Epandrial complex, dorsal view. 2, Aedeagal complex, lateral view. 3, Ejyaculatory apodeme, lateral view. 4, Distiphallus, ventral view. host affiliations is fragmentary, limiting our ability to fully investigate host-use evolu- tion within Agromyzidae. Although most Phytomyza species are leafminers, a number are known to feed in stems or seeds (Spencer 1990), including one group of primarily seed feeders that has apparently radiated onto plants in the Oro- banchaceae. Currently, in North America and Europe, there are 13 Phytomyza species known from this plant family (Table 1). Al- though phylogenetic relationships among these and other Phytomyza species have not been explored, genitalic similarities suggest that there are several distinct lineages with- in this group, with unknown affinities among these lineages. Spencer (1969) first alluded to the pos-_ sibility that Phytomyza subtenella Frost is a seed (or stem) feeder, citing the elongated form of the ovipositor, and later comment- ing that it is “‘almost certainly an internal feeder”? (Spencer 1981). Here, we report the host plant for this species, having reared it from a hemiparasitic Indian paintbrush, Castilleja miniata Douglas (Orobancha- ceae). We also report observations on its natural history and provide a redescription and more detailed drawings of genitalia than were previously available in the liter- ature (only the basiphallus had been figured thus far, cf. Spencer and Steyskal 1986: figs. 1148, 1149). Phytomyza subtenella, 3 genitalia; scale bar = 0.1 mm; aa = aedeagal apodeme, bp = basiphallus, METHODS AND RESULTS Morphological terminology follows McAlpine (1981) and Spencer (1987). Voucher specimens of the flies have been labeled as such and deposited in the Cali- fornia State Collection of Arthropods (CSCA) in Sacramento, CA, and the Na- tional Museum of Natural History (USNM) in Washington, DC; voucher specimens of the plants (LSA#1, 3, 5) have been depos- ited in the Herbarium of the Rocky Moun- tain Biological Laboratory, Gothic, CO. In late summer 1996, numerous fly pu- paria were observed in fruits of C. miniata (voucher specimen LSA #3) at Emerald Lake, Colorado. Pupae were collected from these fruits in September and were stored in | oz. plastic portion cups, half at room temperature and half in a O°C freezer in an attempt to simulate overwintering condi- tions. Adult emergence by either method was low (<5% of ~200 puparia), possibly due to the change in microclimate by re- moval from the field. In July of the follow- ing year, adult flies were collected from the bracts of C. miniata. TAXONOMY Phytomyza subtenella Frost (Figs. 1—4) Phytomyza subtenella Frost 1924: 89. Type locality: “‘Mount Rainier (Washington)”’. VOLUME 106, NUMBER 3 Type: LT? in USNM (designated by Frick 1959, by assumption of holotype). Subsequent references: Frick 1952: 428 (catalog); Frick 1959: 436 (inadvertent lectotype designation, diagnosis, illustra- tions of head and wing); Frick 1965: 805 (catalog); Spencer 1969: 26 (geography), 221 (key), 276 (diagnosis, distribution, il- lustration of basiphallus); Spencer 1981: 365 (key), 425 (redescription, distribu- tion, illustration of basiphallus, comment on relationship with Phytomyza lupini Sehgal); Spencer and Steyskal 1986: 186 (key, diagnosis, distribution, illustration of basiphallus, comment on relationship with Phytomyza eumorpha Frey). Diagnosis.—Face and frons mostly yel- low; gena high; face and gena strongly re- ceding in lateral view; epistoma absent; 4 fronto-orbital setae, posterior 2 reclinate, anterior 2 convergent. Notum and scutellum silver pruinose; dorsocentral setae | +3; sin- gle row (2-6 pairs) of acrostichal setulae; forecoxa yellow; pleuron silvery-grey pru- inose, except yellow only along dorsal and posterior borders of anepisternum, dorsal edge of meron, and surrounding posterior spiracle; femora dark silver pruinose, yel- low apically; tibiae and tarsi dark. Abdom- inal tergites and sternites brown, with yel- low posterior edges. In male, paramere bi- lobed distally, with inner lobe straight, and outer lobe curving and overlapping with in- ner; basiphallus gently S-curved in lateral view with edges along middle of curve wid- ened slightly, strongly bilobed distally in ventral aspect; apex of distiphallus a single, large, membrane-bounded opening with sclerotization giving appearance of being bilobed. In female, oviscape 1.3—1.6 lon- ger than basal width, shining brown, with ring of long setae distally. Adults, d, 2.—Body length 2.3—2.8 mm. Head: Mostly yellow, except ocellar region and behind head (median occipital sclerite, occiput, and postgena) silvery-grey prui- nose, clypeus and palpus light brown, pre- mentum and lateral part of antennal scape 503 brown, and first flagellomere and arista black. Postocellar setae strong, divergent. Ocellar setae proclinate, as strong as pos- terior orbital setae, with bases same dis- tance apart as posterior ocelli. Frons 1.5— 2.0 wider than long, and 2.0—2.8 wider than eye width (when viewed from dorsal aspect); with 4 fronto-orbital setae, poste- rior 2 reclinate and subequal, anterior 2 convergent and subequal or with anterior- most one smaller; with row of small pro- clinate setulae between fronto-orbital setae and orbital margin, anteriorly becoming slightly larger and convergent. Lunule 0.09—0.12 mm high. Gena high, 0.6—0.7 x higher than eye height. Eye 1.1—1.2 high- er than wide (when viewed from lateral as- pect). One strong oral vibrissa, with several smaller setulae along oral margin. Face and gena distinctly receding in lateral view. Thorax: Notum and scutellum entirely sil- very-grey pruinose. Chaetotaxy: 1+3 dor- socentral setae (occasionally with fine su- pernumerary setae), with cluster of setulae just anterior to anteriormost seta; single row (2-6 pairs) of acrostichal setulae; 1 presu- tural intra-alar seta; | postsutural intra-alar seta; | postalar seta; 1 postpronotal seta (with several supernumerary setulae); 2 no- topleural setae; 2 pairs scutellar setae; oth- erwise bare, with few supernumerary setu- lae. Pleuron silvery-grey pruinose, except yellow along dorsal and posterior borders of anepisternum, dorsal edge of meron, and surrounding posterior spiracle. Chaetotaxy: | proepisternal seta; 3—4 anepisternal setae along posterior edge, middle 1—2 strong, outer ones fine, occasionally with 1—2 fine setae in dorsal part, anterior of this edge; 2 katepisternal setae, posterior one stronger. Legs: Forecoxa yellow, getting darker in dorsal 4%; mid- and hindcoxae dark. Femora silvery-grey pruinose, yellow distally. Tib- iae and tarsi dark brown. Wing: Length 2.4—2.8 mm. Distance between end of R, and R,,, 1.4—1.9 longer than that between R,,; and M,,,. Halter yellow. Abdomen: Tergites and sternites brown pruinose, but yellow along posterior edges. Male genita- 504 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON lia (Figs. 1-4): Epandrium rounded, with small setae in addition to covering of small hairs; surstylus fused to epandrium, with densely clustered hairs, many directed me- dially (Fig. 1). Cerci small. Hypandrium U- shaped from dorsal view, with medially di- rected process near middle, and with bi- lobed process distally on each posterior arm (Fig. 2). Paramere (postgonite of other au- thors). bilobed distally, with lobe straight, and outer lobe curving and over- lapping with inner (Fig. 2). Ejaculatory apodeme 2.0 longer than high, with distal edge fan-like (Fig. 3). Aedeagus with bas- iphallus gently S-curved in lateral view, inner with edges along middle of curve widened slightly (Fig. 2); basiphallus in ventral as- pect strongly bilobed distally (Fig. 4). Dis- tiphallus sclerotized in three parts: central sperm duct (Fig. 2), paired, elongated, an- teriorly-directed sclerites, and paired pos- teriorly-directed apical sclerites (Fig. 4); elongated sclerites confluent with apical sclerites through membrane; apex of disti- phallus a single, large, membrane-bounded opening, despite appearance of being truly terminalia: 0.36-0.46 mm wide at base, length 1.3- 1.6X width; brown, with ring of long hairs distally; with bilobed. Female Oviscape greater than basal shining light covering of silver-grey pruinosity in basal “. Type LT? (USNM), Rainier, Washlington]., above Longmire’s, 5,000 ft., Aug. 3, [19]OS [date handwritten |/ Type No. 50021 U.S.N.M handwritten|/Phytomyza subtenella Frost material. [red; number [handwritten; black submargin]. Other material examined.—USA: Wis- consin, PolkiwCo., July, Baker UPLIY; USNM); California, El Dorado Co., Echo Lake [2,250 m el.], E.I. Schlinger, 23-VII- 1955 (1 6, | 2, USNM); Colorado, Gun- nison Co., Emerald Lake, 3,150 m el., 39°00'41"N_ 107°02'32"W, reared from Cas- tilleja miniata, 5.X1.1996 (emer. HI-1997), Lynn Adleré(lud 22! ShGSECAm iiss ilak) USNM), from Castilleja hand collected Mt. - miniata, VII-1998, Lynn Adler (1 d, 3 9, CSCA RLS, 1 Fy USM): Distribution.—Canada (Alberta, British Columbia); United States (California, Col- orado, Washington, Wisconsin, Wyoming). The single paralectotype from Wisconsin is unambiguously labelled (Polk Co. Wis., July, Baker), although the presence of this species in northwestern Wisconsin (where the elevation is below 500 m), seems un- likely, as all other known specimens are from the Rocky Mountains and the Sierra Nevada, at elevations above 2,000 m. Spen- cer (1969) even comments that this species is likely to be limited to this high elevation western zone, despite the record in Wiscon- sin, which he also considered dubious. Behavioral observations.—Adult P. sub- tenella were observed on leaves and bracts of C. miniata in late July 1998 at Emerald Lake, Colorado, where most were in flower. Pairs of adults, or occasionally three adults, were observed in what appeared to be mat- ing positions. Approximately two weeks later, adult flies were no longer observed, but fly larvae were common inside the locules of matur- ing fruits. Up to three larvae were found per single fruit. Because there was no ob- vious damage to these fruits (i.e., no evi- dence of larval entry holes), it seems likely that Ovipositition was directly into fruits. Larvae consumed seeds while leaving be- hind the netlike outer seed coat, causing ex- tensive damage (Adler 2002). By early September, C. miniata fruits had matured and begun dehiscing, and the flies had pupariated. These puparia were found both inside and outside fruits. In the latter case, exit holes were evident in the capsule wall and at times puparia were found pro- truding through these holes. The duration of the puparial stage in the field is not known; field collected puparia emerged after sev- eral months in the laboratory. DISCUSSION The group of Phytomyza presented in Ta- ble | all attack Orobanchaceae, parasitic VOLUME 106, NUMBER 3 505 Table 1. Phytomyza species known to feed on plants in the family Orobanchaceae, arranged by host genus; * = new record; ' = probable feeding site. All host genera are in the hemiparasitic tribe Rhinantheae, except for Orobanche which is in the holoparasitic tribe Orobancheae. Species Distribution Host Genus Feeding Site P. subtenella Frost* Nearctic P. trivittata Frost Nearctic P. rostrata Hering Palaearctic P. affinis Fallen Palaearctic P. flavofemorata Strobl Palaearctic P. isais Hering Palaearctic P. orobanchia Kaltenbach Palaearctic P. diversicornis Hendel Palaearctic P. pedicularicaulis Spencer Nearctic P. pedicularidis Spencer Nearctic P. pedicularifolii Hering Palaearctic 4 o Holarctic Holarctic . tenella Meigen . varipes Macquart Castilleja seeds Cordylanthus seeds! Euphrasia, Melampyrum, stems Odontites, Rhinanthus Euphrasia seeds Melampyrum seeds Odontites seeds Orobanche seeds, stems Pedicularis stems Pedicularis stems Pedicularis seeds Pedicularis leaves Pedicularis seeds Rhinanthus seeds plants that obtain water and nutrients from other plants via root connections. Several species of this group have unknown affini- ties, but genitalic similarities indicate sev- eral distinct lineages. For example, one seed-feeding species, P. flavofemorata Strobl] attacking Melampyrum L., appears, by genitalic similarity, to belong to a group with three leaf-feeding species attacking non-orobanchs, including P digitalis. Hering attacking Digitalis L., and P. crassiseta Zetterstedt and P. veronicicola Hering at- tacking Veronica L. (Plantaginaceae, Digi- talideae). The species P. trivittata Frost, at- tacking Cordylanthus Nutt. ex Benth., ap- pears very closely related by nearly identi- cal male genitalia to P. chelonei Spencer, a species attacking Chelone L. (Plantagina- ceae, Cheloneae). There appear to be at least two other distinct groups based on genitalic similarities. The first includes two stem feeders (P. diversicornis Hendel and P. pedicularicaulis Spencer, both attacking Pedicularis L.) and two seed feeders (P. af- finis Fallén and P. tenella Meigen, the for- mer attacking Euphrasia L. and the latter Pedicularis). The second includes the seed feeder P. subtenella (attacking Castilleja Mutis ex L.f.) and the seed/stem feeding P. orobanchia Kaltenbach (attacking species of the holoparasitic genus Orobanche L.), but also includes several additional species whose host affiliations remain undiscovered (e.g., P. eumorpha Frey and P. orindensis Spencer), or attack species of the non-oro- banch genus Lupinus L. (Fabaceae) (P. lu- pini Sehgal). Interestingly, P. orobanchia ot this latter group has been shown to be of potential importance in controlling Oroban- che species parasitizing tomatoes in Ethio- pia (Elzein et al. 1999). Two Phytomyza species, P. lupini and P. lupinivora Sehgal, feed on hosts of the ge- nus Lupinus, which is in a family quite dis- tinct from Orobanchaceae. The latter spe- cies, known only from the female holotype, is a leafminer, while the former species is an internal stem or flower-head feeder, even pupariating internally. Interestingly, Spen- cer (1981) speculated on a relationship be- tween P. lupini and P. subtenella, based on unstated similarities of the male genitalia, being distinguished by different states of the male epistoma above the mouth margin (which is lacking in P. subtenella) and the color of the forecoxa. Comparing genitalia, the aedeagus is very similar in shape and size from the lateral aspect, and the disti- phallus is very similar from ventral aspect; the same holds true for the genitalia of P. 506 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON orobanchia, P. eumorpha and P. orinden- sis, and in fact, the genitalia of P. eumorpha appear most similar to P. subtenella. Castilleja species, and other Orobancha- ceae, are known to parasitize species of Lu- pinus and obtain alkaloids, secondary com- pounds implicated in resistance to herbi- vores, as well as nitrogen resources from this association (Stermitz and Harris 1987, Stermitz et al. 1989, Arslanian et al. 1990, Boros et al. 1991, Stermitz and Pomeroy 1992). Because of the putatitive close re- lationships among some of these taxa, one can speculate on the possibility of a host switch from feeding within stems and flow- er-heads of lupine to feeding within the par- asitic orobanchs attacking them, or vice- versa. By obtaining compounds from hosts, parasitic plants may expose herbivores to novel secondary compounds normally found in the unrelated host plant species, allowing herbivores to pre-adapt to novel hosts. However, without phylogenetic hy- potheses for relationships among Phytomy- za species and detailed information about host ranges, it is not possible to do more than speculate about how host-plant rela- tionships for Phytomyza species evolved. Few other species of agromyzids attack Orobanchaceae besides the 13 Phytomyza species listed in Table 1. Aside from the single species discussed herein, only Chro- matomyia castillejae (Spencer) and its sub- species nordica Spencer are known to use Castilleja species as hosts (leafminers on Castilleja fissifolia L.f. in Venezuela and C. latifolia Hook. & Arn. in California, re- spectively) (Spencer 1973, Spencer 1981). Outside of Castilleja feeders, the only other orobanch feeders are Ophiomyia strigalis Spencer attacking the lower stem and root of holoparasitic Striga Lour., and the po- lyphagous Chromatomyia horticola Gou- reau attacking Melampyrum and Rhinan- thus L. ACKNOWLEDGMENTS Flies were collected and reared at the Rocky Mountain Biological Laboratory by LSA, who thanks Judah Dinnel for help with field work. This work was funded by National Science Foundation Dissertation Improvement Grant DEB98-00885 to LSA. We also thank Wayne N. Mathis (USNM) for providing access to and help with the lectotype of Phytomyza_ subtenella, Eric Fisher (CSCA) for reviewing the manu- script, and Fred Hrusa (California Depart- ment of Food & Agriculture) for help with understanding botanical issues surrounding the circumscription of Orobanchaceae. LITERATURE CITED Adler, L. S. 2002. Host effects on herbivory and pol- lination in a hemiparasitic plant. Ecology 83: 2700-2710. Arslanian, R. L., G. H. Harris, and E R. Stermitz. 1990. New quinolizidine alkaloids from Lupinus argenteus and its hosted root parasite Castilleja sulphurea: Stereochemistry and conformation of some naturally occurring cyclic carbinolamides. Journal of Organic Chemistry 55: 1204—1210. Becerra, J. X. 1997. Insects on plants: Macroevolu- tionary chemical trends in host use. Science 276(5310): 253-256. Becerra, J. X. and D. L. Venable. 1999. Macroevolu- tion of insect-plant associations: The relevance of host biogeography to host affiliation. Proceedings of the National Academy of Sciences of the Unit- ed States of America 96(22): 12626-12631. Boros, C. A., D. R. Marshall, C. R. Caterino, and F R. Stermitz. 1991. Iridoid and phenylpropanoid glycosides from Orthocarpus spp.: Alkaloid con- tent as a consequence of parasitism on Lupinus. Journal of Natural Products (Lloydia) 54: 506— Sill 3). Ehrilch, P. R., and P. H. Raven. 1964. Butterflies and plants: A study in coevolution. Evolution 18: 586— 608. Elzein, A.E.M., J. Kroschel, A. Admasu and M. Fe- tene. 1999. Preliminary evaluation of Phytomyza orobanchia (Diptera: Agromyzidae) as a control- ler of Orobanche spp. in Ethiopia. Sinet 22: 271— 282. Farrell, B. D. 1998. “‘Inordinate fondness” explained: Why are there so many beetles? Science 281(5376): S55—559: Frick, K. E. 1952. A generic revision of the family Agromyzidae (Diptera) with a catalogue of New World species. University of California Publica- tions in Entomology 8: 339—452. . 1959. Synopsis of the species of agromyzid leaf miners described from North America (Dip- tera). Proceedings of the United States National Museum 108: 347—465. VOLUME 106, NUMBER 3 . 1965. Family Agromyzidae, pp. 794—805. In Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson, eds. A Catalog of the Diptera of America North of Mexico. USDA Ag- riculture Handbook No. 276, 1696 pp. Frost, S. W. 1924. A study of the leaf-mining Diptera of North America. Cornell University Agricultural Experiment Station Memoirs 78: 228 pp. McAlpine, J. EF 1981. Morphology and Terminology— Adults (chapter 2), pp. 9-63. In McAlpine, J. E, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, coords., Manual of Nearctic Diptera, Vol. 1. Research Branch, Agri- culture Canada. Monograph No. 27: 1—674. Scheffer, S. J., and B. M. Wiegmann. 2000. Molecular phylogenetics of the holly leafminers (Diptera: Agromyzidae: Phytomyza): species limits, speci- ation, and dietary specialization. Molecular Phy- logenetics and Evolution 17: 244—255. Spencer, K. A. 1969. The Agromyzidae of Canada and Alaska. Memoirs of the Entomological Society of Canada 64: 311 pp. . 1973. The Agromyzidae of Venezuela. Revis- ta de la Facultad de Agronomia (Maracaibo) 7: 5— 108. . 1981. A revisionary study of the leaf-mining flies (Agromyzidae) of California. University of 507 California Division of Agricultural Sciences, Spe- cial Publication 3273: 489 pp. . 1987. Agromyzidae (chapter 73), pp 869-879. In McAlpine, J. F, ed., Manual of Nearctic Dip- tera, Vol. 2. Research Branch, Agriculture Canada. Monograph No. 28: 675-1332. . 1990. Host Specialization in the World Agro- myzidae (Diptera). Kluwer Academic Publishers, Dodrecht, 444 pp. Spencer, K. A. and G. C. Steyskal. 1986. Manual of the Agromyzidae (Diptera) of the United States. USDA Agriculture Handbook No. 638, 478 pp. Stermitz, E R., G. N. Belofsky, D. Ng, and M. C. Singer. 1989. Quinolizidine alkaloids obtained by Pedicularis semibarbata (Scrophulariaceae) from Lupinus fulcratus (Leguminosae) fail to influence the specialist herbivore Euphydryas editha (Lepi- doptera). Journal of Chemical Ecology 15: 2521— 2530. Stermitz, E R. and G. H. Harris. 1987. Transfer of pyrrolizidine and quinolizidine alkaloids to Cas- tilleja (Scrophulariaceae) hemiparasites from composite and legume host plants. Journal of Chemical Ecology 13: 1917-1925. Stermitz, E R. and M. Pomeroy. 1992. Iridoid glyco- sides from Castilleja purpurea and C. indivisa, and quinolizidine alkaloid transfer from Lupinus texensis to C. indivisa via root parasitism. Bio- chemical Systematics and Ecology 20: 473-475. PROC. ENTOMOL. SOC. WASH. 106(3), 2004, pp. 508-512 TAXONOMIC CHANGES AND NEW GENERIC SYNONYMIES IN THE TIPHIID WASP SUBFAMILY THYNNINAE (HYMENOPTERA: TIPHITDAE) LYNN S. KIMSEY Bohart Museum of Entomology, University of California, One Shields Ave., Davis, CA 95616, USA (e-mail: Iskimsey @ucdavis.edu) Abstract.—A number of taxonomic changes are made in the generic status and species composition of six thynnine genera, four from Australia and two from South America. New generic synonymies are given, with Pogonothynnus Turner made a junior synonym of Zaspilothynnus Turner, Aspidothynnus Turner a junior synonym of 7mesothynnus Turner and Glottynoides Kimsey a junior synonym of Scofaena Turner. Consequently species assignments have to be changed. New combinations for Aspidothynnus are col- laris (Guérin de Meneville), dispersus (Turner), humilis (Erichson), ingrediens (Turner), iridipennis (Smith), platycephalus (Turner), truncatus (Smith), strangulatus (Smith), and zelebori (Saussure); for Zaspilothynnus fenestratus (Smith), fulvohirtus (Turner), morosus (Smith), and vestitus (Smith), and for Scotaena brunnea (Fox), clypearis (Duran), duckei (Smith), fastuosus (Smith), genisei (Kimsey), rosenbergi (Turner), and vigilii (Bréthes). Key Words: ‘Tiphiidae, Thynninae, Pogonothynnus, Zaspilothynnus, Aspidothynnus, Tmesothynnus, Scotaena, Glottynoides Phylogenetic analyses of the genera of Thynninae currently underway have result- ed in the need for some additional generic synonymy. The genera discussed below all end up as sister group pairs in these anal- yses, without diagnostic apomorphies to separate them. This prompted their reex- amination, which led to the synonymies proposed below. Three genera are synony- mized. Two, generic pairs are in the tribe Thynnini from Australia. The third pair is in the Scotaenini from South America. A number of species assignments need to be changed, as given below, as a result of these synonymies and from the examination of primary types in The Natural History Mu- seum, London, and Humboldt Museum, Berlin. Species where the primary types have been examined are indicated by an as- terisk (*). When types have not been seen, the generic placement given by Given (1954) is followed. ASPIDOTHYNNUS TURNER Turner (1910b) described two new gen- era, Aspidothynnus and Tmesothynnus, largely based on male characteristics. At the time, females were known only for 7me- sothynnus. He distinguished the two genera by the structure of the male clypeus and hypopygium. However, after examining the types of the genera and a much broader se- lection of species than Turner had available at the time it is clear that characteristics used to distinguish these genera intergrade between the two extreme forms represented by the types of the genera, Aspidothynnus combustus Turner and Tmesothynnus zele- bori Saussure. The male hypopygium in these species ranges from apically tridentate with parallel sides to having an apicomedial spine or tooth and rounded sides (Figs. 4— 7). Females associated with species placed in both groups are indistinguishable on the VOLUME 106, NUMBER 3 generic level. Males of both genera share several apomorphic features—the subanten- nal sclerite and frons share a well-devel- oped longitudinal medial carina, and in the tongue the stipes is nearly asetose, except for a row of well-separated, short setae along the inner margin, and the prementum is slender, usually elongate, and asetose (Fig. 1). The hypostomal plate on either of the oral fossa is considerably narrower than the oral fossa and is about as broad as or narrower than the prementum (Fig. 1). Oth- er male features that distinguish these gen- era from others closely related include the apical flagellomeres cylindrical (not lobu- late), with two tyloids, the presence of a small red spot on the vertex between the hindocellus and eye margin, and epipygium with a large, transverse carina or swelling before a thin apical rim, which varies from linear, to trilobate to strongly swollen and bilobate (Figs. 2—3). Females have the py- gidium margined laterally by a carina and subtended by a long brush of setae on me- tasomal sternum V, and, most significantly, sternum VI is apically strongly bilobate. None of these characters were found to clearly separate groups of species. Phylo- genetic analyses of Aspidothynnus and Tmesothynnus species resulted in a single, pectinate clade, with many possible trees and no Bootstrap support for any of the branches. Therefore, Aspidothynnus is cho- sen as the senior name and 7mesothynnus the junior synonym (new synonymy). Species now placed in this revised As- pidothynnus are: Lophocheilus collaris Guérin de Meneville 1842, new combina- tion; Aelurus combustus Smith 1859*; Thynnus dispersus Turner 1908*, new com- bination; Aspidothynnus fossulatus Turner 1915*; Thynnus humilis Erichson 1842, new combination; Aspidothynnus ingre- diens Turner 1916*, new combination; Thynnus iridipennis Smith 1859*, new combination; 7mesothynnus platycephalus Turner 1910c*, new combination; 7/ynnus polybioides Turner 1908*; Aspidothynnus rostratus Turner 1908*; Thynnus truncatus 509 Smith 1859*, new combination; Thynnus strangulatus Smith 1879*, new combina- tion, and Thynnus zelebori Saussure 1867, new combination. ZASPILOTHYNNUS ASHMEAD Turner (1910b) described the genus Po- gonothynnus on the page preceding his re- description of Zaspilothynnus Ashmead (1903). The two descriptions are nearly identical except for the shape of the hypo- pygium. Males in both genera have a strongly elevated medial platform on the epipygium, small tooth or spine on meta- somal sternum VI, the same configuration of the antennal lobes and associated carina, ranging from U-shaped to V-shaped, and most species have a well-developed stipal fringe (Fig. 8). The male genital capsule structure is also the same in the types of both genera (Figs. 17, 19), with the gono- coxa dorsally truncate, cuspis with large, sometimes elongate, digitate lobe on the in- ner margin (Figs. 16, 18), and large earlike digitus that folds down dorsally during cop- ulation. Females in these genera have the same densely ridged metasomal terga I and II and snoutlike epipygium, which is nar- rowed basally with a flared, flattened pos- terior plate (Figs. 13—15). No consistent dif- ferences could be found between the two groups. As with Aspidothynnus, phyloge- netic analysis of the combined species of both Pogonothynnus and Zaspilothynnus re- sulted in numerous poorly supported trees, with no support for two discrete clades that might represent these genera. Therefore, Pogonothynnus Turner 1910b becomes a junior synonym of Zaspilothynnus Ash- mead 1903, new synonymy. Included species in the combined Zaspi- lothynnus are: Thynnus andreanus Turner 1908; Thynnus atrocior Turner 1909a; Thynnus biroi Turner 1910d; Thynnus cam- panularis Smith 1868; Thynnus carbonar- ius Smith 1859; Zaspilothynnus cheesman- ae Turner 1940; Zaspilothynnus clelandi Turner 1910c; Thynnus crudelis Turner 1908; Zaspilothynnus cyaneiventris Rohwer 510 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2. fossulatus 3. unidentified sp 4. unidentified sp. J V J 5. zelebori 6. fossulatus 7. combustus 9. dilatatus hd \V 10. 7. interruptus 12. gilesi 11. fulvohirtus 8. morosus | ES: enestr 14. carbonarius interruptus FOLSIELS 15. birot Figs. 1-19. Aspidothynnus species. 1, Posterior view of male head. 2-3, Posterior view of male epipygium. 4-7, Dorsal view of male hypopygium. 8-19, Zaspilothynnus species. 8, Lateral view of male head, with antenna removed. 9-12, Ventral view of male hypopygium. 14—15, Posterior view of female apical metasomal segments. 16, 18, Ventral view of volsella. 17, 19, Dorsal (left) and ventral (right) views of male genital capsule. VOLUME 106, NUMBER 3 1925; Thynnus dilatatus Smith 1859; Thyn- nus excavatus Turner 1908; Thynnus fenes- tratus Smith 1859, new combination; Po- gonothynnus fulvohirtus Turner 1915, new combination; Zaspilothynnus gilesi Turner 1910c; Zaspilothynnus hackeri Turner 1912; Thynnus interruptus Westwood 1845; Zaspilothynnus lasius Montet 1922; Zaspi- lothynnus lignatus Turner 1910c; Zaspilo- thynnus maturus Turner 1909b; Thynnus morosus Smith 1879, new combination; Thynnus multistrigatus Turner 1909a; Zas- pilothynnus neglectus Turner 1910c; Thyn- noides nigripes Guérin de Méneville 1842; Thynnus novarae Saussure 1868; Thynnus ochrocephalus Smith 1868; Thynnus picti- collis Turner 1908; Thynnus pseustes Turn- er 1908; Zaspilothynnus radialis Turner 1910c; Zaspilothynnus rhynchioides Turner 1913; Zaspilothynnus rubropictus Turner 1937; Thynnus rufoluteus Turner 1910d; Zaspilothynnus rugicollis Turner 1915; Thynnus seductor Smith 1868; Thynnus sic- cus Turner 1908; Thynnus simplex Smith 1879; Zaspilothynnus striatifrons Turner 1917; Zaspilothynnus trilobatus Turner 1910c, Zaspilothynnus unipunctatus Turner 1915; Thynnus vernalis Turner 1908; Thyn- nus vestitus Smith 1859, new combination. SCOTAENA KLUG The genus Glottynoides Kimsey was de- scribed for the species genisei Kimsey, which has an unusually modified tongue. In the original description, a strong relation- ship with the genera Scotaena and Rostryn- nus was mentioned. Further examination of male and female characteristics of Glottyn- oides and Scotaena indicates that the only distinctions between these two genera are the species level characteristics of the male tongue seen in genisei. Therefore, Glottyn- oides Kimsey (in Genise and Kimsey 1991) is a junior synonym of Scotaena Klug 1810, new synonymy. Species included in the genus Scotaena include: Elaphroptera brunnea Fox 1898*, new combination; Ornepetes clypearis Duran-Moya 1941*, new combination; 511 Thynnus decora Smith 1859*; Thynnus duckei Smith 1859, new combination; Sco- taena duckei Turner 1909*; Thynnus fas- Smith 1879*, new combination; Thynnus flavovariegatus Smith 1879*; Glottynoides genisei Kimsey 1991*, new combination; Spilothynnus horni Turner 1927*; Elaphroptera pallida Fox 1898*; Scotaena polistoides Turner 1910a*; Thyn- nus pubescens Klug 1842*; Scotaena _ ro- senbergi Turner 1910a*, new combination; Thynnus trifasciata Klug 1810*; Scotaena vetusta Turner 1909b*; Elaphroptera vigilii Bréthes 1910*, new combination. TUOSUS LITERATURE CITED Ashmead, W. H. 1903. Classification of the fossorial, predaceous and parasitic wasps, of the superfam- ily Vespoidea. Canadian Entomologist 35: 95— 107. Brethes, J. 1910. Himenopteros Argentinos. Anales del Museo Nacional de Buenos Aires 20: 205-316. Duran-Moya, L. 1941. Die Thynniden von Chile. Ar- chiv fiir Naturgeschichte, Berlin 10: 71-176. Erickson, W. F 1842. Beitrag zur Insecten-Fauna von Vandiemensland mit besonderer Berucksichtigung die Geographischen Verbreitung der Insecten. Ar- chiv fiir Naturgeschichte, Berlin 8: 83—287. Fox, W. J. 1898. Contributions to a knowledge of the Hymenoptera of Brazil, No. 4. Thynninae and ad- ditions. Proceedings of the Academy of Natural Sciences, Philadelphia 50: 72-75. Genise, J. and L. S. Kimsey. 1991. New genera of South American Thynninae. Psyche 98: 57—69. Given, B. B. 1954. A catalogue of the Thynninae of Australia and adjacent areas. Bulletin of the New Zealand Department of Scientific and Industrial Research (109): 1-89. Guérin de Méneville, FE E. Thynnides. Magasin de Zoologie 4: 1—15. Klug, J. C. E 1810. Einige neue Piezatengattungen. Magazin Gesellschaft Naturforschender Freunde Berlin 4: 31—43. . 1842a. Ueber die Insectenfamilie Heterogyna Latr. und die Gattung 7Thynnus F insbesondere. Physikalische und Mathematische Abhandlungen der KGniglichen Akademie der Wissenschaften, Berlin, pp. 1—44. Montet, G. 1922. Thynnides nouveaux de Muséum d’ Histoire de Genéve. Revue Suisse de Zoologie, Geneva 29: 177-226. Rohwer, S. A. 1925. New thynnid wasps from the Ori- ental and Australian Regions. Philippine Journal of Science 26: 415—420. Saussure, H. L. E 1867. Hymenoptera. Familien der 1842. Matériaux sur les Vespiden, Sphegiden, Pompiliden, Crabroniden und Heterogynen. Reise Novaria-Expedition, Zoologischer Theil 2: 1-156. . 1868. Reise der Osterreich Fregatte Novara, Zoologischer, Hymenoptera, pp. 108—134. Smith, E 1859. Hymenoptera. Catalogue of the Hy- menopterous insects in the collection of the Brit- ish Museum, part 7, Dorylidae and Thynnidae, pp. 10-69. . 1868. Descriptions of aculeate Hymenoptera from Australia. Transactions of the Entomological Society of London 1868: 231—237. . 1879. Descriptions of new species of Hyme- noptera in the British Museum, London, 240 pp. Turner, R. E. 1908. A revision of the Thynnidae of Australia. Pt. Il. Proceedings of the Linnaean So- ciety of New South Wales, Sydney 33: 70—208 (June), 209—256 (August). . 1909a. Remarks on some new or little known species of Thynnidae. Annals and Magazine of Natural History (8) 3: 131—146. . 1909b. Notes on fossorial Hymenoptera. I. Annals and Magazine of Natural History (8) 3: 338-343. . 1910a. On the Thynnidae and Scoliidae col- lected in Paraguay. Zoologische Jahrbiicher, Ab- teilung fiir Systematik 29: 179-227. . 1910b. Hymenoptera, Family Thynnidae. /n Wytsman, P., ed. Genera Insectorum, 105. Brux- elles, 62 pp. . 1910c. Additions to our knowledge of the fos- sorial wasps of Australia. Proceedings of the Zoo- logical Society of London 1910: 253-359 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 1910d. New species of Thynnidae from the Australian and Austro-Malayan Regions. Annales Historico-Naturales Musei Nationalis Hungarici 8: 107-124. . 1912. Notes on fossorial Hymenoptera. XI. On some new Australian and Austro-Malayan Thynnidae. Annals and Magazine of Natural His- tory (8) 10: 533-546. . 1913. New fossorial Hymenoptera from Aus- tralia and Tasmania. Proceedings of the Linnaean Society of New South Wales 38: 608-616. 1915. Descriptions of new fossorial wasps from Australia. Proceedings of the Zoological So- ciety of London 1915: 41—69. . 1916. Notes on fossorial Hymenoptera. XIX. On new species from Australia. Annals and Mag- azine of Natural History (8) 17: 116-136. . 1917. New species of Hymenoptera in the British Museum. Transactions of the Entomolog- ical Society of London 1917: 58—60. . 1927. On a new thynnid wasp from Paraguay. Entomologische Mitteilungen 16: 449. . 1937. Notes on fossorial Hymenoptera. XLVI. On new Australian Thynnidae. Annals and Mag- azine of Natural History (10) 19: 144-150. 1940. Notes on fossorial Hymenoptera. XLVIUI. New Guinea Thynnidae collected by Miss Cheesman. Annals and Magazine of Natural History (11) 5: 91—96. Westwood, J. O. 1845 (1844). Plates 76,77. A decade of Australian thynnideous insects. Arcana Ento- mologica 2(July 1845): 113-124. PROC. ENTOMOL. SOC. WASH. 106(3), 2004, pp. 513-522 RAGLIUS ALBOACUMINATUS (GOEZE) AND RHYPAROCHROMUS VULGARIS (SCHILLING) (LYGAEOIDEA: RHYPAROCHROMIDAE): TWO PALEARCTIC BUGS NEWLY DISCOVERED IN NORTH AMERICA THOMAS J. HENRY Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, % National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC-0168, Washington, DC 20013-7012, U.S.A. (e-mail: thenry @sel. barc.usda.gov) Abstract.—Two invasive Palearctic rhyparochromids (Lygaeoidea: Rhyparochromidae) are recorded for the first time in North America. Raglius alboacuminatus Goeze is reported from California, Oregon, and northern Utah, and Rhyparochromus vulgaris (Schilling), from Oregon and Washington. Although not considered major agricultural pests, these invasive bugs have become serious nuisance pests, invading homes and commercial es- tablishments in large numbers. Provided for each are a summary of the literature, notes on the biology and habitat, diagnosis, description, and photographs of the adult to help separate these nonindigenous bugs from other North American Rhyparochromidae. In addition, Xanthochilus saturnius (Rossi), a Palearctic species known in North America only from California, is reported for the first time from Oregon. Key Words: Heteroptera, Lygaeoidea, Rhyparochromidae, Raglius alboacuminatus, Rhy- parochromus vulgaris, North America, new records, California, Oregon, Utah, Washington, invasive Both invasive Palearctic seed bugs re- ported in this paper, Raglius alboacumina- tus (Goeze) and Rhyparochromus vulgaris (Schilling), were discovered after they at- tracted attention as serious nuisance pests by homeowners and various local business- es. For Raglius alboacuminatus, these events occurred in California and Utah; for Rhyparochromus vulgaris, similar situa- tions were documented in Oregon and Washington. While these nonindigenous in- sects probably will not become important agricultural pests, they may profoundly in- fluence our environment in other ways. Both species already have affected local economies, causing anxiety among hom- eowners and creating costly control expens- es for commercial establishments. The first specimens of Raglius alboacu- minatus were submitted to the Systematic Entomology Laboratory, ARS, USDA, Washington, D.C., in October 1999 by Alan Roe at Utah State University, Logan. Jay Karen (Utah State University), stated that specimens were found invading a local li- brary and causing concern about possible damage to library books. Shortly thereafter, other reports of large numbers of this bug in the region invading homes and schools attracted the attention of the local televi- sion, radio, and newspaper media. One lo- cal Utah newspaper (The Herald Journal) published articles (by M. R. Weibel) with headlines, such as “‘New Library is Fight- ing Bugs” (October 11, 1999) and ‘North Logan Library Shakes Heebie-Jeebies”’ 514 (October 15, 1999). In 2000, very large populations of these bugs invaded several buildings of a prominent trade school at Ogden, Utah. A visit to Logan, Ogden, and surrounding areas in 2001 indicated that the populations of R. alboacuminatus apparent- ly had collapsed, making it difficult to col- lect even a few dozen bugs. In August 2002, however, Dr. Karen reported to me that large populations had again developed in the Ogden area in numbers similar to the 2000 outbreak. I also received a specimen of R. alboacuminatus from Alan Hardy (California Department of Food & Agricul- ture, Sacramento) in October 2002 from San Jose, California, that was collected in August 2002 from a population of thou- sands of specimens invading a home from surrounding old fields. One report from the latter site indicated that one wall of the home was so heavily covered that it ap- peared ‘“‘alive with bugs” (A. Hardy, per- sonal communication). More recently, spec- imens from Benton County, Oregon, were discovered by John D. Lattin (Oregon State University, Corvallis). In November 2001, I received an e-mail and digital image of another rhyparochrom- id from Eric LaGasa (Washington Depart- ment of Agriculture, Olympia). The images were tentatively identified as representing Rhyparochromus vulgaris and later con- firmed by examining a series of specimens collected at Orting, Washington. These bugs were present in large numbers in and around a home, many of which were found on sidewalks, siding, and trapped in win- dow sills. A follow-up survey of the area revealed smaller numbers of specimens over a wider area around the Orting area, including a few specimens in a local nurs- ery. Shortly after the Washington discovery, communication with James LaBonte and Kathleen Johnson (Oregon Department of Agriculture, Salem) and John D. Lattin re- vealed additional specimens collected in Oregon, indicating that this rhyparochromid is well established in the Pacific Northwest. In this paper, | summarize the known in- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON formation on the two Palearctic bugs, Rag- lius alboacuminatus recently discovered in California, Oregon, and Utah, and Rhypar- ochromus vulgaris recently discovered in Oregon and Washington, and provide for each a diagnosis, description, and illustra- tions of the adult. In addition, Xanthochilus saturnius (Rossi), a Palearctic species known in North America only from Cali- fornia, is reported for the first time from Oregon. The following acronyms are used for de- positories cited in this paper: ODA (Oregon Department of Agriculture, Salem); OSU (Oregon State University, Corvallis); USU (Utah State University, Logan); USNM ({United States] National Museum of Nat- ural History, Smithsonian Institution, Washington, D.C.); and WSDA (Washing- ton State Department of Agriculture, Olym- pia). Raglius alboacuminatus (Goeze) (Figs. 1—3) Cimex alboacuminatus Goeze 1778: 266 (original description). Rhyparochromus alboacuminatus: Slater 1964: 1310 (catalog); Péricart 1998: 271 (description, distribution, biological notes, illustrations). Raglius alboacuminatus: Péricart 2001: 208 (catalog). See Slater (1964) and Péricart (1998, 2001) for extensive bibliogra- phies. Diagnosis.—This medium-sized species (Figs. 1, 2) is best recognized by the overall fuscous to black coloration, with a pale or whitish posterior pronotal lobe, a pale an- terior half of the corium (with alternating linear brown and pale areas producing a striped appearance), and a white spot at the apex of corium and the apex of hemelytral membrane. Description—Male (n = 10): Length 4.64-5.28 mm, width 1.53—1.72 mm. Head: Width 0.98—1.03 mm, vertex 0.63— 0.65 mm. Rostrum: Length 2.13—2.30 mm, extending to bases of mesocoxae. Antenna: VOLUME 106, NUMBER 3 Nn nr Figs. 1-2. Segment I, length 0.36—0.43 mm; II, 0.87— 1.00 mm; III, 0.85—0.93 mm; IV, 1.03—1.08 mm. Pronotum: Length 0.95—1.13> mm, basal width 1.43—1.48 mm. Female (n = 10): Length 4.96—6.56 mm, width 1.64—2.08 mm. Head: Width 0.98— 1.13 mm, vertex 0.63—0.75 mm. Rostrum: Length 2.17—2.50 mm, extending to meso- coxae Antenna: Segment I, length 0.38— 0.55 mm; II, 0.85—1.15 mm; HI, 0.78—1.05 mm; IV, 0.93-0.115 mm. Pronotum: Length 0.98—1.30 mm, basal width 1.48— 1.88 mm. Small to medium sized, slender, elon- gate-oval, generally dark brown to black with pale or white areas on the pronotum, hemelytra, and hemelytral membrane (Fig. 1). Head shiny black. Labium fuscous. An- tennal segment I black, apex narrowly pale brown; segment II pale yellowish brown, narrowly fuscous at base; segments III and Adult female of Raglius alboacuminatus. 1, Dorsal aspect. 2, Lateral aspect. IV uniformly black. Pronotum rectangular, basal margin deeply emarginate, lateral margin narrowly explanate; anterior lobe uniformly shiny black, impunctate, gently and evenly swollen; narrower posterior lobe pale or whitish, basal angles black, darkly punctate across base and through middle, punctures fading laterally. Scutellum equi- lateral, finely punctate, dull black with nar- row apex yellowish brown. Hemelytron punctate, most densely along veins; basal half largely pale or whitish; inner half of clavus dark brown, outer half white; corium pale or whitish, narrowly brown along cla- val suture and veins giving a striped ap- pearance; apical half of corium largely fus- cous to black with a large white blotch at apex bordering membrane; membrane black with a round to triangular white spot at middle of apex. Ventral surface black ex- cept for white posterior pleural area of 516 pronotum, thoracic metanotum, and pro-, meso- and meta-acetabulae. Legs: Coxae black; trochanters brown; swollen profemur black, yellowish brown at base and apex, with one large and two to three smaller spines ventrally on apical third; meso- and metafemora black on apical third to half, yellowish brown basally; tibiae brown to yellowish brown, fuscous at bases; tarsi and claws brown. Habitat.—Raglius alboacuminatus colo- nizes open clear areas and borders of woods, and old fields where tall vegetation grows, and seems to prefer sandy, calcare- ous soils (Wagner 1961, Péricart 1998). In California and Utah, large populations de- veloped in old or fallow fields with mixed vegetation. Late in the season (August to November), adults and late-instar nymphs migrated to irrigated lawns and congregated in and around buildings in enormous num- bers, resulting in media attention and con- cern to businesses and homeowners. Biology.—According to Péricart (1998), Raglius alboacuminatus Overwinters as adults under the bark of trees, often in large groups, sometimes several meters above ground. Its feeds on fallen seeds and, very often, climbs on vegetation to feed on de- veloping seeds. In the Ukraine, it is fre- quently found near Stachys spp. and other Lamiaceae, such as Marrubium sp. and Bal- lota sp., or Scrophulariaceae, such as Ver- bascum lychnitis (Putshkov 1969). Over- wintering adults emerge in early spring as the weather warms. They deposit one or two eggs, one at a time, in ground litter or on the soil in late April or early May and continue until mid August. Populations can have a second generation, with nymphs de- veloping until September. In the southern Ukraine, a third generation can develop. In England, mating begins in early May and first-generation adults appear by late July to begin a second generation. Eggs are depos- ited singly in leafy or woody litter (South- wood and Leston 1959), Distribution.—In the Old World, R. al- boacuminatus is widespread from Great PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Britain, throughout much of Europe to the Middle East; into northern Africa, including Algeria, Morocco, Tunisia, and the Canary Islands; and western parts of Asia, includ- ing Armenia, Iran, Kazakhstan, Russia, Tur- key, Tadzhikistan, and Uzbekistan (Péricart 2001). This species is recorded for the first time in the United States (Fig. 3) from one coun- ty in California (Santa Clara), three coun- ties in Oregon (Benton, Multnomah, and Wasco), and three counties in Utah (Box Elder, Cache, and Weber). New United States records —CALIFOR- NIA: 1 2, Santa Clara Co., San Jose, 5356 Colony: Field’ Drive, 21 August..2002, 1P Hamilton (USNM); 3 66, 16 2, Santa Clara Co., San Jose, 5356 Colony Field Drive, 121.84°N, 37.26°W, 24 August 2002, M. Nachand (USNM). OREGON: | 4, 1 2, Benton Co., Adair, #3-02, SWPM re- cycler, 23-IX-2002, EWB Survey, ex. fun- nel traps with alpha-pinene & ethanol (ODA); 1 6, 1 2, Benton Co., Adair, #30— 02, SWPM recycler, 23-IX-2002, EWB Survey, ex., funnel trap with ethanol lure (ODA); 4 22, Benton Co., Adair, #03-02, SWPM recycler, 30-IX-2002, EWB Survey, ex. funnel traps with ethanol lure (ODA, OSU); 1 2, Benton Co., Corvallis, #03-01, SWPM recycler, 30-[X-2002, EWB Survey ex. funnel traps with ethanol lure (ODA); | 3, Multnomah Co., Portland, #51-03, SWPM importer, 23-IC-2002, EWB Sur- vey, ex. funnel traps with chalcoprax (ODA); 1 2, Multnomah Co., #51-03, SWPM importer, 12-IV-2002, EWB Sur- vey, ex. funnel traps with ethanol lure (ODA); 1 2, Wasco Co., The Dalles, #65- 42, urban vicinity, volatile blend lure (ODA). UTAH: 1 G2 2:9 Box, Eldegi@oe Brigham City, Poplar Drive, July 2000, T. Houguad (USNM); 4 66, 2 22, Cache Co., Logan, 14 October 1999, J. Karen, in Library storage facility (USNM); 2 66, 2 22, Cache Co., North Logan, 28 October 1999 (USNM); 8 66, Cache Co., North Logan City Library, 41°46’40’N, 111°49’ 11”W, 16 July 2001, elev. 4,800’, T. J. Hen- VOLUME 106, NUMBER 3 S)IL7/ Fig. 3. Distribution of Raglius alboacuminatus in the United States. ry and A. G. Wheeler, Jr. (USNM); 4 36, Cache Co., River Heights, 7-12 November 1995, Dale Nielson (USNM, USU); 13 36, 62 22, Weber Co., Ogden, Ogden Weber Tech Ctr., 41°5’56"N 111°58'10"W, 16 July 2001, elev. 4,400’, T. J. Henry and A. G. Wheeler, Jr., taken on walls and walks of the center (USNM). Rhyparochromus vulgaris (Schilling) (Figs. 4-6 ) Pachymerus vulgaris Schilling 1829: 65 (original description). Rhyparochromus vulgaris: Southwood and Leston 1959: 95 (biological notes, illus- trations); Wagner 1961: 85; Slater 1964: 1322 (catalog); Slater and O’Donnell 518 4 Figs. 4-5. 1995: 178 (catalog); Péricart 1998: 304 (description, biological notes, illustra- tions); Péricart 2001: 211 (catalog). See Slater (1964) and Péricart (1998, 2001) for extensive bibliographies. Diagnosis.—This relatively large species (Figs. 4—5) is distinguished by the black head, antennae, femora, anterior half of the pronotum, scutellum, and membrane (sometimes with a small pale or white mark at apex); and the dark punctured, pale or yellowish-brown hemelytra, with a large quadrate black spot on the inner angle of the corium. Description.—Male (n = 10): Length 6.48—7.27 mm, width 2.38—2.62 mm. Head: Width 1.40-1.57 mm, vertex 0.75—0.83 mm. Rostrum: Length 2.85—3.01 mm, ex- tending to middle coxae. Antenna: Seg- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Adult female of Rhyparochromus vulgaris. 4, Dorsal aspect. 5, Lateral aspect. ment I, length 0.55—0.61 mm; II, 1.40—1.66 fom: TE eh tS =. 3mm: VV 2S 4a Pronotum: Length 1.38—1.55 mm, basal width 2.13—2.38 mm. Female (n = 10): Length 7.62—8.26 mm, width 2.75—2.98 mm. Head: Width 1.45— 1.53 mm, vertex 0.85—0.88 mm. Rostrum: Length 3.14—3.22 mm, extending to middle coxae. Antenna: Segment I, length 0.58— 0.63 mm; I, 1.43—1.50 mm; III, 1.08—1.28 mm; IV, 1.25—1.38 mm. Pronotum: Length 1.55—1.65 mm, basal width 2.48—2.73 mm. Large, oval, shiny, dark brown to black with pale or white areas on the pronotum, hemelytra, and sometimes the apex of the hemelytral membrane. Head shiny black, covered with short, dense, silvery setae, ex- cept for more glabrous vertex posterior to midline of eyes. Labium uniformly fuscous VOLUME 106, NUMBER 3 or black. Antenna uniformly fuscous to black. Pronotum quadrate, basal margin nearly straight, lateral margins explanate and weakly recurved; anterior lobe shiny black including explanate margins, weakly swollen; posterior lobe, including margin pale yellow to white, narrowly fuscous across base, uniformly dark punctured, less so laterally. Scutellum equilateral, shiny black, finely punctate. Hemelytron pale or yellowish brown; clavus black on inner half, with a pale or white blotch at base; corium yellowish brown with rows of dark punctures, apical third with a fuscous or black blotch, area bordering membrane pale or whitish; membrane black, sometimes with a small pale blotch at middle of apex. Ventral surface shiny black, with dorsal third of posterior pleural area of pronotum, posterior half of metanotum, and acetabulae pale yellowish or white. Legs: Coxae and trochanters fuscous to black; femora black, narrowly yellowish brown apically; meta- tibia black, pro- and mesotibiae yellowish brown, more fuscous basally and apically; tarsi and claws brown to fuscous. Habitat.—Raglius vulgaris frequents clearings, forest edges, mixed forests, parks, woods, borders of gardens, and other shaded biotypes (Péricart 1998), often oc- curring under loose soil litter and stones (Wagner 1961). This species overwinters as adults and nymphs in sheltered places, often in company with other lygaeoids. Putshkov (1969) found as many as 200 adults and 5" instars under the bark of a beech tree near mid August in Russia. Adults and nymphs may become active on warm days during fall and winter. Biology.—According to Péricart (1998), females begin ovipositing in early May, de- positing eggs one at a time in ground litter. Eggs hatch through July. This species is known to feed on the seeds of numerous plants, including raspberry, nettles, and sage, and on the fallen seeds of elms, poplar, and other plants. The first adults appear in the Ukraine in early July (Putchkov 1969). Most 519 individuals feed until late September before seeking overwintering shelter. Distribution.—In the Old World, R. vul- garis is widespread in Europe; north Africa, including Algeria, Morocco, and the Canary Islands; and Asia from western Russia to China, Mongolia, and Korea (Péricart 2001). This species is recorded for the first time in the United States (Fig. 6) from two coun- ties in Oregon (Multnomah and Wasco) and one county in Washington (Pierce). New United States records OREGON: 1 5, Multnomah Co., Portland, funnel trap, alpha-pinene & ethanol lure, 18 September 2001 (USNM); 2 66, Multnomah, Port- land, #51-04 SWPM importer, 23-IX-2002, EWB Survey, ex. funnel traps with ethanol lure (ODA); | 3d, Multnomah Co., Portland, #51-03, SWPM importer, 23-VII-2002, EWB Survey, ex. panel traps with spruce volatile blend lure (ODA); 1 ¢, Multnomah Co., Portland, #51-19, SWPM recycler, 19- [X-2002, EWB Survey, ex. funnel traps with alpha-pinene & ethanol (ODA); 1 d, Multnomah Co., Portland, #51-04, SWPM importer, 2- X (Oct) 2002, EWB Survey, ex. funnel traps with exotic /ps lure (ODA); 1 3d, Multnomah Co., Portland, #51-07, 16- & (Oct) 2002, EWB Survey, ex. funnel traps with apha-pinene & ethanol (ODA); 3 22, Multnomah Co., Portland, #51-19, SWPM recycler, 3-X (Oct) 2002, EWB Sur- vey, ex. funnel traps with alpha-pinene & ethanol & panel trap with ethanol (ODA); 1 2, Wasco Co., The Dalles, funnel trap, ethanol lure, mill site, 25 September 2001 (USNM); 1 2, Yamhill Co., Newberg, #71- O01, SWPM recycler, 23-IX-2002, EWB Survey, ex. funnel traps with chalcoprax (ODA). WASHINGTON: 6 36, 8 22, Pierce County, NW Orting, N 47°06.000, W 122°12.860, 5 November 2001, Veronica Kuebler coll., hand collected, ODA 999- 1010 S (USNM). Xanthochilus saturnius (Rossi) Xanthochilus saturnius is a widespread Palearctic species, ranging throughout much of Europe, northern Africa, and west- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON PEND OREMLLE CREGON ern Asia (Péricart 2001). Henry and Adam- ski (1998) gave the first North American report of this species (as Rhyparochromus saturnius) based on specimens from 17 counties in California. Since the Henry and Adamski (1998) re- port, specimens of X. saturnius have been intercepted in Japan on produce exported from California. Dr. J. A. Slater (personal communication) identified two specimens sent to him by Goro Takihiro (Yokahama Plant Protection Station, Yokahama, Japan) in September 1999, adding a new twist to the impact invasive insects may have in for- eign lands. Distribution of Rhyparochromus vulgaris in the United States. The following specimens identified by J. D. Lattin represent the first state record of X. saturnius in Oregon. Specimens examined.— OREGON: | o, Benton Co., Corvallis, #03-01, SWPM im- porter, 29-X (Oct) 2002, EWB Survey, ex. funnel traps with exotic /ps lure (ODA); | ?, Benton Co., Corvallis, #03-01, SWPM importer, 30-IX (Sept) 2002, EWB Survey, ex. funnel traps with exotic /ps lure (ODA). DISCUSSION As Hoebeke and Page (2002) aptly not- ed, ‘“‘Because of dramatic increases in in- ternational travel, combined with a boom- VOLUME 106, NUMBER 3 ing world-trade industry, plant and animal species have become globetrotters.’” Henry and Adamski (1998) indicated it was not surprising that the Palearctic rhyparochrom- id Xanthochilus saturnius (Rossi) had be- come established in California, considering that over the previous nine years, it had been intercepted more than 40 times at ports-of-entry in the United States by APHIS/PPQ personnel, almost exclusively on pallets of tile and marble imported from the Mediterranean Region. Since that report (Henry and Adamski 1998), there have been more than 20 additional U.S. intercep- tions of X. saturnius, including new records reported in this paper for Oregon. Similarly, Raglius alboacuminatus has been intercept- ed 22 times since 1990 and Rhyparochro- mus vulgaris, 67 times during the same pe- riod. Lattin and Wetherill (2002) recently documented the establishment of another Old World heteropteran, the oxycarenid Metopoplax ditomoides (Costa), which was found on commercial hazelnut, Corylus av- ellana (L.), and swarming in large numbers around homes in several Oregon counties; and Hoebeke and Carter (2003) reported large numbers of the pentatomid Haly- omorpha halys (Stal) in eastern Pennsyl- vania, a potential agricultural pest from eastern Asia. Between 1989 and 2002, M. ditomoides was intercepted 73 times at U.S. ports-of-entry, and H. halys, 15 times. The frequency of interceptions for the above ex- otic bugs makes it fairly certain that they arrived in this country as stowaways in in- ternational commerce. While invasive species such as the Hal- yomorpha halys are potential agricultural pests, others, such as Raglius alboacumi- natus, Rhyparochromus vulgaris, and Xath- ochilus saturnius might affect our environ- ment in other ways. Hoebeke and Page (2002) pointed out that nonindigenous in- sects have the potential to disrupt native en- vironments, threaten fragile ecosystems, and reduce local biodiversity. The environ- mental effects of the two alien rhyparo- 521] chromids reported in this paper must await further assessment. ACKNOWLEDGMENTS I am grateful to Alan Hardy (California Department of Food & Agriculture, Sacra- mento), Eric H. LaGasa (WDA), James R. LaBonte and Kathleen J. R. Johnson (ODA), John D. Lattin (OSU), Jay B. Karen and Alan H. Roe (USU), and Michelle Nac- hand (Division of Agriculture, County of Santa Clara, San Jose, California) for their help in documenting the distribution of Raglius alboacuminatus and Rhyparochro- mus vulgaris, and James A. Slater (Univer- sity of Connecticut, Storrs [UCS]) for shar- ing his notes on the export of Xanthochilus saturnius from California to Japan. Without their generous help, these bugs would have gone undetected. I express special gratitude to Jay Karen and Richard W. Baumann (Brigham Young University, Provo, Utah) for their local hospitality and assistance with fieldwork while surveying for R. al- boacuminatus in Utah in 2001; Raymond J. Gagné (Systematic Entomology Laborato- ry, ARS, USDA, % National Museum of Natural History, Washington, D.C. [SEL]) and Eric Guilbert (Muséum National d’Histoire Naturelle, Entomologie, Paris) for translating parts of several French arti- cles; Michele A. Touchet (SEL) for the pho- tographs used in this paper; and Robert W. Carlson (SEL, Beltsville, Maryland) for searching the Systematic Entomology Lab- oratory Identification System (SELIS) da- tabase for records of invasive bugs inter- cepted at U.S. ports-of-entry by APHIS/ PPQ (Animal and Plant Health Inspection Service, Plant Protection and Quarantine). I also give special thanks to A. G. Wheeler, Jr. (Clemson University, Clemson, South Carolina) for companionship while con- ducting fieldwork and for our many discus- sions involving invasive species. John D. Lattin, Michael G. Pogue (SEL), James A. Slater (UCS), David R. Smith (SEL), and A. G. Wheeler, Jr. kindly reviewed the man- uscript and offered suggestions for its im- provement. LITERATURE CITED Goeze, J. A. E. 1777-1783. Entomologische Beytrage zu des Ritter Linne zwolften Ausgabe des Natur- systems. 3 volumes. Weidmann, Leipzig. [1777, 1: i-xvi + 1-736; 1778, 2:I-Ixxu + 1-352; 1779, 3(1): I-xl + 1-390; 1780, 3(2): i—xxiv + 1—350; 1781, 3(3): i-xlvini + 1-439; 1783, 3(4): 1-xx + 1-178]. Henry, T. J. and D. Adamski. 1998. Rhyparochromus saturnius (Rossi) (Heteroptera: Lygaeoidea: Rhy- parochromidae), a Palearctic seed bug newly dis- covered in North America. Journal of the New York Entomological Society 106: 132-140. Hoebeke, E. R. and M. E. Carter. 2003. Halyomorpha halys (Stal) (Heteroptera: Pentatomidae): A po- lyphagous plant pest from Asia newly detected in North America. Proceedings of the Entomological Society of Washington 105: 225-237. Hoebeke, E. R. and G. Page. 2002. Longhorned beetles of the genus Anoplophora and lithography: alien invaders in the eye of the beholder. American En- tomologist 48: 200—206. Lattin, J. D. and K. Wetherill. 2002. Metopoplax di- tomoides (Costa), a species of Oxycarenidae new PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON to North America (Lygaeoidea: Hemiptera: Het- eroptera). Pan-Pacific Entomologist 78: 63—65. Péricart, J. 1998. Faune de France. France et Régions Limitrophes. Hémipteres Lygaeidae Euro-Médi- terranéens. Systématique: Troisieme Partie Rhy- parochrominae (2). Fédération Frangaise des So- ciétés de Sciences Naturelles, Paris. 3 volumes. 84A: 468 pp; 84B: 453 pp.; 84C: 487 pp. . 2001. Family Lygaeidae, pp. 35-220. In Au- kema, B. and C. Rieger, eds. Catalogue of the Het- eroptera of the Palearctic Region, Vol. 4. Penta- tomomorpha I. Putshkoy, V. G. 1969. Vypusk 2: Ligeidi. Fauna Ukrai- ni. Tom 21. Kiev. 388 pp. (In Ukrainian; not seen.) Schilling, P. S. 1829. Hemiptera Heteroptera Silesiae systematice disposuit. Beitrage zur Entomologie 1: 34-92. Slater, J. A. 1964. A catalogue of the Lygaeidae of the World. 2 volumes. University of Connecticut, Storrs, 1,688 pp. Slater, J. A. and J. E. O'Donnell. 1995. A Catalogue of the Lygaeidae of the World (1960-1994). New York Entomological Society, New York, 410 pp. Southwood, T. R. E. and D. Leston. 1959. Land and Water Bugs of the British Isles. Frederick Warne & Co., London, 436 pp. Wagner, E. 1961. Zur Systematik der Gattung Rhypar- ochromus Hahn, 1826 (Hem. Het. Lygaeidae). Deutsche Entomologische Zeitschrift 8: 73-116. PROC. ENTOMOL. SOC. WASH. 106(3), 2004, pp. 523-530 LIFE HISTORY OF MAYATRICHIA PONTA ROSS (TRICHOPTERA: HYDROPTILIDAE) IN HONEY CREEK, OKLAHOMA YI-KUANG WANG, JAMES H. KENNEDY Department of Biological Sciences, P.O. Box 310559, University of North Texas, Den- ton, TX 76203-0559, U.S.A. (JHK e-mail: kennedy @unt.edu); Y-KW current address: Department of Zoology, Michigan State University, East Lansing, MI 48824, U.S.A. Abstract.—The life history of Mayatrichia ponta Ross (Trichoptera: Hydroptilidae) was investigated in Honey Creek, Turner Falls Park, Murray Co., Oklahoma, U.S.A. from August 1994 to August 1995. This study provides the first comprehensive life history of a species of Neotrichiini. Mayatrichia ponta had an asynchronous multivoltine life history with considerable cohort and generation overlap; five generations were estimated, repre- senting the largest number of annual generations among Trichoptera. The winter genera- tions of M. ponta had wider head capsule widths (136—165 jm) than summer generations (121—145 pm). The growth of M. ponta was slower in the winter than the summer. The sex ratio of adults was 1.43 d: | @ and fecundity ranged from 46 to 150 eggs/female. Fifth instars and pupae aggregated on the bottom side of rocks whereas early instars were distributed evenly on all sides of rocks. Key Words: Trichoptera are a diverse and ecological- ly important order whose larvae live in a wide range of flowing and standing waters. Hydroptilidae, a species diverse family in the order Trichoptera, account for approxi- mately 19% of all Nearctic trichopteran species. With over 300 species, Hydroptil- idae are the second most diverse trichop- teran family, surpassed only by Limnephil- idae (Morse 1993, 2002). Despite their di- versity relatively few detailed studies on the biology of Nearctic hydroptilids have been published. Leucotrichia pictipes Banks was reported to be univoltine with an overwin- tering final instar in Montana (McAuliffe 1982). Resh and Houp (1986) described a univoltine cycle for Dibusa angata Ross in Kentucky, with an egg diapause through the summer and autumn and emergence in April and May. More recent studies on Ne- arctic hydroptilids were by Keiper and Wal- Mayatrichia ponta, life history, voltinism, fecundity, microhabitat ton (1999) and Keiper and Foote (2000). Keiper and Foote (2000) described univol- tine populations based on field observations in an Ohio stream for Hydroptila comsi- milis Mosely, Ochrotrichia spinosa (Ross) and O. wojcickyi Blickle. The biology of Hydroptilidae has been reported in a num- ber of studies in other biogeographical re- gions of the world. A study of hydroptilids in northeastern Spain, reported univoltine populations for Hydroptila vectis Curtis and Oxyethira frici Klapalek, bivoltine for Hy- droptila insubrica Ris, and possibly three generations per year for Hydroptila martini Marshall (Recasens and Puig 1991). Wells (1985) reported univoltine and multivoltine populations life cycles for Australian hy- droptilids, but gave no details. The biology of several eastern Palearctic hydroptilids has been described (Ito 1988, 1998; Ito and Kawamura 1980). Palaeagapetus ovatus Ito 524 and Hattori populations were reported as having approximately three generations per year while Eubasilissa regina (McLachlan) were univoltine (Ito 1988). In another study, Ito (1998) reported populations of Palaeagapetus flexus Ito as univoltine and P. ovatus as having 3—4 generations in 2 years. The genus Mayatrichia Mosely was first established in 1937 (Mosely 1937). The ge- nus is in the tribe Neotrichiini under the subfamily Hydroptilinae. Mayatrichia has sIx species distributed over North and Cen- tral America, and northern South America. Honey “Creekw an *TLumer? Pallse’ Park; Oklahoma is the type locality for Mayatri- chia ponta Ross (Ross 1944). The species has also been reported in Texas and Wyo- ming (Moulton and Stewart 1996). The at- tenuated head of Mayatrichia make their larvae easy to identify in all instars (Wig- gins 1996a). There have been no published studies on the ecology and life history of any Maytrichia species or any other Neo- trichiini. The purpose of this study is to de- scribe the life cycle and aspects of the bi- ology of M. ponta through field studies. MATERIALS AND METHODS Study site and sampling.—Honey Creek is a second order permanent limestone stream, originating in the Arbuckle Moun- tains, and flowing northeastward into the Washita River. The geological formation of Honey Creek changes sequentially from cherty limestone and sandstone to ‘Fern- vale’ limestone and, eventually, to Washita alluvium (Ham 1969). Honey Creek de- scends from an elevation of 420 m at its origin to 235 m at Washita River (Ham 1969). The most rapidly descending areas are Bridal Veil Falls and Turner Falls. The pH, raised by photosynthetic activity from aquatic plants and algae, causes precipita- tion of calcium and magnesium carbonates to form thick layers of travertine in the study area (Minkley 1963). The study riffles were positioned within a 900 m zone upstream of Bridal Veil Falls PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON in Honey Creek; Turner ‘Falls Pack, Oklahoma (34°25.407'N and 97°04.210'W). The upstream riffles were mainly composed of cobbles and pebbles; downstream riffles were mainly composed of travertine sub- strates with sparse to aggregated stones. The creek width ranged from 15 to 30 m. Riffle depth ranged from | to 2 cm in the travertine beds to 20 cm in the cobble sub- strate under base flow condition. The mean discharge is 1.06 m?/s. The mean pH was 8.0 and the dissolved oxygen was always near saturation. These values were similar to ones reported by Reisen (1975, 1976) suggesting that the water conditions in this stream have been stable over the last 25 years. Water temperature ranged from 9 °C in the winter to 25 °C in the summer during the study period. Air temperature ranged from —6. / Moa lie: Samples of larvae, pupae, and adults were taken from August 1994 until August 1995, weekly from August to September 1994, and biweekly for the remainder of the study. Adults.—Mayatrichia ponta adults were collected with an 8-watt UV light during each field trip. The trap was set approxi- mately one meter from the shoreline near downstream riffles and was operated for one hour after sunset. Flight periodicity was determined from samples taken with aerial nets. Fecundity was studied by dissecting field-collected females, and lab-reared fe- males when no field collected adults were available. Larvae.—Samples indicated that the ma- jority of M. ponta larvae were found on rocks greater than 15 cm across that were resting on the travertine stream substrates. On each sampling date six rocks ranging from 15 to 30 cm in diameter were collect- ed along a randomly chosen transect across the stream. The upper size limit was estab- lished based on our ability to handle and process the material in the field. Organisms and debris on the surface of the rock were brushed into a bucket filled with filtered stream water. The contents of the bucket VOLUME 106, NUMBER 3 were then passed through a sieve to con- centrate the sample. A 150-~m sieve was initially used for collections in August 1994; for subsequent sampling a 38-ym sieve was used to collect the earliest instars. Collected materials were preserved in Kah- le’s solution then stored in 80% ethanol un- til processed in the laboratory.. Head capsule width and length were measured using an Olympus SZH dissect- ing microscope coupled with an Olympus CUE-2 image processing system. Instars were separated by a scatter plot of head capsule width against head length. The prominent sclerites of the fifth instar also were used to help distinguish instars. The length of the head capsule (HCL) was mea- sured from the anterior margin of the frons to the posterior margin of head sclerites. The width of the head (HCW) was mea- sured as the distance of the head capsule across the eyes. Diet examinations of larvae were deter- mined from collections made during July 1996. Freshly collected larvae were relaxed in carbonated water for 5 minutes to reduce regurgitation of the gut contents and then preserved in ethanol. Fore- and midguts were dissected, placed in glycerin on.a mi- croscope slide, and scanned under a com- pound microscope with 800—1,000 magni- fication. The proportion of each component of the diet was quantified by relative per- centage of area. Larval microhabitat——The effect of wa- ter current on the spatial distributions of M. ponta larvae and pupae on each surface (top, bottom, front, back, left, and right) of individual rocks was examined in March 1996. This sampling occurred during a pe- riod with high numbers of larvae and pupae were present. Prior to each biological sam- pling, water flows were estimated in front of individual rocks by a Pygmy flow meter and on each rock surface exposed to water current using a modified Gessner current meter (Hynes 1970, Wallace 1975). After measuring currents, each rock surface was brushed, and the contents were concentrat- Nn i) nN ed in a sieve (38 wm) and preserved with Kahle’s solution in separate bottles. Larvae were sorted to instar and counted in the lab. The area of each side of each rock was measured using the aluminum foil weight- area method described by Doeg and Lake (1981). Counts of each instar and pupae on each side of each rock were converted to a stan- dard density, number of individuals/0.01 m’. This area was selected because it ap- proximates the mean surface area of all stones collected. Both population size and surface area data were normally distributed (Shapiro-Wilk test for normality, a = 0.05). A one-way ANOVA was used to test if the means of current velocities on six sides were significantly different. A Pearson Cor- relation Analysis was used to determine the coefficient between the population size of M. ponta and surface area of substrates. A one-way ANOVA was also used to examine the mean densities of each instar on the six sides. The Student-Newman-Kuels multiple comparison test was used to determine sta- tistically significant differences between means. All statistical analyses followed Zar (1984) using SAS software (1991). Rearing.—Live larvae and pupae were transported to the laboratory and placed in aquaria containing natural substrates and aerated stream water for rearing. Pupae were reared individually for association with adults. Behaviors were observed in these aquaria. Larval behaviors were also observed under a dissecting microscope. The overall behavior, fecundity, life cy- cle, voltinism, and life history of M. ponta were interpreted from information gathered by the field activities described above and laboratory observations. Voucher materials, including representatives of all life stages, are deposited in the University of North Texas Elm Fork Natural Heritage Museum. RESULTS AND DISCUSSION Eggs.—Despite numerous attempts, we were unable to induce oviposition in fe- males collected in the field or reared in the 160 140 | 120 100 80 Head Capsule Width (micron) 60 50 100 skele PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 300 200 250 350 Head Capsule Length (micron) Bice ie lab. However, female M. ponta taken in light traps often released egg masses when they became immersed in preservative. These egg masses were contained in a light yellow gelatinous fluid. The diameter of eggs in female abdomens was 52—76 pm (x = 65 pm, n = 8). Eggs of Agraylea mul- tipunctata Curtis and Oxyethira costalis Curtis, were also reported to be in a gelat- inous mass (Nielsen 1948). However, D. angata laid eggs singly (Resh and Houp 1986), indicating that oviposition strategies vary among the Hydroptilidae. Larval distribution.—Instars were sepa- rated by a scatter plot of head capsule width against head capsule length (Fig. 1). Five instars were determined for M. ponta, which is typical for Hydroptilidae (Nelson 1948). Nearly all larvae of M. ponta were found on or under rocks. However, a few 5" instar larvae were found under moss mats beneath fast flowing water and in abandoned black- fly pupal cases. A one-way ANOVA showed significant difference in flows Scatter plot of Mayatrichia ponta head capsule width versus length illustrating instars. among the six sides of rocks (n = 10, a = 0.05, P = 0.0001). The populations on the bottom side of the rocks were separated as a distinct group from those on other surfac- es (Student Newman Kuels test, a = 0.05). The top surface had the highest mean flow of 3244 cm/s (SD =110:9%cmi/s); Densities of instar I through IV on the six surfaces of rocks were not significantly different. However, densities of fifth instars and pupae were significantly greater on the bottom of the rocks than other sides (Stu- dent Newman Kuels test, a = 0.05). These observations suggest that instars I—IV of M. ponta are dispersed evenly on all sur- faces, but final instars preferred the bottom surface were low flows occurred. These shifts in microhabitats are similar to those reported for Hydropsyche pellucidula Cur- tis (Boon 1979, Muotka 1990), and H. an- gustipennis Curtis (Muotka 1990). Diet.—Wiggins (1996b) indicated that Mayatrichia species are scrapers. Exami- nation of 13 full larval midguts found that all gut contents were 100% unidentifiable VOLUME 106, NUMBER 3 (Sa a = Fi 400 Peak 50% of larvae 2 | Oviposition and pupae @ 300