be (Vallee fnntohaty Ad 2 Baofim ei eR 29 niet tie ep aera ate Uv Ye Paiste Vee a meat 4) gi eM Aeh wt Syhapnesy etynninn ce eens tat V8 a STN 9 Fs Bang STM Be as eens eidrhdnddbalic deatiocate a tinny ena bey as = areas ernie ne repre paler SM ychetarhe ama 4a FDOT a Da BA Rrra ne PONS V ANSI sonra eh Sinai shame a LAY a, Weare fe th 1 He rat ah i Thu p : i ; } t ‘ , 4 me WRG) a A oh | \ i Di a et } ea, t I pea Ait ; Na AM VG i TAGE ie 1 j , wey ' i i ee f : , L r i ‘ht \ iy h 1 . ‘ , t i j i : f i r, ne i ree | 4 At Ne i ; te Va Ve nal a , Dey i 7 } : ; EW al Naat VAR AH AIM THE 7 ne | St j ae , ain wy, Mh Wi y ‘ Ta na ah i r " Th, nna i | yt I) vehi ra ' i ( iY of My hit i 1. Pw py iy i if 1 y Wy aq i nN j : i Tike ya | i ; oe Tat gy sta in? anne NM is a a a ; ma 7 r 1 fel ' i! 1 ie i : i ye : hic i vi rout) vy ah : ; i} in a tea ie - i Di esi : i a : i ae 7 iy i i 1) 7 } : Fat eae ij De iy 1 ii hy ! ‘ A } t Aye i oy i y 1 ie | fin ate ; ; ul ! i Wid r i i i Me ii, Te ry ia ety a) era rN ta oe : a 1 iL i 1 # ai ‘ - Nit Payee tier ira fa a i f { . i ty i: i ni a von I mn i : ¥ 27 || ” i ae f ; 1 - 7 t i r i in in ik ms : ‘ a if f at iy 1) ey Ps eT ii na yO wet a et ' if im, i H i oy i i 1 ial ; i ; i : 7 i) ith ; Wah a) i * : 1 ij \ j ¥ f i i NUN 6 i ; , fi ; nt et hae ie | ’ i { ie i M ty ; i . ie A ¥ com ey I! i : i ms) a al a gia iy yt i : DF i : id yi ait RON BU ewe AY ND ; a , 1 i} i eee { | a i Nant ; fi}, ‘ ; i f panes Ht ea 7 : i ean ; | i Dt Aa a) i Ad ; 1 HH 7 Via) j oa VOL. 101 JANUARY 1999 “x PROCEEDINGS of the %, PUBLISHED QUARTERLY CONTENTS ADAMSKI, DAVID—Blastobasis graminea, new species (Lepidoptera: Gelechioidea: Coleo- phoridae: Blastobasinae), a stem borer of sugar cane in Colombia and Venezuela .......... BAIER, TEVIS M.—A revision of the flavidus group of the genus Chrysops Meigen (Diptera: SEATS TAN AC ee ee LS eae Nichia) oe eacr oh eae MRE Spe Set eoe ate SRE ee alata RPS oe aa cTE eee cde saa ee ean state BATRA, SUZANNE W. T.—Biology of Andrena (Scrapteropsis) fenningeri Viereck (Hyme- nopteraswAndLenidae) harbinger of: Springy asa) cclssticmn cee neldemiarre msslinesheisicle Re avai seh eattela ta retels BRAILOVSKY, HARRY and GERASIMOS CASSIS—New genus and new species of Amorbini (Heteroptera:-Coreidae) strom Australian oye. asecaedasalets tee oe sae scene come ee ead as meeee BREIDENBAUGH, MARK S. and BRADLEY A. MULLENS—Two new western Nearctic Culicoides Latreille (Diptera: Ceratopogonidae) descibed from all stages ..................... CARVALHO, CLAUDIO JOSE BARROS DE—Revision, cladistics and biogeography of the Neotropical genus Souzalopesmyia Albuquerque (Diptera: Muscidae) ....................-005- EASTON, EMMETT R. and WING-WAH PUN—Observations on twelve families of Homoptera Inv NViacaussoutheastenti@ hina mrom,)) 98 9) tontheypreseny minyaus als amas eens se ae euatela ys GAGNE, RAYMOND J. and JOHN F. DOANE—The larval instars of the wheat midge, Sito- diplosis mosellana (Gehin))(Wiptera: ECecidomyiidac))e..34ce ee es sae nee ae sees eie see tese GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Life history and description of imma- ture stages of Trupanea arizonensis Malloch (Diptera: Tephritidae) on Trixis californica Kellogg var. californica (Asteraceae) in southern California ...............0...0ee cence eee ee eee HARRIS, S. C. and L. J. DAVENPORT—New species of Hydroptilidae (Trichoptera) from the AINAZONEre SLONAOMMOLNeASLELM) RELUmec ee seach ole Ae te ca bs Mee ates MERU seo etait cleeehe a ania ee JENSEN, ANDREW S. and MANYA B. STOETZEL—An examination of the North American aphid species currently placed in Ovatus van der Goot (Hemiptera: Aphididae) with the ESCH OD Wiia NE MP SMUS wee sees mas Cheat err ee A AMS hse cl Coe en wera liad uia Fey earn 2 eee ata JOHNSON, NORMAN F., LUCIANA MUSETTI, JAMES B. JOHNSON, and KERRY KA- TOVICH—The larva of Pelecinus polyturator (Drury) (Hymenoptera: Pelecinidae) ........ (Continued on back cover) NO. 1 (ISSN 0013-8797) 164 175 106 69 149 123 99 57 Tf) 26 39 THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 1999 MICHAEL E. SCHAUFF, President MICHAEL G. POGUE, Treasurer DAvib G. FurtH, President-Elect JOHN W. Brown, Program Chair STUART H. McKAmey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair HoL.is B. WILLIAMS, Corresponding Secretary WARREN E. STEINER, Past President ANDREW S. JENSEN, Custodian DAviD R. SmitH, Editor Publications Committee THOMAS J. HENRY WAYNE N. MATHIS GARY L. MILLER, Book Review Editor Honorary President LouIsE M. RUSSELL Honorary Members ALAN STONE KARL V. KROMBEIN RONALD W. HODGES 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 7:30 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. 599-600 OF THE JULY 1998 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: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC- West, Beltsville, MD 20705. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 19 January 1999 Second Class 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. 101(1), 1999, pp. 1-25 EGG ARCHITECTURE OF NAUCORIDAE (HETEROPTERA): INTERNAL AND EXTERNAL STRUCTURE OF THE CHORION AND MICROPYLE ROBERT W. SITES AND BECKY J. NICHOLS Enns Entomology Museum, Department of Entomology, University of Missouri, Co- lumbia, MO 65211, U.S.A. (e-mail: bugs @showme.missouri.edu) Abstract.—The chorion and micropyles of 27 species of Naucoridae, including 21 spe- cies of Ambrysus, are described and each description is supported with scanning electron micrographs. Photomicrographs of thick sections through the chorion or micropyle of 10 of these species also are presented. Chorionic sculpturing differs interspecifically in Am- brysus and based on the species studied, eggs of Ambrysus, Limnocoris, and Pelocoris have 2—3 micropyles. Key Words: The Naucoridae (sensu lato), or creeping water bugs, comprise 394 described species worldwide (see La Rivers 1971, 1974, 1976; Polhemus and Polhemus 1982, 1988, 1994; Nieser et al. 1993; Liu and Zheng 1994; Polhemus 1994; Nieser and Chen 1996). As is typical among insect families, the adults have received the greatest amount of morphological research; accord- ingly, taxonomic treatments emphasize adult characters. Although nymphal stages have been described for several species of Naucoridae, details of egg structure largely have been ignored. Line diagrams from light microscopy have been presented for a few species; however, elucidation of fine detail is not possible with this technique. Recent reports for several species [Ambry- sus lunatus Usinger (Sites and Nichols 1990), Cryphocricos hungerfordi Usinger (Sites and Nichols 1993), Pelocoris poeyi Guérin Méneville (Sites 1991), and several species of South American Ambrysus and Pelocoris (L6pez Ruf 1989)] have included scanning electron micrographs that have re- vealed interspecific differences in chorionic patterns. Egg, chorion, micropyle, Naucoridae Eggs of most naucorid species, for which Oviposition is known, either are adhered to plants (exophytic oviposition) or to rock substrata (Hinton 1981). For example, eggs of A. /unatus are adhered to plants (Sites and Nichols 1990), Ambrysus mormon Montandon to pebbles (Usinger 1946), Aphelocheirus aestivalis (Fabricius) proba- bly to rocks (Larsén 1927), C. hungerfordi to rocks (Sites and Nichols 1993), Lacco- coris limigenus Stal to hard substrata (Clarke and Baroudy 1990), Naucoris ma- culatus F. to plants (Lebrun 1960), and Pel- ocoris femoratus (Palisot de Beauvois) to plants (Torre Bueno 1903, Hungerford 1927, McPherson et al. 1987). More spe- cifically, P. femoratus eggs are glued to leaf- lets of Nitella and other aquatic plants with a ‘fairly generous quantity of white adhe- sive” (Hungerford 1927). In contrast, the oviposition of Ilyocoris cimicoides (L.) is endophytic, inserting eggs into submergent plant tissue (Cobben 1968), such as into stems of Ranunculus or water peppermint (Rawat 1939). Eclosion occurs through the anterior pole and, generally, a crescentic slit is made 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON through the chorion. In Coleopterocoris kleerekoperi Hungerford and species of Cryphocricos, a predetermined fracture line exists, and in J. cimicoides, a well-defined operculum faces the water, whereas the re- mainder of the egg is embedded within the plant (see Rawat 1939). Internally, the chorion of //yocoris and Cryphocricos is bilayered (Cobben 1968 and Sites and Nichols 1993, respectively), with a thick chorionic outer layer, which is perforated by pore canals, and a thinner, un- perforated chorionic inner layer. Externally, scanning electron micrographs of six spe- cies of Pelocoris and two species of Am- brysus from Argentina revealed that the chorionic surface differed among species (L6pez Ruf 1989). For the two species of Ambrysus, interspecific internal differentia- tion was subtle; however, marked differenc- es existed externally. Therefore, L6pez Ruf (1989) suggested that the external chorionic pattern is valuable as a taxonomic character and that internal chorionic attributes may be useful at the generic level. Presented herein are scanning electron micrographs of the chorion and micropyles of 21 species of Ambrysus (subfamily Cry- phocricinae) and selected species represent- ing six additional genera and five additional subfamilies. Photomicrographs of thick sec- tions of the micropyle and chorion for some species also are presented. Egg morphology is described for each species. MATERIALS AND METHODS Eggs were obtained by both oviposition and dissection. Oviposited eggs were pre- ferred for examination because they were fully developed structurally. Thus, we brought live female naucorids into the lab- oratory and maintained them individually in glass petri dishes with enough water to sub- merge them. To provide a potential ovipo- sition substrate, an aquatic plant stem (usu- ally Justicia americana) was placed in each petri dish and live food provided for each naucorid generally as one corixid (Corisel- la, Ramphocorixa, Sigara) per naucorid, daily. Most naucorids, including members of those species inhabiting lotic environ- ments, Oviposited on plants or on the dish. For the several species that did not oviposit in the laboratory or for which we did not have live specimens, eggs were dissected from females preserved in alcohol. Eggs were taken from only the common oviduct or vagina (sometimes erroneously referred to as ovarian eggs), rather than from the ovarioles, and were considered to be struc- turally well-developed because they were near the end of the reproductive tract. To allow a rapid evaluation of the reproductive tract to determine position of eggs and to minimize external damage to the specimen, a dissection technique was developed. With the insect in alcohol and ventral side up, the tip of a pair of jeweler’s forceps was in- serted into the membrane of the lateral mar- gin of the 7th abdominal segment. By mov- ing the forceps anteriorly, the sternum and laterotergites were separated from the terga of segments 4—7. The venter then was pulled laterally, separating along segmental sutures, thereby exposing the abdominal cavity. Eggs were gently removed from the abdomen with forceps and kept in 3.7 ml snap-cap glass vials in 80% ethyl alcohol. Dissected eggs often had tissue from the reproductive tract adhered to them. To re- move this tissue, an ultrasonic cleaner was used, which had a peak output of 40 watts and a frequency of 60 Hz. Vials containing eggs in ethyl alcohol were placed in the cleaner with a small amount of water for ca. 10 minutes: the exact amount of time depended on the amount of tissue to be re- moved. Various solublizers [e.g., Triton X- 100, sodium dodecyl] sulfate (SDS)] were ineffective in removing tissue. Subsequent- ly, the eggs were examined under a micro- scope and any remaining tissue was care- fully removed. Eggs then were transferred to fresh 80% ethyl] alcohol. To prepare eggs for critical point drying, both dissected and oviposited eggs in 80% ethyl alcohol were fixed with 2% glutaral- dehyde in 0.1 M phosphate buffer, washed VOLUME 101, NUMBER 1 Bigs I through micropylar plug distad of base. Ambrysus circumcinctus, cross section in 0.1 M phosphate buffer, and dehydrated through a graded alcohol series to 100% ethyl alcohol, and then placed into 100% acetone. Eggs were then critical point dried and sputter-coated. For light microscopy, the above procedure was followed through the glutaraldehyde primary fixation. After washing in 0.1 M phosphate buffer, eggs were subjected to a secondary fixation in 2% osmium tetroxide in 0.1 M phosphate buffer. Subsequent dehydration techniques were as for the scanning microscopy prep- aration. Eggs were embedded in Spurr’s resin, sectioned with a diamond knife, slide-mounted, and stained with toluidine blue. Voucher specimens of adults and eggs are housed in the Enns Entomology Muse- um, University of Missouri—Columbia. DESCRIPTIONS OF EGGS Little detail of the micropylar plug of most genera is evident externally that would allow the determination of the num- ber of micropyles contained therein. Cross sections taken at levels above the base of the plug reveal numerous canals, including transverse canals, which represent convo- lutions of the same canal(s) throughout the plug (Fig. 1). However, the number of mi- cropyles can be observed by sectioning through the base of the plug where it enters the egg. Thus, photomicrographs of thick sections taken through the base of the plug are presented for some species. Citations for the original description and subsequent descriptions of adults or imma- ture stages are given for each species. Also given are collecting localities of females from which eggs were obtained, egg mea- surements (mm + SE), sample size, and method of obtainment (1.e., oOviposited or dissected). For all naucorid species exam- ined, egg color was creamy white to beige. Subfamily Cryphocricinae Montandon 1897a Genus Ambrysus Stal 1862 Ambrysus (Acyttarus) funebris La Rivers (Figs. 2—5) Ambrysus funebris La Rivers 1948a: 103-— LOF: USA: California, Death Valley National Monument Length, 1.04; width, 0.38; n = 1; dissected. Overall appearance elongate with round- ed, asymmetrical ends. Reticulation pattern consisting generally of pentagonal to hep- tagonal units, delimited by distinctly raised lines (Fig. 2). Within each unit, poorly de- fined, low, irregular protruberances. Aero- pyles small, distinct, numerous, more than 50 per unit (Fig. 3). Anterior pole with re- ticulation and aeropyles lacking, with amor- phous micropylar plug set in shallow con- cavity (Fig. 4). Exochorion thicker than endochorion. Pore canals widest at base (Fig. 5). Ambrysus (Syncollus) circumcinctus Montandon (Figs. 6-9) Ambrysus circumcinctus Montandon 1910: 442-444. USA: Texas, Kimble Co., Junction Length, 1.11 = 0:01; width, 0:48 = 0:01; n = 10; dissected. Overall appearance elongate-oval. Retic- ulation pattern only faintly visible as im- pressed lines. Within each unit formed by 4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-7. 2-5, Ambrysus funebris. 2, 3, Chorion surface. 4, Micropylar plug. 5, Chorion section. 6, 7, A. circumcinctus. 6, Chorion surface. 7, Micropylar plug. pattern, 5—8 aeropyles set in deep individual Section through base of micropylar plug sockets (Fig. 6). Anterior pole with reticu- with two micropyles (Fig. 8). Protuberances lation pronounced with raised lines and _ other than micropyle lacking. with aeropyles absent. Micropylar plug at Exochorion ca. 4 X thicker than endo- anterior pole, ovate, and with two distinct, chorion. Pore canals widest at base lateral helical micropylar tubes (Fig. 7). (Fig.9). VOLUME 101, NUMBER 1 5 Figs. 8-13. 8, 9, Ambrysus circumcinctus. 8, Micropyle section at base of plug. 9, Chorion section. 10, 11, A. montandoni. 10, Anterior pole with micropylar plug. 11, Chorion surface. 12, 13, A. arizonus, chorion surface. Ambrysus (Syncollus) montandoni La Length, 1.34; width, 0.64; n = 1; dissected. Rivers Overall appearance elongate-oval. Retic- (Figs. 10, 11) ulation pattern generally consisting of te- Ambrysus montandoni La Rivers 1963: 1-5. tragonal to heptagonal units, delimited by VENEZUELA: Territorio Amazonas, To- sulci between adjacent hemispherical, fu- bogan runcular mounds (Fig. 10). Within each 6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON unit, 15—30 aeropyles, becoming less dis- tinct toward anterior pole. Aeropyles in sul- ci as well as on mounds. Numerous glob- ules adhered to surface (Fig. 11). (Although these globules may be artifacts, they were persistent even after sonication in 100% acetic acid.) Reticulation less conspicuous anteriorly because mounds become flat- tened until immediate vicinity of micropyle where smaller reticulation units are evident (Fig. 10). Micropylar plug at anterior pole and amorphous. Protuberances other than micropyle, mounds, and granules lacking. Ambrysus (Ambrysus) arizonus La Rivers (Figs. 12-15) Ambrysus arizonus La Rivers 1951: 320— 32): USA: Arizona, Gila Co., Jakes Corner Eength.1:45°-=0:01> width: 02/7,<=0:01: n = 8; oviposited. Overall appearance elongate-oval. Retic- ulation generally consisting of pentagonal to heptagonal units, delimited by depres- sions. Each depression with double row of elongate papillae defining unit boundaries (Fig. 12). Within each unit, chorion raised and coarsely papillose (Fig. 13). Aeropyles indistinct, evident as pitted appearance among papillae; number 20—SO per cell. Tu- bercles lacking. Anterior pole with reticu- lation and papillae less distinct. Micropylar plug amorphous; section through base of micropylar plug with two micropyles (Fig. 14). Exochorion distinctly thicker than endo- chorion. Pore canals widest in basal third (Big 15): Ambrysus (Ambrysus) buenoi Usinger (Figs. 16, 17) Ambrysus buenoi Usinger 1946: 199-200. USA: Texas, Kimble Co., Junction Length, 1.32 + 0.02; width, 0.65 + 0.02; n = 11; oviposited. Overall appearance elongate-oval. Sur- face comprising a series of anastomosing mounds with irregularly produced protuber- ances (Fig. 16). Bases of mounds with large aeropyles distributed randomly. Chorionic surface, including swells and protuberanc- es, granular (Fig. 17). Micropyle amor- phous. Chorionic surface immediately sur- rounding micropyle lacking regular surface features, although poorly defined mounds may occur. Ambrysus (Ambrysus) crenulatus Montandon (Figs. 18, 19) Ambrysus crenulatus Montandon 1897a: 13-14. ECUADOR: Napo Province, Puerto Napo Length, 1.12 + 0.01; width, 0.49 + 0.01; n = 10; dissected. Overall appearance elongate-oval. Retic- ulation generally consisting of pentagonal to heptagonal units, delimited by distinctly raised, thin walls (Fig. 18). Each unit ap- pearing as a deep socket, with approxi- mately 8-17 large, irregularly distributed aeropyles (Fig. 19). Tubercles lacking. An- terior pole with reticulation less distinct. Micropylar plug amorphous. Protuberances other than micropyle and raised reticulation lacking. Ambrysus (Ambrysus) fossatus Usinger (Figs. 20, 21) Ambrysus fossatus Usinger 1946: 191-192. ECUADOR: Napo Province, Puerto Napo Length; 1-232: 0:03; width).0'57 == 0/022 n = 3; dissected. Overall appearance elongate-oval. Retic- ulation pattern generally consisting of pen- tagonal to heptagonal units and only faintly visible (Fig. 20). Within each unit, 10—20 conspicuous aeropyles. Chorionic surface devoid of protruberances other than micro- pyle. Middle of pore canals generally par- allel-sided (Fig. 21). Micropylar plug amor- phous, acentric. Ambrysus (Ambrysus) hungerfordi Usinger (Figs. 22—25) Ambrysus hungerfordi Usinger 1946: 192- 194. VOLUME 101, NUMBER 1 ~~] ~ 9 he : > ye “£22 % i“; ® Figs. 14-19. 14, 15, Ambrysus arizonus. 14, micropyle section at base of plug. 15, Chorion section. 16, 17, A. buenoi, chorion surface. 18, 19, A. crenulatus, chorion surface. USA: Texas, Presidio Co., Big Bend Ranch Overall appearance oval (Fig. 22). Retic- State Natural Area ulation pattern generally consisting of pen- Length, 1.18 + 0.01; width, 0.63 + 0.01; n tagonal to heptagonal units, delimited by = 9; dissected. low, raised ridges (Fig. 23). Within each 8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON eo we Rd . Figs. 20-25. 20, 21, Ambrysus fossatus. 20, Chorion surface. 21, Chorion section. 22-25, A. hungerfordi. 22, Whole egg. 23, Chorion surface. 24, Micropylar plug. 25, Chorion section. unit, 50-120 aeropyles more or less evenly — with reticulation less pronounced, aeropyles distributed. Protruberances, tubercles, and fewer. Chorionic inner layer 60% as thick papillae lacking. Micropylar plug anterior as chorionic outer layer (Fig. 25). Pore ca- and amorphous (Fig. 24). Anterior pole nals narrowest at middle. VOLUME 101, NUMBER | 9 Figs. 26-31. 26, Ambrysus inflatus, chorion surface. 27—29, A. lunatus. 27, Chorion section. 28, Pore canal containing bacteria. 29, Micropyle section at base of plug. 30, 31, A. mormon, chorion surface. Ambrysus (Ambrysus) inflatus La Rivers Length, 1.37 + 0.01; width, 0:68 = 0.01; n (Fig. 26) = 10; dissected. Ambrysus inflatus La Rivers 1953a: 1316— Overall appearance elongate with 1318. rounded ends. Reticulation pattern gener- MEXICO: Jalisco, Chapala ally consisting of pentagonal to heptago- 10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 32-37. 32, Ambrysus mormon, anterior pole with micropylar plug. 33, A. occidentalis, chorion surface with antheform processes. 34, 35, A. plautus, chorion surface. 36, 37, A. portheo. 36, Anterior pole. 37, Chorion surface with antheform processes. nal units, delimited by a series of irregu- larly raised ridges with numerous tiny pa- pillae between (Fig. 26). Scattered groups of papillae depressed, creating pitted ap- pearance. Anterior pole with amorphous micropylar plug, without raised reticula- tion or papillae. The appearance of the chorion is virtu- ally indistinguishable from that of A. /una- tus Usinger (see Sites and Nichols 1990). VOLUME 101, NUMBER 1 Ambrysus (Ambrysus) lunatus Usinger (Figs. 27—29) Ambrysus lunatus Usinger 1946: 203—205. Ambrysus lunatus: Sites and Nichols 1990: 800-808. USA: Texas, Kimble Co., Junction Endochorion subequal to exochorion in thickness (Fig. 27). Pore canals parallel-sid- ed and widest at base. Presence of bacteria in pore canals as detected by transmission electron microscopy (Fig. 28). Micropylar plug with three micropyles (Fig. 29). Original description of egg was given by Sites and Nichols (1990). Ambrysus (Ambrysus) mormon Montandon (Figs. 30-32) Ambrysus mormon Montandon 1909: 48—49. Ambrysus mormon: Usinger 1946: 186— 187, Plate X. USA: New Mexico, Lincoln Co., Hondo Rensth1-70"= 0:03? width; 0:98 = 0.02; n = 6; oviposited. Overall appearance elongate-oval. Retic- ulation pattern generally consisting of te- tragonal to heptagonal units, delimited by a series of depressions between composite tu- mescences (Fig. 30). Tumescences rarely entire, usually completely or incompletely divided into two to four components, oc- casionally with a smaller, central tumes- cence or depression (Fig. 31). Tumescences, depressions, including reticulation, covered with papillae. Aeropyles generally distrib- uted, neither clustered nor concentrated. Tu- mescences becoming flatter and less divid- ed toward anterior pole. Micropylar plug amorphous, acentric (Fig. 32). Usinger (1946) indicated that eggs are glued to the surface of pebbles and are sub- oval with a buttonlike micropyle at the an- terior pole. Ambrysus (Ambrysus) occidentalis La Rivers (Fig. 33) Ambrysus_ occidentalis La_ Rivers 322-325. LOS ie USA: Arizona, Gila Co., Jakes Corner Length, 1.28 + 0.01; width, 0.62 + 0.00; n = 3; dissected. Overall appearance elongate-oval. Reticu- lation pattern generally consisting of hexag- onal units, delimited by raised boundaries. Single antheform process extending outward from center of each unit, distal end concave and expanded (Fig. 33). Margins surrounding distal concavity irregular, never in contact with adjacent antheform process. Base of an- theform process widest, gradually narrowing distally. Papillae covering surface from raised reticulation to base of antheform process. 25— 40 irregularly distributed aeropyles distinctly visible around base of antheform process. Anterior pole with micropylar plug, with pat- tern less distinct, antheform processes and pa- pillae absent. Ambrysus (Ambrysus) plautus Polhemus and Polhemus (Figs. 34, 35) Ambrysus plautus Polhemus and Polhemus 1982: 326-328. MEXICO: Chihuahua, Cusarare Keneth 1-122: 0:01; width; 0:54 >= 0:02; n = 3; dissected. Overall appearance elongate-oval. Retic- ulation pattern generally consisting of pen- tagonal to heptagonal units, delimited by depressed lines. Within each unit, chorion raised and distinctly papillose (Fig. 34). Aeropyles large, distinct; number 3—12 in interior of unit, 1O—18 in depressed perim- eter of unit (Fig. 35). Tubercles lacking. Anterior pole with pattern and papillae less distinct. Micropylar plug amorphous. Ambrysus (Ambrysus) portheo La Rivers (Figs. 36-39) Ambrysus portheo La Rivers 1953a: 1320-— 1321. MEXICO: Nuevo Leon, La Nogalera Length, 1.56 + 0.02; width, 0.89 + 0.01; n = 2; oviposited. 12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 38-43. A. pudicus. 40, Anterior pole with micropylar plug (dislodged). 41, Chorion surface. 42, Micropyle section at base of plug. 43, Chorion section. General appearance robust with apices slightly truncate (Fig. 36). Reticulation pattern generally consisting of pentagonal to hexagonal units, delimited by raised, fence-like boundaries. Ectal edge of 38, 39, Ambrysus portheo. 38, Micropyle section at base of plug. 39, Chorion section. 40—43, boundary with irregularly-spaced, deep notches extending ca. Y; distance to base. Single antheform process extending out- ward from center of each unit, the distal end of which is concave and expanded VOLUME 101, NUMBER 1 (Fig. 37). Margins surrounding distal con- cavity irregular, never in contact with ad- jacent antheform process. Base of anthe- form process widest, gradually narrowing distally. Papillae covering surface _ be- tween unit boundaries and base of anthe- form process. Irregularly spaced aeropy- les occasionally visible in gaps between papillae. Anterior pole with pattern re- duced around micropylar plug, antheform processes and papillae absent. Micropylar plug with two micropyles (Fig. 38) Exochorion slightly thicker than endo- chorion. Pore canals bulbous at base. An- theform processes solid, without ducts (Fig. 59): Ambrysus (Ambrysus) pudicus Stal (Figs. 40—43) Ambrysus pudicus Stal 1862: 460. USA: Texas, Kimble Co., Junction Penothy Pi2 == 0:01; width; 0:61 = 0.01; n = 11; oviposited. Overall appearance elongate-oval. Retic- ulation pattern generally consisting of pen- tagonal to heptagonal units, delimited by slightly raised boundaries (Fig. 40). Within each unit, protuberances of two sizes: larger tubercles and smaller pustules (Fig. 41). Pe- rimeter of each unit with a series of ca. 12— 25 pustules; additional pustules usually near center of unit and among tubercles. Tuber- cles number 1-5 per unit and situated near center, occasionally surrounding one or more pustules. Outline of pustules round, of tubercles amoebiform. Anterior pole with pattern faintly visible, protuberances other than micropyle lacking. Micropylar plug amorphous, acentric, with two micropyles (Fig. 42). Exochorion thicker than endochorion (Fig. 43). Pore canals wide, ca. 0.4 X length. Ambrysus (Ambrysus) pulchellus Montandon (Figs. 44, 45) Ambrysus pulchellus Montandon 1897a: 16. USA: Texas, Kimble Co., Junction 13 Keneth, 1°36 0:02; width; 0:65 = 0:01; n = 11; oviposited. General appearance elongate-oval with rounded apices. Reticulation pattern gener- ally consisting of pentagonal to heptagonal units, delimited by sulci between tumes- cences (Fig. 44). Each tumescence extends around perimeter of unit and abuts adjacent tumescences (Fig. 45). Single, smaller, ir- regularly-shaped tumescence within each perimeter tumescence. Occasionally, central tumescence absent, replaced by depression near center. Tumescences generally gla- brous. Clusters of approximately 12-20 aeropyles distributed over surface of tu- mescences, concentrated near margins. Mi- cropylar plug slightly acentric and amor- phous. Ambrysus (Ambrysus) puncticollis Stal (Figs. 46—48) Ambrysus puncticollis Stal 1876: 143. USA: Texas, Kimble Co., Junction Length, 1.36 + 0.01; width, 0.66 + 0.01; n = 11; oviposited. Overall appearance elongate-oval. Retic- ulation pattern generally consisting of pen- tagonal to heptagonal units, delimited by depressed lines (Fig. 46). Within each unit, 20-80 large aeropyles (aeropyles filled with debris in SEMs) (Fig. 47). Protruberances, tubercles, and papillae lacking. Micropylar plug anterior and amorphous. Anterior pole with pattern less distinct. Exochorion ca. 4.5 thicker than endo- chorion (Fig. 48). Pore canals slightly di- vergent entally and occur at somewhat reg- ular interval. Ambrysus (Ambrysus) spiculus Polhemus and Polhemus (Figs. 49-51) Ambrysus spiculus Polhemus and Polhemus 1981: 400-401. MEXICO: Chihuahua, Rio Concheno Length, 1.10 + 0.01; width, 0.48 + 0.03; n = 3; dissected. 14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 44-49. 44, 45, Ambrysus pulchellus, chorion surface. 46—48, A. puncticollis. 46, 47, Chorion surface. 48, Chorion section. 49, A. spiculus, whole egg. Overall appearance elongate-oval (Fig. lose (Fig. 50). Perimeter of each unit with 49). Reticulation pattern generally consist- 8-16 large, distinct aeropyles; near the cen- ing of pentagonal to heptagonal units, de- ter of unit 1—4 aeropyles. Anterior pole limited by depressed boundaries. Within with reticulation and papillae less evident. each unit, chorion raised and finely papil- Micropylar plug amorphous (Fig. 51). VOLUME 101, NUMBER 1 15 51, Ambrysus spiculus. 50, Chorion surface. 51, Anterior pole with micropylar plug. 52— 54, A. thermarum. 52, 53, Chorion surface. 54, Chorion section. 55, A. tridentatus, chorion surface. Figs. 50-55. 50, Ambrysus (Ambrysus) thermarum La Length, 1.43 + 0.06; width, 0.72 + 0.02; n Rivers = 10; oviposited. (Figs. 52-54) Overall appearance elongate-oval. Retic- Ambrysus thermarum La Rivers 1953b: 1-3. ulation pattern generally consisting of pen- USA: New Mexico, Taos Co., Arroyo Hondo tagonal to heptagonal units, delimited by 16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 56-61. 56, Ambrysus tridentatus, chorion surface. 57, 58, A. woodburyi, chorion surface. 59-61, Gestroiella limnocoroides. 59, Whole egg. 60, Chorion surface. 61, Micropylar plug. double row of rounded to elongate papillae (Fig. 52). Chorionic surface generally cov- ered with papillae. Within each unit formed by reticulation, a single amorphous tumes- cence (Fig. 53). Aeropyles distinct and ran- domly distributed on chorion except on tu- mescence; number 15—30 per unit. Pattern, tumescences, papillae, aeropyles becoming VOLUME 101, NUMBER 1 less distinct toward anterior pole. Micro- pylar plug amorphous. Exochorion slightly thicker than endo- chorion (Fig. 54). Pore canals widest at base. Ambrysus (Ambrysus) tridentatus La Rivers (Figs. 55, 56) Ambrysus tridentata La Rivers 1962: 129— 132. MEXICO: Nuevo Leon, Potrero Redondo Length, 1.14; width, 0.58; n = 1; dissected. Overall appearance elongate-oval. Retic- ulation pattern generally consisting of pen- tagonal to heptagonal units, delimited by depressed boundaries. Chorion domed within each unit (Fig. 55). Aeropyles sub- circular near center of dome, ellipsoid off- center; 20—40 per cell. Chorion smooth; pa- pillae and tubercles lacking (Fig. 56). An- terior pole with domes diminishing in size, aeropyles becoming less evident. Micropy- lar plug amorphous. Ambrysus (Ambrysus) woodburyi Usinger (FISs: 574.9) Ambrysus woodburyi Usinger 1946: 194— 1953 USA: Arizona, Cochise Co., Portal Kenseth 156)25 0:02: width, 0.57 = 0:02; n = 10; oviposited. Overall appearance elongate-oval. Retic- ulation pattern generally poorly-defined. Within each unit formed by reticulation, a single amorphous tumescence (Fig. 57). Each unit with approximately 7—12 large, distinct aeropyles randomly distributed around the tumescence. Chorionic surface generally finely papillose (Fig. 58). Ante- rior pole with tumescences and papillae less developed. Micropylar plug amorphous, acentric. Subfamily Cheirochelinae Montandon 1897b Genus Gestroiella Montandon 1897b Gestroiella limnocoroides Montandon (Figs. 59-61) Gestroiella limnocoroides Montandon 1897b: 371-372. 17 THAILAND: Chiang Mai Prov., Chiang Mai Eength; 3/04 = "0:02: width, 1.35 + 0:02: n = 4; dissected. Overall appearance elliptical with poles slightly acuminate (Fig. 59). Reticulation pattern generally consisting of pentagonal to octagonal units, delimited by low, ele- vated ridges (Fig. 60). Within each unit, 300-800 aeropyles. Micropyles incorpo- rated into low, broad mound at anterior pole (Fig. 61). Other than micropyle, pro- truberances, tubercles, and papillae lack- ing. Subfamily Aphelocheirinae Fieber 1851 Genus Aphelocheirus Westwood 1833 Aphelocheirus femoratus Polhemus and Polhemus (Figs. 62—64) Aphelocheirus femoratus Polhemus and Polhemus 1988: 214-216. THAILAND: Songkhla Prov., Ton Nga Chang National Park Eength> 1213 20-012 width;.0:51' 2-001; n = 10; dissected. General appearance oval and robust with anterior pole slightly truncate (Fig. 62). Re- ticulation pattern generally consisting of pentagonal to heptagonal units, delimited by broad, slightly elevated ridges (Fig. 63). Within each unit, 100-300 aeropyles. Mi- cropyle amorphous (Fig. 64). Chorionic surface immediately surrounding micropyle lacking well-defined surface features. Other than micropyle, protruberances, tubercles, and papillae lacking. Subfamily Laccocorinae Stal 1876 Genus Heleocoris Stal 1876 Heleocoris ovatus Montandon (Figs. 65-67) Heleocoris ovatus Montandon 1897c: 451— 452. THAILAND: Yala Province, Than To, Banglang National Park Length,-1.55. 5 0:01;-widths0740:012n = 9; dissected. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 6 ) 0.3 mm 65 0.5 ae . 63, Chorion surface. 64, Anterior pole with micropylar plug. 65—67, Heleocoris ovatus. 65, Whole egg. 66, C on surface. 67, Micropylar plug in concavity at anterior pole. Overall appearance elongate, parallel- tiform ridges (Fig. 66). Within each unit, sided with rounded ends (Fig. 65). Reticu- 25—75 aeropyles. Chorionic surface gener- lation pattern generally consisting of tetrag- ally granular. Anterior pole with amorphous onal to heptagonal units, delimited by cos- micropylar plug set in acetabular depression VOLUME 101, NUMBER 1 19 Figs. 68-73. 68-71, Limnocoris moapensis. 68, Whole egg. 69, 70, Micropyles. 71, Chorion section. 72, 73, Ilyocoris cimicoides. 72, Whole egg. 73, Anterior pole. (Fig. 67). Pattern in depression becoming tus. Locality and ecological data associated indistinct toward micropyle. with the collection of the adults from which Pending taxonomic revision, this species eggs were obtained are available in Sites et has been determined probably to be H. ova- al. (1997). 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Subfamily Limnocorinae Stal 1876 Genus Limnocoris Stal 1858 Limnocoris moapensis (La Rivers) (Figs. 68-71) Usingerina moapensis La Rivers 1950: 368-373. Limnocoris moapensis: Sites and Willig 1994: 810. USA: Nevada, Clark Co., Moapa Length, 0.98 = 0.01; width, 0.56 + 0.01; n = 8; oviposited. Overall appearance robust, elongate-oval (Fig. 68). Reticulation pattern generally con- sisting of pentagonal to heptagonal units, de- limited by faintly impressed lines. Within each unit, chorion smooth, perforated by 15— 60 aeropyles set in shallow individual sock- ets. Chorion smooth, devoid of papillae, tu- bercles, and protruberances other than mi- cropyle. Anterior pole with micropyle set in concavity, immediately surrounded by mod- ified chorion devoid of pattern and aeropy- les. 2—3 distinct micropyles (vy = 2.8, n = 10) arising from central point, extending outward in arcuate fashion (Figs. 69, 70). Exochorion ca. % as thick as endocho- rion. Pore canals widest at base (Fig. 71). Subfamily Naucorinae Stal 1876 Genus Ilyocoris Stal 1861 Ilyocoris cimicoides (Linnaeus) (Figs. 72-77) Nepa cimicoides Linnaeus 1758: 440. Ilyocoris cimicoides: Stal 1861: 201. Naucoris cimicoides: Rawat 1939: 127, Figs. 1-3. CZECHOSLOVAKIA: southern Bohemia, Veselnad Lu Nic Eensth;, 2:25: = 0.05; width,’0:56"= 0:03: n = 8; dissected. 123— Overall appearance cylindrical and elon- gate with a 45 degree bend near middle (Fig. 72). Anterior pole flattened (Fig. 73), posterior pole rounded. Reticulation gener- ally consisting of pentagonal to heptagonal units, delimited by faintly impressed boundaries (Fig. 74). Within each unit, cho- rion smooth, perforated by 50-90 aero- pyles. Flattened anterior pole with elongate tumescences radiating outward from acen- tric micropylar plug (Fig. 75). Micropylar plug with four micropyles (Fig. 76), each of which is raised slightly above the re- mainder of the plug (Fig. 77). Rawat (1939) described the egg as ap- proximately 2 mm in length and cylindrical with an operculate, recurved end. Lebrun (1960) illustrated the position of the micro- pylar plug on the operculum (although not labeled as such) and Hinton (1981) reported the presence of three to four micropyles. This is the only naucorid documented to have endophytic oviposition (Rawat 1939, Cobben 1968). Genus Pelocoris Stal 1876 Pelocoris femoratus (Palisot de Beauvois) (Figs. 78-81) Naucoris femorata Palisot de Beauvois 18202377 Pelocoris femoratus: Stal 1876: 144. Pelocoris femorata: Torre Bueno 1903: 168-172. Pelocoris carolinensis: Hungerford 1927: 80-82, Plate VI. Pelocoris femoratus: McPherson et al. LOSi7es29 ile USA: Missouri, Boone Co., Columbia Length, 1.17 + 0.01; width, 0.64 + 0.01; n = 10; oviposited. Overall appearance elongate-oval (Fig. 78). Reticulation pattern generally consisting of pentagonal to heptagonal units. Within each unit, irregular and sometimes discontin- uous elongate tumescence approximating boundary (Fig. 79). Anterior pole with mi- cropylar plug set in shallow concavity. Mi- cropylar plug somewhat amorphous and in- consistent in appearance, with micropyles opening laterally (Fig. 80). Number of mi- cropyles indistinct, but apparently 2—3. Exochorion ca. 3.5 X thicker than en- dochorion. Pore canals widest at middle and base (Fig. 81). Using light microscopy, McPherson et VOLUME 101, NUMBER 1 2] 78 0.3 mm Figs. 74-79. 74-77, Ilyocoris cimicoides. 74, Chorion surface. 75, Anterior pole. 76, Micropylar plug. 77, Micropylar opening. 78, 79, Pelocoris femoratus. 78, Whole egg. 79, Chorion surface. al. (1987) reported that the chorion has a DISCUSSION primarily irregular hexagonal pattern and the micropylar plug is situated at the ante- Differences were evident in the chorion rior end. between incompletely developed and well- 22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 80-81. developed eggs. Eggs that were incomplete- ly developed generally appeared to have a single point within each reticulation unit, which was raised and around which the chorion appeared to have flowed down onto the surface of the egg. This ‘poured’ ap- pearance probably represented the site of chorion deposition for each follicular epi- thelial cell. Nonetheless, for dissected eggs, even though we selected eggs from the common oviduct or vagina for descriptions, the possibility exists that egg structure may have continued to develop prior to ovipo- sition. For the 21 species of Ambrysus ex- amined, chorionic sculpturing differs inter- specifically and generally is species-specif- ic. Although these differences were noted, other specific features were common among some of the species. Previously, eggs were described and electron micrographs presented for A. lun- atus (Sites and Nichols 1990), a member of the signoreti group. Other members of the signoreti group represented here are A. in- flatus, A. occidentalis, and A. portheo. Gen- erally, these four species share egg features including an acutely raised, fence-like retic- ulation forming a polygonal pattern, and minute papillae distributed generally over the surface. In addition, two species possess elongate, antheform processes. Eggs of oth- er members of the signoreti group are likely to possess these features. Pelocoris femoratus. 80, Micropylar plug. 81, Chorion section. Eggs of the genus Ambrysus have been reported to have six micropyles (Hinton 1981), and those of an unspecified species from Aruba, Netherlands Antilles, usually have at least five (Cobben 1968). The in- dividual micropylar tubes of Ambrysus are fused into a single, prominent plug (Cobben 1968). Our internal examinations have re- vealed two micropyles in each of three spe- cies of Ambrysus and three in two other species. Although it is likely that intraspe- cific variation exists in micropyle number for species of Ambrysus, as has been ob- served in species of other naucorid genera [e.g., C. hungerfordi (Sites and Nichols 1993)], we have observed only two and three micropyles. Thus, our data do not cor- roborate Hinton’s (1981) report of six nor Cobben’s (1968) report of five or more mi- cropyles for species of Ambrysus. In sharp contrast to Ambrysus, the num- ber of micropyles for species of Limnocoris is clear with external examination because micropylar fusion is minimal. For Limno- coris lutzi and Limnocoris sp. [Ecuador, see Sites (1990)], two micropyles are clearly evident. Of 10 eggs of Limnocoris moap- ensis (La Rivers), eight had three micro- pyles whereas the other two had two mi- cropyles. Previous reports for Limnocoris micropyles are nonexistent. The number of micropyles for species of Pelocoris is unclear and the degree of fu- VOLUME 101, NUMBER 1 sion differs interspecifically. Sites (1991) revealed two partially-fused micropyles for P. poeyi. Surprisingly, for P. femoratus, the micropyle number has not previously been given despite three separate descriptions of eggs [Torre Bueno 1903, Hungerford 1927 as P. carolinensis (see La Rivers 1948b), McPherson et al. 1987]. Pelocoris femora- tus micropyles are fused into a plug [‘mi- cropylar boss’ of Torre Bueno (1903)] sim- ilar to that of Ambrysus. The form of the plug is inconsistent, and a canal leading to a micropylar opening may be observed in some specimens. In the only report for eggs of species of the subfamily Potamocorinae, which is con- sidered by some to represent a distinct fam- ily level taxon (e.g., Stys and Jansson 1988), Cobben (1968) indicated that C. kle- erekoperi has a single micropylar opening with several external mucous projections. SYSTEMATIC VALUE The family Naucoridae is not blessed with even a modicum of somatic characters that varies among the higher taxa that may be used to elucidate systematic relation- ships. The principal characters that have been used for interspecific taxonomic dis- tinctions have been adult male and female external genitalic features. Characters of nymphs and eggs have not been used, al- though Lépez Ruf (1989) suggested that chorionic attributes may be valuable taxo- nomic characters externally at the species level and internally at the generic level. We concur with this assessment. Specifically, intergenerically variable characters include the relative widths of the chorionic inner and outer layers and pore canal configura- tion. Although the number of micropyles does not vary appreciably among these gen- era, the degree of external fusion of the in- dividual micropylar tubes may be a taxo- nomically valuable character at generic or higher levels. External chorionic patterns are quite valuable as an interspecific diag- nostic character in certain genera (e.g., Am- brysus). However, the pattern is invariant 23 among the four species of Limnocoris that we have examined. Thus, the utility of this character in providing systematic resolution appears to be restricted to particular genera. ACKNOWLEDGMENTS We thank D. L. Pinkerton and J. A. White, Electron Microscopy Facility, Col- lege of Agriculture, University of Missouri, for assistance with the electron microscopy and preparation of micrographs; and J. T. Polhemus and Jitka Vilimnova for loan or gift of selected females. We also thank T. J. Henry, J. E. McPherson, and J. T. Polhemus for critical reviews of this manuscript, and T. J. Henry for advice on nomenclature. Funding for RWS was provided in part by MU project #PSSLO232. This is Missouri Agricultural Experiment Station journal se- ries paper No. 12,744. LITERATURE CITED Clarke, FE and E. Baroudy. 1990. Studies on Lacco- coris limigenus (Stal) (Hemiptera: Naucoridae) in Lake Naivasha, Kenya. The Entomologist 109: 240-249. Cobben, R. H. 1968. Evolutionary trends in Heterop- tera: Part I: Eggs, architecture of the shell, gross embryology and eclosion. Centre for Agricultural Publishing and Documentation, Waginengen. Fieber, EF X. 1851. Genera Hydrocoridum secundum ordinem naturalem in familias disposita. Abhand- lung der K6niglische BGhemischen gesellschaft der Wissenschaften in Prag, Series 5, 7: 181-212, 4 plates. 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Six new taxa of Ne- pomorpha from Sulawesi and Mindanao. Notes on Malesian aquatic and semiaquatic bugs (Heterop- tera), VI. Tijdschrift voor Entomologie 139: 161— 174. Palisot de Beauvois, A. M. F. J. 1820. Insectes recueil- lis en Afrique et en Amérique, dans les royaumes d’Oware et de Benin, a Saint-Domingue et dans les Etats-Unis, pendant les années 1786-1797. Fain, Paris 14: 225-240. Polhemus, D. A. 1994. A new species of Aphelochei- rus from Sumatra, and addenda to the world checklist (Heteroptera: Naucoridae). Journal of the New York Entomological Society 102: 74-78. Polhemus, J. T. and D. A. Polhemus. 1981. Three new species of Ambrysus from Mexico (Hemiptera: Naucoridae). Pan-Pacific Entomologist 57: 397— 401. . 1982. Notes on neotropical Naucoridae I. A new species of Ambrysus and review of the genus Potamocoris (Hemiptera). Pan-Pacific Entomolo- gist 58: 326-329 (1983). . 1988. The Aphelocheirinae of tropical Asia (Heteroptera: Naucoridae). Raffles Bulletin of Zo- ology 36: 167-300. . 1994. A new species of Ambrysus Stal from Ash Meadows, Nevada (Heteroptera: Naucoridae). Journal of the New York Entomological Society 102: 261-265. Rawat, B. L. 1939. On the habits, metamorphosis and reproductive organs of Naucoris cimicoides L. (Hemiptera-Heteroptera). Transactions of the Roy- al Entomological Society of London 88: 119-138. Sites, R. W. 1990. Naucorid records from Amazonian Ecuador (Heteroptera: Naucoridae). Florida En- tomologist 73: 334-335. . 1991. Egg ultrastructure and descriptions of nymphs of Pelocoris poeyi (Guérin Méneville) (Hemiptera: Naucoridae). Journal of the New York Entomological Society 99: 622-629. Sites, R. W. and B. J. Nichols. 1990. Life history and descriptions of immature stages of Ambrysus lun- atus lunatus (Hemiptera: Naucoridae). Annals of VOLUME 101, NUMBER 1 the Entomological Society of America 83: 800-— 808. . 1993. Voltinism, egg structure, and descrip- tions of immature stages of Cryphocricos hunger- fordi (Hemiptera: Naucoridae). Annals of the En- tomological Society of America 86: 80—90. Sites, R. W. and M. R. Willig. 1994. Efficacy of men- sural characters in discriminating among species of Naucoridae (Insecta: Hemiptera): multivariate approaches and ontogenetic perspectives. Annals of the Entomological Society of America 87: 803— 814. Sites, R. W., B. J. Nichols, and S. Permkam. 1997. The Naucoridae (Heteroptera) of southern Thai- land. Pan-Pacific Entomologist 73: 127-134. Stal, C. 1858. Bidrag till Rio Janeiro-traktens Hemip- ter-fauna. Konglica Svenska Vetenskaps-Akade- miens Handlingar 2(7): 1—84. . 1861. Nova methodus familias quasdam Hem- ipterorum disponendi. Ofversight af Kongliga Svenska Vetenskaps-Akademiens Forhandlingar 18: 195-212. . 1862. Hemiptera mexicana enumeravit spe- 25 ciesque novas descripsit (continuatio). Stettin En- tomologische Zeitung 23: 437-462. . 1876. Enumeratio Naucoridarum /n Enumer- atio Hemipterorum. Konglica Svenska Veten- skaps-Akademiens Handlingar 14(4): 141-147. Stys, P. and A. Jansson. 1988. Check-list of recent fam- ily-group and genus-group names of Nepomorpha (Heteroptera) of the world. Acta Entomologica Fennica 50: 1—44. Torre Bueno, J. R. de la. 1903. Brief notes toward the life history of Pelocoris femorata Pal. B. with a few remarks on habits. Journal of the New York Entomological Society 11: 166-173. Usinger, R. L. 1946. Notes and descriptions of Ambry- sus Stal with an account of the life history of Am- brysus mormon Montd. (Hemiptera, Naucoridae). The University of Kansas Science Bulletin 31: 185-210. Westwood, J. O. 1833. On the connecting links be- tween the Geocorisae and Hydrocorisae of Latreil- le, or the land and water bug tribes. Magazine of Natural History and Journal of Zoology, Botany, Minerology, Geology and Meterology 6: 228-229. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 26-38 NEW SPECIES OF HYDROPTILIDAE (TRICHOPTERA) FROM THE AMAZON REGION OF NORTHEASTERN PERU S. C. HARRIS AND L. J. DAVENPORT (SCH) Department of Biology, Clarion University, Clarion, PA 16214, U.S.A. (e-mail: harris @mail.clarion.edu); (LJD) Department of Biology, Samford University, Birming- ham, AL 35229, U.S.A. Abstract.—Two new species of Neotrichia, N. orejona and N. tirabuzona, five new species of Oxyethira, O. presilla, O. peruviana, O. vaina, O. picita, and O. hozosa, and one new species of Orthotrichia, O. shimigaya are described from the upper Amazon region in Peru. Key Words: In 1992, Harris and Davenport described five new species of microcaddisflies from the Rio Sucusari and Rio Yanamono in the upper Amazon region of Peru (Fig. | in that paper). The collections were made in 1991 by Davenport during educational expedi- tions to the Explorama Lodge and the Ex- plornapo Camp. Davenport made additional trips to the same region (as described in Har- ris and Davenport 1992) in January of 1993 and 1995. This paper reports on seven new species, two in the genus Neotrichia, one in the genus Orthotrichia, and four in the ge- nus Oxyethira, from the most recent collec- tions. An additional Oxyethira is described from an earlier collection in northern Peru. Morphological terminology follows that of Marshall (1979). Length is measured from the top of the head to the wing tip and is given as a range with more than one specimen. Type material is deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (NMNH), and in the collection of the senior author (SCH). Neotrichia orejona Harris and Davenport, new species (Fig: 1) Diagnosis.—In many respects, this species is similar to N. yanomonoa Harris and Dav- Microcaddisflies, Trichoptera, Hydroptilidae, Peru, Neotropics, new species enport, NV. cayada Harris and N. browni Har- ris. All three share the elongate posterolateral extensions from the ninth abdominal segment and all have a hooklike phallic apex. The new species is readily separated by the triangular inferior appendages, the bifid subgenital plate and the serrated phallic processes. Male.—Length 1.3 mm. Antenna with 18 segments. Brown in alcohol. Abdominal seg- ment VIII annular. Segment [X in lateral view with narrow, elongate process from postero- lateral margin; in ventral view with elongate, narrow lobes laterally, mesally with pair of thin, elongate processes each bearing small seta at apex. Segment X fused with IX, in dorsal view deeply incised mesally, pair of small setiferous lobes anteriorly. Inferior ap- pendages triangular in lateral view, apex trun- cate bearing numerous setae; in ventral view rectanguloid, curving mesad. Subgenital plate beaklike in lateral view with narrow seta- bearing process dorsally; wide basally in ven- tral aspect with bifid processes, narrowing posteriorly. Phallus tubular, apex divided into two flattened serrate processes, ejaculatory duct protruding subapically, thin paramere encircling shelf near midlength. Female.—Unknown. Type material.—Holotype, ¢. Peru, Lor- VOLUME 101, NUMBER |! Zi Fig. 1. Neotrichia orejona, male genitalia. A, Lateral. B, Ventral. C, Dorsal. D, Phallus, dorsal. eto, edge of Rio Sucusari backwater, ad- Neotrichia tirabuzona Harris and joining Explornapo Camp, 16 January Davenport, new species 1993, L. J. Davenport (NMNH). (Fig. 2) Etymology.—Named for the Orejone in- Diagnosis.—This new species falls with- dians which live in the area. in the caxima group, as established by Mar- 28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Neotrichia tirabuzona, male genitalia. A, Lateral. B, Ventral. C, Dorsal. D, Phallus, dorsal. shall (1979), with greatest similarity to N. rotundata Flint and N. dientera Harris. The hooklike inferior appendages and _ spiral process from the phallus apex are charac- teristic for N. tirabuzona. Male.—Length 1.5-1.7 mm. Antenna with 18 segments. Brown in alcohol. Ab- dominal segment VIII annular. Segment IX in lateral view square, curving ventrally to posteroventral projection; in ventral view square, emarginate on posterior and anterior margins. Segment X lobate in lateral view; VOLUME 101, NUMBER 1 in dorsal view partially fused with IX at narrow base, widening distally with trun- cate incision apically. Inferior appendages hook-shaped in lateral view; in ventral view square and widely separated, sclerotized bands basally, posteriorly, and mesally. Subgenital plate a narrow shelf in lateral view; in ventral view rounded distally, pair of elongate setae mesally. Phallus wide at base and at rounded apex, spiral process apically, ejaculatory duct thin and elongate, protruding apically, elongate paramere en- circling shaft beyond midlength. Female.—Unknown. Type material—Holotype, 6. Peru, Lor- eto, edge of Rio Sucusari backwater, ad- joining Explornapo Camp, 16 January 1993, L. J. Davenport (NMNH). Paratypes, Peru, same data as holotype, 7 6 (NMNH, SCH). Etymology.—Spanish, corkscrew, refer- ring to the spiral process from the phallus apex. Orthotrichia shimigaya Harris and Davenport, new species (Fig. 3) Diagnosis.—The genus Orthotrichia is represented by six species in the Nearctic region, with two, O. aegerfasciella (Cham- bers) and O. cristata (Morton), extending into the Neotropical region. Orthotrichia shimigaya is the first species of the genus to be reported exclusively from South America. The species is easily recognized by the structure of the inferior appendages. Male.—Length 2.5—2.6 mm. Antenna with 32 segments. Brown in alcohol. Ab- dominal segment VII with elongate poster- omesal process from venter. Segment VIII annular. Segment IX reduced ventrally to narrow bridge, a rounded lobe posteroven- trally; in dorsal view incised posterolater- ally, mesally a truncate lobe. Segment X in lateral view divided into pair of thin, elon- gate processes, turned laterad at apex; in ventral view, narrowly incised on postero- lateral margin, left lobe wider and more truncate than right. Inferior appendages in 29 lateral view short and triangular; in ventral view asymmetrical, left appendage with pair of thin, seta-bearing processes from posterolateral margin, right appendage nar- rowing mesad and curving downward, sin- gle seta-bearing process from posterolateral margin. Subgenital plate in lateral view a narrow shelf; in ventral view tonguelike, pair of setae from rounded, posterior mar- gin. Phallus tubular, apical half with ring- like crenulations, tipped with pair of narrow lateral lobes, paramere encircling shaft at midlength. Female.—Unknown. Type material—Holotype, ¢. Peru, Lor- eto, small stream just outside grounds of Explorama Inn, 20 January 1995, L. J. Dav- enport (NMNH). Paratype. Peru, Loreto, backwater creek at outlet of Lake Shimigay, ca. 2 km. upstream Rio Napo from mouth of Rio Sucusari, 15 January 1993, L. J. Davenport, | 6 (NMNH). Etymology.—Named for Lake Shimigay, one of the type localities for the species. Oxyethira presilla Harris and Davenport, new species (Fig. 4) Diagnosis.—This new species is most similar to O. rareza Holzenthal and Harris, with which it shares the asymmetrical gen- italic features. The multiple processes at the apex of the phallus and the long looping processes from the venter of segment IX will readily identify O. presilla. Male.—Length 2.2 mm. Antenna with 38 segments. Brown in alcohol. Abdominal segment VII annular, lacking posteromesal process from the venter. Segment VIII ta- pering posteriorly, deeply incised on pos- terior margin in ventral and dorsal views. Segment IX complex, elongate anteriorly and tapering posteriorly with several asym- metrical processes, ventrally divided into two processes, lowermost upturned at mid- length and slightly widening, uppermost process thin, posteriorly curving into a loop; dorsal process narrow anteriorly, wid- ening to transverse plate at midlength; in 30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Orthotrichia shimigaya, male genitalia. A, Lateral. B, Ventral. C. Dorsal. D, Phallus, dorsal. ventral view deeply incised, right lateral margin with two processes, apical process narrow, subapical process wider and pro- jecting more mesad, left lateral margin di- vided into two elongate thin processes. Ter- gum X apparently fused with LX as apex of rectangular plate on IX. Inferior append- ages and subgenital plate not evident. Phal- lus wide at base, narrowing at midlength, apical portion widening with several scler- VOLUME 101, NUMBER 1 31 Fig. 4. Oxyethira presilla, male genitalia. A, Lateral, left side. B, Lateral, right side. C, Ventral. D, Dorsal. E, Phallus, dorsal. FE Phallus, lateral. Ww i) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON otized processes on lateral margins, ejacu- Type material.—Holotype, d. Peru, Lor- latory duct protruding distally as sclero- eto, Yanamono Creek at jungle’s edge, near tized, sinuate process. Female.—Unknown. Explorama Lodge, 12 January 1995, L. J. Davenport (NMNH). VOLUME 101, NUMBER 1 Etymology.—Spanish, loop, refering to the distinctive process from segment IX. Oxyethira peruviana Harris and Davenport, new species (Fig. 5) Diagnosis.—This species appears to be most similar to O. spissa Kelley, a mem- ber of the pallida group of Kelley (1984). Both species have a prominent posterolat- eral process from segment IX and incon- spicuous subgenital plate, but O. peruvi- ana differs in the elongate processes from the phallus apex and the longer inferior appendages. Male.—Length 2.3 mm. Antenna with 29 segments. Brown in alcohol. Abdomi- nal segment VII annular with short pos- teromesal process from the venter. Seg- ment VIII annular in lateral view; deeply incised posteriorly in ventral view; round- ed posteriorly in dorsal view. Segment IX triangular in lateral view, greatly reduced dorsally, acute, downturned curved pro- cess posterolaterally, posteroventrally with short, triangular process; in ventral view, posterior margins with three pro- cesses, lateralmost thin and acute, ventro- mesal processes triangular, dorsomesal processes tapering and angled inward. Segment X a short, membranous lobe; in dorsal view wide and rounded posteriorly. Inferior appendages short and rounded in lateral view, short spine from venter; in ventral view fused mesally, triangular lat- erally with small bilobed sclerite from mesal margins. Subgenital plate rectan- guloid and slanted posteroventrally in lat- eral view; in ventral view a narrow, trans- verse band; lacking bilobed process. Phal- lus tubular, widening at apex which bears an elongate paramere subapically and short, transverse process at apex; ejacu- latory duct bifid at apex. Female.—Unknown. Type material.—Holotype, ¢. Peru, Lor- eto, tributary to Rio Yanamono at Explor- ama Lodge, 11 March 1991, L. J. Daven- port (NMNH). 33 Etymology.—Named for the country of Peru. Oxyethira vaina Harris and Davenport, new species (Fig. 6) Diagnosis.—This new species is most similar to O. orellanai Harris and Dav- enport, a member of the Tanytrichia sub- genus of Kelley (1984), which was also collected at the same locality on the Rio Sucusari. The new species is separated by the shorter anteroventral extension of seg- ment IX and the triangular inferior ap- pendages, which in ventral view are fused mesally. Male.—Length 2.3 mm. Antenna with 28 segments. Brown in alcohol. Abdomi- nal segment VII annular with short pos- teromesal process from the venter. Seg- ment VIII tapering posteroventrally in lat- eral view; in ventral view deeply incised mesally on posterior margin; posterior margin of dorsum with mesal truncate in- cision. Segment IX reduced posterodor- sally to narrow bridge, narrowing and ex- tending anteriorly through segment VII. Tergum X lobate, membranous. Inferior appendages short and triangular in lateral view; fused in ventral view, round mesal incision creating thin triangles laterally, stout seta from apex, dorsal lobes along mesal margins. Subgenital plate in lateral view a narrow shelf with bilobed process curving over it posteriorly; in ventral view a narrow rectangle with thin lateral arms angled mesad, bilobed processes widely separated with setae at tips. Phal- lus tubular with lateral sheath, narrow sclerite running contiguous with ejacula- tory duct. Female.—Unknown. Type material—Holotype, d. Peru, Lor- eto, edge of Rio Sucusari backwater, ad- joining Explornapo Camp, 16 January 1993, L. J. Davenport (NMNBH). Etymology.—Spanish, sheath, referring to the lateral ribbonlike band of the phallus. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Oxyethira vaina, male genitalia. A, Lateral. B, Ventral. C, Dorsal. D, Phallus, lateral. VOLUME 101, NUMBER 1 Oxyethira picita Harris and Davenport, new species (Fig. 7) Diagnosis.—Although this species ap- pears to be a member of the Tanytrichia subgenus of Kelley (1984), there is some resemblance to O. zilaba (Mosely) of the subgenus Loxotrichia. The new species is distinguished by the elongate subgenital plate and the pair of small lateral spines on the tenth tergite. Male.—Length 2.3 mm. Antenna with 26 segments. Brown in alcohol. Abdominal segment VII annular with short postero- mesal process from the venter. Segment VIII tapering posteroventrally to rounded apex, short process dorsolaterally; in ven- tral view deeply incised posteriorly; in dor- sal view also deeply incised with three acute processes mesally. Segment IX ex- tending anteriorly into segment VI, poste- riorly short and reduced dorsally to thin process. Segment X lobate in lateral view; in dorsal view rectanguloid with rounded apex, pair of short spines posterolaterally. Inferior appendages in lateral view elongate and tapering posteriorly, apex with ventral spine and dorsal seta-bearing process; in ventral view fused posteriorly, apex with lateral seta-bearing processes and median truncate process, heavy seta between pro- cesses, lateral margins gently emarginate. Subgenital plate curving anteroventrally, posteriorly elongate and narrowing to acute apex, transverse bilobed process thin and elongate; in dorsal and ventral views divid- ed at base into two elongate processes which narrow and cross apically, bilobed process thin, diverging distally. Phallus tu- bular, small lateral spines below midlength, widening apically, pair of lateral processes which are curved distally, ejaculatory duct between the two processes. Female.—Unknown. Type material.—Holotype, d. Peru, Lor- eto, edge of Rio Sucusari backwater, ad- joining Explornapo Camp, 16 January 1993, L. J. Davenport (NMNH). 35 Etymology.—Spanish, small sharp point, referring to the small spines of the tenth tergite. Oxyethira hozosa Harris and Davenport, new species (Fig. 8) Diagnosis.—This species is closely relat- ed to O. scaeodactyla Kelley, particularly in the structure of the phallus and the in- ferior appendages. The new species differs in the acute, distal point of the subgenital plate, the short bilobed process, and the more complete ninth segment. Male.—Length 2.4 mm. Both antenna broken. Brown in alcohol. Abdominal seg- ment VII annular with short, posteromesal process from the venter. Segment VIII rounded posteriorly in lateral view; in ven- tral view, deep mesal incision; emarginate dorsally. Segment IX tapering anteriorly, a narrow band posterodorsally, narrow pro- cess posteroventrally; ventrally with lateral margins produced into sharp points which extend beyond VIII, pair of fingerlike pro- cesses mesally; dorsally fused posteriorly with X. Segment X a short membranous lobe in lateral view; dorsally square in shape with posterior margin truncate. Infe- rior appendages short and truncate; in ven- tral view, incised mesally and fused, nar- rowing posterolaterally and bearing stout seta. Subgenital plate in lateral view strong- ly curving ventrad, apex with acute apical point; in ventral view wide with lateral edg- es rounded, mesally with knoblike process; bilobed processes short in lateral view; in ventral view widely separated and sinuate. Phallus short, distally narrowing to con- spicuous hook, ejaculatory duct sinuate and enclosed within membranous lobe. Female.—-Unknown. Type material—Holotype, d. Peru, Lor- eto, Rio Yanamono just below Explorama Lodge, 10 January 1993, L. J. Davenport (NMNH). Etymology.—Spanish, sicklelike, refer- ring to the distinctive phallus. Bigs 7 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Oxyethira picita, male genitalia. A, Lateral. B, Ventral. C, Dorsal. D, Phallus, dorsal. VOLUME 101, NUMBER 1 PN xt ott eds Fig. 8. Oxyethira hozosa, male genitalia. A, Lateral. B, Ventral. C, Dorsal. D, Phallus, dorsal. ACKNOWLEDGMENTS Thanks to the staff at Explorama Tours in Iquitos for their help in organizing the trips to Peru; to the two manuscript review- ers; and to Sarah Harris who typed most of the original draft of the manuscript. LITERATURE CITED Harris, S. C. and L. J. Davenport. 1992. New species of microcaddisflies from the Amazon region, with especial reference to northeastern Peru (Trichop- tera: Hydroptilidae). Proceedings of the Entomo- logical Society of Washington 94: 454—470. Kelley, R.W. 1984. Phylogeny, morphology and clas- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sification of the micro—caddisfly genus Oxyethira — Marshall, J. E. 1979. A review of the genera of the Eaton (Trichoptera: Hydroptilidae). Transactions Hydroptilidae (Trichoptera). Bulletin of the British of the American Entomological Society 110: 435— Museum (Natural History), Entomological Series 463. 39: 135-239. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 39-56 AN EXAMINATION OF THE NORTH AMERICAN APHID SPECIES CURRENTLY PLACED IN OVATUS VAN DER GOOT (HEMIPTERA: APHIDIDAE) WITH THE DESCRIPTION OF A NEW GENUS ANDREW S. JENSEN AND MANYA B. STOETZEL Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, Bldg. 046, BARC-W, Beltsville, MD 20705, U.S.A. (e-mail: ajensen@ sel.barc.usda.gov; mstoetze @sel.barc.usda.gov) Abstract.—The current placement in Ovatus of four endemic North American aphid species is critically examined. Results of cladistic analyses clearly indicate that these four species are not congeneric with Palearctic Ovatus and that they are more closely related to Myzus Passerini. Cladistic evidence is provided to justify the erection of a new genus, Abstrusomyzus n. gen., for the four endemic North American species formerly placed in Ovatus. This resulted in four new combinations: Abstrusomyzus leucocrini (Gillette & Palmer) n. comb., Abstrusomyzus phloxae (Sampson) n. comb., Abstrusomyzus reticulatus (Heie) n. comb., and Abstrusomyzus valuliae (Robinson) n. comb. Information on each of the four species is compiled and presented. Illustrations are provided, along with a key to the species of the new genus and notes on the single remaining species of Ovatus in North America, Ovatus crataegarius (Walker). Key Words: Most aphids are extremely host specific, feeding on one or a few species of plants that are usually closely related (Eastop 1973). Many of the most pestiferous aphids known differ dramatically from this normal pattern by being polyphagous on plants in widely divergent families. Well-known po- lyphagous pests include the green peach aphid [Myzus persicae (Sulzer)], the potato aphid [Macrosiphum euphorbiae (Thomas)], and the cotton aphid (Aphis gossypii Glov- er). These aphids can cause serious problems even when they feed on a crop in low num- bers, since they can transmit plant viruses between their phylogenetically disparate host plants. Very few polyphagous aphids are not pests. Given the fact that many of the most serious pest aphids are polypha- gous, it is important to study newly-discov- ered cases of polyphagy in aphids. As po- polyphagy, Myzus, Ovatus, new genus, cladistics lyphagous aphids are likely to some day be- come pests, prior knowledge of their biolo- gy, taxonomy, and evolution will facilitate an effective reaction to their emergence as pests. Ovatus phloxae (Sampson) is one of the few polyphagous aphids that so far is not a pest. It was described in 1939 from Phlox subulata from Berkeley, California. Since then it has been found on eighteen plants in fourteen families (Table 1). It has been col- lected in most regions of the United States and probably occurs in most temperate parts of North America. Most plants on which this aphid has been found have a basal rosette of leaves, or tend to be prostrate. The aphid usually lives in the crown or on the lower leaves. It is possible that this species is more habitat specific than host specific, much as in Rhopalosiphum nymphaeae (L.), which 40 Table 1. Host plants and distributions for five species heretofore placed in Ovatus. Aphid Species Ovatus crataegarius (Walker) Ovatus leucocrini (Gillette & Palmer) Ovatus phloxae (Sampson) Ovatus reticulatus Heie Ovatus valuliae (Robinson) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Host Plants Distribution Rosaceae Almost Crataegus spp. worldwide, Cydonia spp. with a Malus spp. Palearctic Lamiaceae origin Mentha spp. some other Lamiaceae Liliaceae Colorado Leucocrinum montanum Apocynaceae Canada: Apocynum androsaefolium British Apocynum sp. Columbia Asteraceae Achillea sp. U.S.A:: Agoseris sp. California Centaurea sp. Colorado Brassicaceae Maryland Capsella hursa-pastoris Mississippi Caryophyllaceae Nebraska Cerastium vulgatum Oregon Stellaria crispa Pennsylvania Cyperaceae Utah Carex densa Virginia Fabaceae Trifolium sp Hydrophyllaceae Phacelia nemoralis Liliaceae Seilieske Plantaginaceae Plantago major Polygonaceae Polygonum paronychia Ranunculaceae Ranunculus sp. Polemoniaceae Phlox subulata Rubiaceae Galium sp. Violaceae Viola sp. Oxalidaceae North Carolina Oxalis ?stricta Rosaceae Manitoba Fragaria vesca feeds widely on aquatic and semiaquatic plants. If this were true, O. phloxae could emerge at any time as a pest of a crop with an appropriate growth habit. Therefore a study of North American Ovatus was un- dertaken to provide needed information re- garding O. phloxae and three closely related endemic North American aphids currently placed in Ovatus. The relationship of these four species to Palearctic Ovatus is exam- ined using cladistics, and a new genus is proposed for them based on these analyses. VOLUME 101, NUMBER 1 MATERIALS AND METHODS Most specimens studied are housed in the National Collection of Aphidoidea (USNM, located at the Systematic Entomology Lab- oratory, USDA, Beltsville, Maryland, USA). Others were obtained on loan from The Nat- ural History Museum, London (BMNH); Agriculture Canada, Vancouver, British Co- lumbia (UBC); University of California, Berkeley (UCB); and Oregon State Univer- sity, Corvallis (OSU). Aphids were mounted on microscope slides in Canada balsam or Hoyer’s medium, and observed under phase contrast micros- copy. Terminology follows Miyazaki (1987). Drawings were made by the first author us- ing a camera lucida. Cladistic analyses were performed using PAUP 3.1.1 (Swofford 1993) and MacClade 3.01 (Maddison and Maddison 1992) on 23 taxa using 25 characters. Apterous and alate viviparae were included in the character analysis, supplemented by one character of the first instar nymph. All characters were treated as unordered. Ovatus van der Goot Ovatus has most recently (Remaudiére and Remaudiére 1997) been used for four- teen Myzus-like aphids, which generally mi- grate between Pomoidea and various Lami- aceae, or are monoeceous on either of these groups. Nine of the fourteen species fit this pattern of host plant association. One excep- tion is the single member of the subgenus Ovatoides, Ovatus (Ovatoides) inulae (Walker). This species feeds on Asteraceae and differs morphologically from most Ova- tus by the nearly complete lack of spinula- tion on the head and first two antennal seg- ments, and the long, setose ultimate rostral segment. The other four species that do not conform with the Pomoidea/Lamiaceae host plant association are the four endemic North American species [Table 1, excluding Ova- tus crataegarius (Walker)]. These species feed on herbs from a diverse set of plant families, and none of them is known from 41 Pomoidea or Lamiaceae. Morphologically these species differ from most Palearctic Ovatus by the dark bars and blotches on the abdominal tergum of the alate vivipara (Figs. 17, 18), first tarsal chaetotaxy of 3,3,2, and the peculiarly shaped siphunculus (Figs. 9-12). The first to place one of these species in Ovatus was Hille Ris Lambers (1966), who transferred Phorodon phloxae to Ovatus without any explanation. Heie (1972) de- scribed Ovatus reticulatus, and Eastop and Hille Ris Lambers (1976) transferred Myzus leucocrini Gillette and Palmer and Myzus valuliae Robinson without explanation. The obviously close relationship among _ these four species dictated that they be placed to- gether in the same genus, but their morpho- logical and biological differences from Pa- learctic Ovatus causes doubt about their cur- rent generic placement. CLADISTIC ANALYSES This study attempts to determine whether the four North American species currently placed in Ovatus will cluster together, and whether they will form their own group, form a part of Ovatus, or fall within another genus. Doing this requires that the cladistic analysis include several species from one or more genera to which the species in question might be more closely related than they are to Ovatus. These cladistic analyses are meant to explore the proper generic place- ment of the four endemic North American species currently placed in Ovatus and are not meant to resolve species-level issues for the other aphids included in the analysis. Choice of appropriate comparative groups for a test of the proper placement of the four endemic North American species is partic- ularly troublesome in the case of Ovatus for at least three reasons. First, the fact that the analysis deals with aphids presents problems in itself (see Jensen 1997 for some discus- sion of this issue). Aphids are highly pro- genetic sensu Gould (1977), and because of their conservative morphology most genera lack apomorphic characters. Instead, there is 42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON essentially a large pool of characters that oc- cur across a wide range of aphid taxa, and many genera are recognized on the basis of a unique combination of these characters. Aphid genera are also frequently defined by the absence of one or more characters typi- cal in genera to which they are most similar morphologically. These facts, in combina- tion with the lack of work on the evolution of the commonly used aphid characters, make deductions about relatedness difficult. The second problem that Ovatus poses is biogeographical. As currently understood, the genus is Holarctic, with ten species na- tive to Eurasia, and four native to North America. Perhaps, then, comparative species should be drawn from related genera on both continents. Nearly all species of Myzina, the subtribe to which Ovatus belongs, are en- demic to Eurasia, with the subtribe’s greatest diversity in southern and eastern Asia. Therefore, choosing Eurasian species for comparison is easy. But examination of the native aphid fauna of North America shows that there are only a handful of Myzina na- tive to this continent. These native species are mostly specialized moss and sedge feed- ers such as Myzodium Borner and Caroli- naia Wilson. There are few other native North American Myzina besides the four pu- tative Ovatus, and several species of Hyalo- myzus. In other work (Stoetzel, Miller and Jensen, in preparation), evidence has been found for the monophyly of Hyalomyzus Richards of North America. This leaves few North American groups to which the four Ovatus could belong. Thirdly, the four North American species currently placed in Ovatus are extremely similar and possess some unique character- istics. For example, they all have siphunculi more or less cylindrical, but slightly expand- ed apically (Figs. 9-12). Two of the species also tend to have more than three pairs of setae on the cauda, whereas most Myzina have two or three pairs. The four North American species also have a distinctive pat- tern of pentagonal or hexagonal reticulation on their terga in the apterous viviparous fe- male. These peculiarities make them isolated morphologically from most of the Myzina and make the choice of appropriate species for the current analysis more difficult. Myzus has for a long time been used for a great diversity of species of Myzina. This can be illustrated by examining the list of Eastop and Hille Ris Lambers (1976). They list 53 valid species of Myzus, and 40 others that were described in Myzus, but subse- quently transferred to 27 other genera. The range of species included in the genus has varied from one author to another during this century. In North America most workers have dealt with the genus in a broad sense (Mason 1940, Palmer 1952). Mason states that Myzus can be recognized among the Macrosiphina in his tribe Aphidini by the convergent frontal tubercles. Thus in his re- vision of the genus, Mason (1940) included 20 species that are today scattered among six genera. Despite all this modification or re- striction of the definition of the genus, My- zus still contains a wide array of species and is quite likely polyphyletic. The diversity of species in Myzus and the lack of another ge- nus that is clearly related to the four species in question indicate that Myzus is the best choice for use in this analysis. Myzus pro- vides a wide array of species to which the four endemic North American species might prove to be closely related. SPECIES INCLUDED IN THE ANALYSES Included in the analysis were Ovatus phloxae and three very similar North Amer- ican species, O. leucocrini, O. reticulatus, and O. valuliae. For purposes of testing their relationship to Palearctic Ovatus, six other Ovatus species were studied, including Ova- tus (Ovatoides) inulae, the only member of its subgenus. These were chosen based on their availability in the collection of the USNM and material obtained on loan. Thus ten of the world’s fourteen Ovatus species were analyzed. Adequate material of the oth- er Eurasian species currently placed in Ova- tus was not available. Ten species of Myzus were included, VOLUME 101, NUMBER 1 Mable 2: polymorphism, and a + indicates a 0/2 polymorphism. SOS SSS 999°C Ss sss 5 55 5 55 2 )2)5 eriobotryae Jussiaeae mitchellensis ascalonicus cerast certus cymbalariae dycei . hemerocallis lythri ornatus persicae varians crataegarius glechomae INSItUS inulae leucocrini malisuctus mentharius phloxae reticulatus valuliae Val aba alae 123456789012345678 000000000000001110 00000000010000111 000111010000101110 +20170101010101110 000000000000000000 000110041010101010 120100001110001110 000040001110100110 0201100+1000100010 000010000000100010 100110000010110110 000100021010001010 000000001010100110 001100000100000110 100100001000010110 001100001100000110 011101101000110010 031100101011102001 0111000010101001T0 1001000000001t0110 00010000111110201t 030100021011102000 1301000010111120C0 144444 901234 ALAMOS A +++000 000000 000100 010000 010110 000011 000000 000110 000100 000110 Ab abal ab(O)al 110001 ial avab(oyal ah ab lays 000100 Oalalenlek@ 111001 O+O0O0t 2??20? 000100 Data matrix used in the cladistic analyses. Question marks indicate missing data, + indicates a 0/1 = SP rPRPrPODOOF SS je representing three of four subgenera, and 16% of the world fauna. These species were chosen to represent all possible sub- genera, and are all the species of which adequate material was available. Three outgroup species were chosen from North American Hyalomyzus. The dataset is shown in Table 2. CHARACTERS Apterous viviparae Head capsule: 1. Anterior setae on dorsum of head cap- sule: less than 0.5 times basal width of antennal segment III (0); more than 0.5 times basal width of antennal segment mE by: Ornamentation on dorsum of head cap- sule: spinules or nodules present anteri- orly and along margins, smooth in mid- dle posteriorly (0); without spinules or nodules, sometimes with wrinkles (1); spinules or nodules entirely covering NS) dorsum of head (2); dorsum of head with posterior surface in middle with polyg- onal reticulation (3) (Figs. 5—8). 3. Venter of head capsule adjacent to eyes: spinulose (0); smooth (1). 4. Ventral tubercles at rear of head capsule: present (0); absent (1). Antenna: 5. Ratio of processus terminalis to the base of antennal segment VI: greater than 3 (0); less than 3 (1). 6. Venter of antennal segment I: bumpy (0); smooth (1). Mouth Parts: 7. Pairs of setae on rostral segment III: al- most always 2 (0); more than 2 (1). 8. Ultimate rostral segment: longer than hind tarsal segment II (0); shorter than hind tarsal segment II and without ac- cessory setae (1); shorter than hind tarsal segment II and with accessory setae (2). 9. Number of setae on ultimate rostral seg- 44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON {77} 53 M. ascalonicus 2} (77) M. cymbalariae 62 M. cerasi M. lythri M. hemerocallis M. certus M. persicae O. leucocrini O. reticulatus — O. valuliae O. phloxae M. ornatus (77} M. varians M. dycei O. malisuctus {2} {65} {53} O. glechomae 51 @ O. crataegarius {52} id O. insitus oe O. inulae O. mentharius 1 H. mitchellensis {2} H. jussiaeae H. eriobotryae Fig. 1. Tree number one of two equally parsimonious trees found using the heuristic search option in PAUP with successively weighted data (weighted consistency index: 0.67; retention index: 0.84; and rescaled consis- tency index: 0.57). This tree was also one of 135 trees found using unweighted data (unweighted consistency index: 0.47; retention index: 0.64; and rescaled consistency index: 0.30). Numbers below some branches indicate branch support values from decay analyses, plain numbers indicating unweighted branch support, and numbers in braces indicating rescaled branch support from the weighted data (Bremer 1994). Numbers above branches display results of bootstrap analyses, plain numbers indicating results using unweighted data, and numbers in braces representing weighted data. ment: O—3 (usually 2) or rarely 4 acces- irregular closed shapes with smooth sory setae (0); usually 3 or more (rarely space’ between ithem® (0) (ess): 2) accessory setae (1). smooth (1); pentagonal or hexagonal (2) (Fig. 4). 13. Spinal tubercles on abdominal tergite VII: usually present (0); absent (1). 14. Setae on abdominal tergite VIII: much shorter than basal width of antennal segment III (0); subequal to basal width of antennal segment IIIf (1). Abdomen: 15. Shape of siphunculi: cylindrical or ta- 12. Tergum patterning or sculpturing: pering (0); distinctly swollen (1); very maze-like with no distinct polygons, or slightly expanded near apex (2). Legs: 10. Dorsal base of hind tibia: smooth or slightly wrinkled (0); distinctly sca- brous or imbricated (1). 11. First tarsal segments chaetotaxy for- mula: 3—3-3 (0); 3—3-—2 (1). VOLUME 101, NUMBER 1 Fig. 2. O. leucocrini O. reticulatus O. valuliae O. phloxae M. certus M. persicae M. ascalonicus M. cymbalariae M. cerasi M. lythri M. hemerocallis M. ornatus M. varians M. dycei O. malisuctus O. glechomae O. crataegarius Tree number one of four equally parsimonious trees found using the heuristic search option in PAUP with successively weighted data (weighted consistency index: 0.77; retention index: 0.84; and rescaled consis- tency index: 0.65). This tree was one of 14 trees found using unweighted data (unweighted consistency index: 0.47; retention index: 0.64; and rescaled consistency index: 0.30). Numbers below some branches indicate branch support values from decay analyses, plain numbers indicating unweighted branch support, and numbers in braces indicating rescaled branch support from the weighted data (Bremer 1994). Numbers above branches display results of bootstrap analyses, plain numbers indicating results using unweighted data, and numbers in braces representing weighted data. 16. Apical spinulation of cauda: entire, without blank spaces dorsally (0); re- duced with blank spaces dorsally (1). 17. Abdominal tergum pigmentation: pig- mented, with complete or nearly com- plete dorsal shield (0); pale, or with only cross bands on tergites VII and/or VI. 18. Setae on cauda: 2 or 3 lateral pairs (0); 4 or more lateral pairs (1). Alate viviparae Antenna: 19. Secondary rhinaria on antennal seg- ment V: absent (0); present (1). 20. Secondary rhinaria on antennal seg- ment IV: absent (0); present (1). 21. Secondary rhinaria on antennal seg- ment III: restricted to approximately half the circumference of the segment (O); distributed around the entire cir- cumference of the segment (1). Mouthparts: 22. Ornamentation of lateral part of man- dible (base of mouthparts): spinulose or scabrous (0); smooth (1). Abdomen: 23. Lateral abdominal tubercles: present in at least some specimens (0); absent (1). 24. Pigmented abdominal patch or bands: present (0); absent (1). Nymphs Legs: 25. Apical spinulation of hind tibia: present (0); absent (1). CLADISTICS RESULTS AND DISCUSSION The full data set was analyzed using the heuristic search option in PAUP. One hun- 46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON [ 0.026 mm Figs. 3, 4. dred iterations of random addition sequence were performed, finding 135 equally parsi- monious trees of length 91 steps. These had a consistency index (CI) of 0.47, retention index (RI) of 0.64, and rescaled consistency index (RC) of 0.30. One of these trees is presented in Fig. 1, with numbers below a few branches indicating results of a decay analysis (see Bremer 1994), and numbers above branches indicating the level of boot- strap support from 100 replications of heu- ristic bootstrap searching. Only branches supported by more than 50% of the boot- strap trees are labeled in Fig. 1. In all 135 trees, the group of four North American Ovatus is placed among the Myzus species, separate from other Ovatus, and supported by more than 50% of the bootstrap trees. Old World Ovatus are placed as close rel- atives to the outgroup Hyalomyzus. The close relationship between Ovatus and Hy- alomyzus has been discussed by Eastop (1966), Nielsson and Habeck (1971), and Blackman and Eastop (1994). The four North American Ovatus formed a clade in all 135 trees and were often placed with M. certus and M. persicae, two | 0.024 mm Dorsal sculpturing. 3, Ovatus crataegarius. 4, Abstrusomyzus leucocrini. species with swollen siphunculi that repre- sent the subgenus Nectarosiphon. Moving the four North American species from with- in Myzus to the more basal parts of the tree along with the other Ovatus caused an in- crease in tree length of three or four steps, depending on the branch to which they were attached. For example, moving the branch composed of these four species to become the terminal Ovatus, along with Ovatus malisuctus (Matsumura), caused an increase in tree length of three steps. But it is clear from the bootstrap results that the most strongly supported clade of more than two species is that composed of the four North American species related to O. phlox- ae, henceforth referred to as the “‘phloxae group.” A successive weighting procedure using the default settings in PAUP was conduct- ed, finding two of the 135 trees found using unweighted data, with the following weighted statistics: length, 22, 140; CI, 0.67; RI, 0.84; RC, 0.57. The tree shown in Fig. 1 is one of these trees. Bootstrap and decay analyses were performed using the weighted dataset, and their results are dis- VOLUME 101, NUMBER 1 played in Fig. 1. Decay indices in braces in Fig. 1 are rescaled (Bremer 1994). The two successive weighting trees even more strongly support the monophyly of the phloxae group and its separation from Pa- learctic Ovatus. Monophyly of the phloxae group was supported in 78% of the boot- strap trees and through 2 rescaled steps of decay. The results of this analysis clearly indi- cate that the phloxae group does not belong to Ovatus. Consistent placement of the phloxae group within the clade of Myzus species indicates that they are more closely related to Myzus than they are to Ovatus of the Palearctic. It should also be pointed out that O. malisuctus is placed among the My- zus Clade in some of the 135 trees, and may be more closely related to Myzus than are the other Ovatus. Further studies focusing on Palearctic aphids in and related to Ova- tus will be required to finalize the classifi- cation of this and other problematic species not included in this study. The unique characters of the four North American “‘Ovatus’’ species (peculiar si- phuncular shape, generally more setose ros- trum and cauda, reticulate tergum) may jus- tify the erection of a new genus. Therefore, another analysis was conducted focusing only on the relationship of these four spe- cies to the Myzus included in the analysis. The objective of this analysis was to deter- mine whether the phloxae group and the available Myzus species would form sepa- rate monophyletic groups. The same set of characters was used. Three Palearctic Ova- tus were included, the first two as outgroup taxa: O. crataegarius, Ovatus glechomae Hille Ris Lambers, and O. malisuctus. In- group species were the phloxae group, and all ten Myzus in the first analysis. A “branch and bound” search found fourteen trees of length 71, with a CI of 0.55, RI of 0.56, and RC of 0.31. One of these trees is presented in Fig. 2. Similar to the previous analysis, the phloxae group is part of a clade with two Myzus (Nectaro- siphon) species. These Myzus have swollen 47 siphunculi and, similar to the phloxae group, two setae on the first segment of the hind tarsus. Rearranging the trees to make the phloxae group and Myzus monophyletic requires only a two step increase in tree length. A successive weighting analysis was conducted, which selected four of the original fourteen trees of length 71 (weight- edt statistics: teneth, 19,61 15"Cl, 0.775: Ri- 0:84: RE} 0:65). the tree in Fig: 2 1s ‘one of these trees. The results of weighted and unweighted decay and bootstrap (500 rep- lications) analyses are presented in Fig. 2 as well. Unweighted data supported the uni- ty of the phloxae group through | step of decay and in 52% of the bootstrap trees. Few branches were strongly supported by either measure using unweighted data. Weighted data yielded very strong support for the phloxae group, which held up through six steps of rescaled decay and was present in 83% of the bootstrap trees. These results point to the necessity for the erection of a new genus to house the phloxae group. First of all, it is clear that the phloxae group is not congeneric with Ovatus from the Palearctic. The full anal- ysis that used Hyalomyzus as outgroup was overall rather weak, but where it was rela- tively strong was in the separation between the basal Hyalomyzus/Ovatus clade and the more apical Myzus/phloxae group clade. This showed that the phloxae group is more closely related to Myzus than to Palearctic Ovatus. The next step in the analysis ex- amined more closely the relationship be- tween the phloxae group and the other My- zus. The results showed that there was strong support for the monophyly of the phloxae group, but not for the relationship between it and the Myzus with which it was placed in the trees. Thus not only is there no support for the placement of the phloxae group in Ovatus, but the group also appears to be only weakly related to any species of Myzus. Our cladistic results as well as the peculiar morphology of the phloxae group provide ample justification for the erection of a new genus, as we do below. 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Abstrusomyzus Jensen and Stoetzel, new genus (Figs. 4-18) Type species: Phorodon phloxae Sampson Diagnosis.—Abstrusomyzus can be sep- arated from Ovatus, Myzus and other My- zus-like genera in North America by the pe- culiar shape of the siphunculi in all stages (Figs. 9-12), the distinctly pentagonal/hex- agonal reticulated pattern of the tergum (Fig. 4), 3,3,2 first tarsal segment chaeto- taxy, and the usual presence of pigmented cross-bands on the abdomen of alate vivi- parae (Figs. 17, 18). The completely pig- mented tergum in apterous viviparae of the three species other than A. phloxae is also unusual. One North American species that may be closely related to Abstrusomyzus is Aphthargelia symphoricarpi (Thomas). The latter species shares with Abstrusomyzus the same type of dorsal reticulate sculpturing, moderately prominent Myzus-like antennal tubercles, a relatively large number of cau- dal setae, large lateral tubercles, and dark transverse abdominal bands in the alate vi- viparae. Abstrusomyzus and A. symphori- carpi differ significantly, in that the latter species has 3,3,3 first tarsal chaetotaxy, very short almost barrel-shaped siphunculi, and more numerous secondary rhinaria on antennal segments III-V. Description.—Apterous vivipara: Nymph: hind tibiae of nymphs without spinules api- cally. Adult: body length 0.83—-1.94 mm. Dorsum of head capsule (Figs. 5—8) pale to black, with small sparse spinules often ar- ranged in curving rows. Spinal region of head capsule with more or less triangular area of reticulate sculpturing; spinal tuber- cles sometimes present. Dorsum of head normally with 4 setae in posterior row, with 3 pairs farther forward; small lateral ocelli or traces thereof often present. Antennal tu- bercles moderately to strongly produced, rough, with several setae. Frons often pro- duced slightly as a median tubercle. Setae on head short, blunt, much shorter than bas- al width of antennal segment III. Eyes nor- mal, with ocular tubercles and interfacetal spaces brownish pigmented. Ventral surface of head capsule more or less evenly, but sometimes very lightly, spinulose, with spi- nules often arranged in rows. Antennal tu- bercles with large ventral projections bear- ing a few short setae. Ventral head setae sometimes as short as the dorsal setae, sometimes about twice as long. Mandibular area of mouthparts smooth. Antennae nor- mally shorter than body; antennal segment I rough, roughest medially, with several se- tae, these about equal in length to those on dorsum of head capsule; antennal segment II rough, roughest ventrally and medially, with course imbrications; antennal segment III covered with imbrications, rarely with 1 or 2 secondary rhinaria; remainder of an- tenna roughly imbricated. Rostrum reaching middle of thorax, with segment II strongly ornamented with rows of spinules; segment III with 2 pairs of setae, these located in apical %; ultimate rostral segment about equal in length to tarsal segments II, or about 0.1 mm, with 3—9 accessory setae. Pronotum with 3 pairs of short setae: 1 spi- nal, 1 pleural, and 1 lateral; the lateral pair associated with a frequently present pair of lateral tubercles that are sometimes bi- or trifid; setae about equal in length to those on the dorsum of the head. Surface of pron- otum reticulate, the reticulations usually somewhat flattened front to back. Meso- and metathoracic terga reticulate and often pigmented, setae extremely short, blunt. Coxae of all legs covered with spinulose imbrications. Femora with sparse imbrica- tions, especially anteriorly, with setae about equal in length to dorsal head setae. Tibiae sparsely setose, basal setae much shorter than those at apex. First tarsal segments with 3,3,2 setae (i.e., hind tarsus I with 2 setae), second tarsal segments imbricated. Mesothoracic furca sessile or with short stalk. Abdomen with tergum reticulate throughout, sometimes pigmented. Dorsal abdominal setae blunt, very short, except sometimes longer on tergite VIII. Abdom- inal segments II-VI often with lateral tu- 49 VOLUME 101, NUMBER | [0.156 mm G yi ’ afebe ’ [0.180 mm [0.170 mm Head with first three antennal segments of apterous viviparae. 5, Abstrusomyzus leucocrini. 6, A. Figs. 5-8. phloxae. 7, A. reticulatus. 8, A. valuliae. 50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON [ 0.079 mm Figs. 9-16. Siphunculi and caudas of apterous viviparae. 9, 13, Abstrusomyzus leucocrini. 10, 14, A. phloxae. 11, 15, A. reticulatus. 12, 16, A. valuliae. bercles of various sizes. Siphunculus cylin- drical over most of its length, slightly swol- len toward the tip (Figs. 9-12), imbricated throughout. Cauda moderately long, with 4—15 setae, more or less pointed apically. Tergite VIII with 4—7 setae, normally 4. Ventral abdominal setae pointed. Gonapo- physes 3 in number. VOLUME 101, NUMBER 1 & a Bag JeSo ade 51 Figs. 17, 18. Abdominal tergum of alate viviparae. 17, Abstrusomyzus leucocrini. 18, A. phloxae. Alate vivipara: Body length 0.98—1.92 mm. Head capsule dark, slightly wrinkled or smooth, not reticulate. Lateral and me- dian ocelli present, median ocellus creating a strong median tubercle. Antennal tuber- cles only slightly rough, with several setae. Ventral surface of head capsule more or less smooth, sometimes with a few sparse spi- nules. Antennal segment III with secondary rhinaria scattered over most or all of its length, limited to one side, normally less than 20 in number; antennal segment IV oc- casionally with a few secondary rhinaria; antennal segment V without secondary rhi- naria. Thoracic tergites smooth. Sclerites of thorax dark brown, of normal design. Ab- domen with dark lateral sclerites, and often dark cross bands on some segments. Dark abdominal cross bands often reticulate in a similar fashion to the tergum of apterous vivipara. Otherwise essentially as in apter- ous vivipara. Etymology.—The generic name is taken from the Latin word “‘abstrusus,”’ meaning hidden or concealed and Myzus, which is based on the Greek “‘myzo,”’ meaning suck. The name is meant to draw attention to the way the type species of the genus is often hidden in ant-created shelters or on the low- er leaves of its host plants. The gender is masculine. KEY TO APTEROUS VIVIPARAE OF ABSTRUSOMYZUS 1. Dorsum of apterous vivipara unpigmented; an- tennal tubercles strong, with prominent con- verging processes (Fig. 6); dorsum with hex- agonal reticulation usually faint due to the pale, uNSCleEnOtZeEd \CEROUIMs hee a ag. cates none mel elles, Saree Abstrusomyzus phloxae (Sampson) — Dorsum of abdomen of apterous vivipara dark pigmented, usually black; antennal tubercles strong, but without converging processes (Fig. 5, 7, 8); dorsum with distinct hexagonal retic- i) Wlationy (Higa) nes oo secrete a Rene Apterous vivipara without traces of lateral ocelli, and without spinal tubercles on head; antennal segments I & II nearly black, antennal segment III unpigmented, except extreme tip slightly dusky; dorsum entirely dark, concolo- rous with cauda and siphunculus i) FTP SO RES Abstrusomyzus reticulatus (Heie) — Apterous vivipara usually with traces of lateral ocelli, and sometimes with low spinal tubercles on head; antennal segment I brown, antennal segment IIT much paler than I, antennal segment Il with tip brown; dorsum entirely dark or with some pale areas, cauda and siphunculus pigmentation variable 3. Tergite VIII with middle pair of setae pointed, about twice as long as lateral pair of setae on tergite VIII, and about three times as long as setae on tergite VII; cauda usually with fewer than 9 setae (Fig. 16) zn Een eee Abstrusomyzus valuliae (Robinson) — Tergite VIII with middle pair of setae blunt, about equal in length to the lateral pair on ter- gite VIII and those on tergite VII; cauda usu- ally with more than 9 setae (Fig. 13) .. Abstrusomyzus leucocrini (Gillette and Palmer) KEY TO KNOWN ALATE VIVIPARAE OF ABSTRUSOMYZUS 1. Setae on abdominal tergite VIII distinctly lon- ger than setae on tergite VII, and about equal to basal width of antennal segment III; lateral tuberclesysmalllior- absent aaa... Juokopekcke Peet Abstrusomyzus valuliae (Robinson) — Setae on abdominal tergite VIII about equal in length to the setae on abdominal tergite VII, and much shorter than the basal width of an- tennal segment III; lateral tubercles various . . . Cauda relatively setose, usually with more than 9 setae; lateral abdominal tubercles relatively large (Fig. 17); antennal tubercles moderately prominent as in Fig. 5 .. Abstrusomyzus leucocrini (Gillette and Palmer) — Cauda usually with fewer than 9 setae; lateral abdominal tubercles small or absent (Fig. 18); antennal tubercles sometimes strongly pro- duced similar to the apterous vivipara (Fig. 6) BES Aa cirri Abstrusomyzus phloxae (Sampson) i) NO Abstrusomyzus leucocrini (Gillette and Palmer 1929), new combination (Figs:.425, 9; 13, 17) Myzus leucocrini Gillette and Palmer 1929: 470; Gillette and Palmer 1934: 202: Palmer 1952: 338. Ovatus leucocrini: Eastop and Hille Ris Lambers 1976: 328; Smith and Parron 1978: 225; Remaudiére and Remaudiére 19973135: This species was described from several collections made by L.C. Bragg from Fort Collins, Colorado, in May of 1916. The aphids were found on Leucocrinum mon- tanum, a small herb of the Liliaceae. We know of no other collections of this aphid 52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from Leucocrinum. There are a few indi- vidual alate viviparae found in traps and on various plants that seem to be this species, but lack of good host plant records makes any deductions about the biology of this species difficult. The presence of many ap- terous and alate viviparae on an herb (Leu- cocrinum) in May suggests a monoecious life cycle. Further collecting will be needed to determine whether this species is mo- nophagous on Leucocrinum, or feeds on other plants as well. Within Abstrusomyzus, this species is most similar to A. reticulatus and A. valu- liae. In the material at hand (19 apterous, and 15 alate viviparae), the apterous vivi- parae of A. /eucocrini almost always have partially developed lateral ocelli, and often spinal tubercles on the head as well (Fig. 5). Abstrusomyzus reticulatus lacks both of these, and A. valuliae always lacks the lat- ter, but usually lacks both. The cauda of A. leucocrini is usually more setose, ranging from 9-15 setae in both alate and apterous viviparae, compared to 4—9 setae on the cauda (Figs. 13, 15, 16) of the other two species. This species and A. phloxae have characteristic brown to orange regions of pigmentation surrounding the bases of the siphunculi. It is not known whether the oth- er two species in the group have this un- usual pigmentation pattern. Differences be- tween A. /eucocrini and A. phloxae are dis- cussed under the latter species. Adequate descriptions of this species have been pub- lished previously (Mason 1940, Palmer 1952): Types of this species are located in the USNM. A single paratype slide was also obtained on loan from the BMNH, and an- other from UCB. Other material examined were specimens from the same series as the types, some of which were cleared and re- mounted for this study. Abstrusomyzus phloxae (Sampson 1939), new combination (Figs. 6, 10, 14, 18) Phorodon phloxae Sampson 1939: 174. Myzus plantagineus Passerini (misidentifi- VOLUME 101, NUMBER 1 cation): Williams 1911: 65; Davis 1910: 495; Davis 1911: 23; Mason 1940: 17. Ovatus phloxae: Hille Ris Lambers 1966: 600; Heie 1972: 450; Eastop and Hille Ris Lambers 1976: 329; Smith and Par- ron 1978: 225; Remaudiere and Remau- diére 1997: 135. Abstrusomyzus phloxae was described from California based on specimens col- lected on Phlox subulata. Sampson de- scribed the species in Phorodon Passerini under a broad concept of the genus that in- cluded species that are today considered Ovatus and Myzus. Since Sampson (1939), the species has been collected on many oth- er plants (Table 1). In the eastern half of the U.S.A., A. phloxae is most often found on the crown, young leaves, and roots of Plan- tago major. This led to the misidentification of this species by Williams (1911) and Ma- son (1940) as Myzus plantagineus Passeri- ni. Hille Ris Lambers (1966) was the first to understand the identity and wide host range of this species. This species has been found on many un- related host plants, but mostly only in iso- lated collections. It has been best studied in eastern U.S.A. on Plantago major. It was collected on this plant in Illinois, Maryland, Nebraska, Pennsylvania, and Virginia in May, June, and July of various years. When feeding on this plant A. phloxae is often tended by ants (Lasius alienus Foerster in Maryland) which build earthen “‘tents”’ sur- rounding the young leaves on which the aphids feed. Such “‘tents’”’ we found in Mar- yland housed a mixture of aphids, including A. phloxae, Aphis gossypii, and Nearctaphis bakeri (Cowen). We reared A. phloxae on Plantago on potted plants indoors. To test acceptance of two host plants used by other Abstrusomyzus species, we allowed the population of A. phloxae on Plantago to in- crease, and provided Fragaria sp. (the host genus of A. valuliae) and Oxalis sp. (the host genus of A. reticulatus) in adjacent potted plants. Many A. phloxae individuals attempted to colonize Fragaria and Oxalis, DS, but no colonies were established during the two weeks of attempts. The A. phloxae pop- ulation eventually killed its host plants. This bolstered the somewhat weak morpho- logical separation between A. phloxae and A. reticulatus and A. valuliae, since despite the polyphagy of A. phloxae, it was unable to colonize the hosts of these two species. The aphids reared on potted plants were heavily parasitized by the aphelinid Aphel- inus asychis Walker. Little is known about the life history of this species. It has been collected in every month except October and November. In Maryland the earliest collection was Feb- ruary. In Oregon it is most commonly found in August and September. The single known collection of oviparae was found in Abbotsford, British Columbia on 2 January 1992. Given that males have never been found, and considering that it has been col- lected almost throughout the year, we sus- pect that A. phloxae is primarily anholo- cyclic. Concerted collecting efforts during autumn and winter will be required to de- termine the life history of this species. When A. phloxae is found in nature, it almost always lives on the lower leaves of its host plant, and generally on plants that are low to the ground or have a basal ro- sette. When reared on potted plants indoors, the aphid thrived on all parts of the plant, but moved to the upper parts only when populations were very large. This propen- sity for living near the ground, but on many different plant species, suggests that this aphid may be a rare example of a habitat- specific, rather than host-specific aphid. Other examples of this phenomenon are known in the aphids, such as the polypha- gous tree feeding aphid Longistigma caryae (Harris), which feeds specifically on bark, and Rhopalosiphum nymphaeae, which feeds on aquatic herbs. Chromosome number is often useful as a taxonomic character in aphids. For most species of Myzus in which it is known, the chromosome number is 2n = 12, with a few having 2n = 13 or 14. One slide obtained 54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON on loan from the BMNH contains a sample of apterous viviparae of A. phloxae that was karyotyped as 2n = 18 by R.L. Blackman. Karyotyping of the other Abstrusomyzus species may show that chromosome num- bers will also support the separation of Abs- trusomyzus from similar Myzus-like genera. Within Abstrusomyzus, apterous vivipa- rae of A. phloxae are easily recognized by their pale tergum and large, distinctly con- verging antennal tubercles (Fig. 6). Alate viviparae are far more difficult to separate from other members of the group. The alate form of A. reticulatus is unknown. The long middle pair of setae on abdominal tergite VIII is diagnostic for A. valuliae. This leaves A. /eucocrini as the primary species that may be confused with A. phloxae in the alate stage. Abstrusomyzus phloxae has rel- atively more prominent antennal tubercles, and usually has fewer caudal setae; A. phloxae normally has eight or fewer caudal setae (but sometimes up to 10), compared to 9-13 in A. leucocrini (Figs. 13, 14). Many A. leucocrini have spinal tubercles on the head, which are always absent in A. phloxae. Finally, A. phloxae tends to have smaller lateral tubercles, and often fainter or less extensive abdominal pigmentation (Figs. 17, 18). Good descriptions of A. phloxae can be found in Sampson (1939) and Mason (1940). Many paratypes of this species were seen, having been borrowed from the UCB. Other material examined included material from all the host plants and states listed for this species in Table 1. Seventy eight ap- terous viviparae, and 59 alate viviparae were seen during this study. Abstrusomyzus reticulatus (Heie 1972), new combination (Bigs-7 7. leet) Ovatus reticulatus Heie 1972: 447; Eastop and Hille Ris Lambers 1976: 329; Smith and Parron 1978: 225; Remaudiére and Remaudiére 1997: 135. This species was described based on nine specimens collected on Oxalis ?stricta in North Carolina. These specimens are still the only ones known for the species. They include seven apterous viviparae, one Ovi- para, and one brachypterous male. Abstrusomyzus reticulatus is apparently monoecious holocyclic based on the occur- rence of viviparae, a male, and ovipara to- gether on the herbaceous Oxalis. According to Heie (1972) the aphids cause the leaves of Oxalis to curl. Further collecting will be required to determine the host range of this species. This species can be distinguished from A. phloxae as described above. Abstrusomyzus valuliae is distinct from this species and others in the group because of the long se- tae on abdominal tergite VIII. Abstruso- myzus reticulatus is most similar to A. leu- cocrini. These species can be separated based on the small number of caudal setae in A. reticulatus, its complete lack of spinal tubercles on the head, and relatively dark antennal segment II. This is the only spe- cies of Abstrusomyzus for which the male is known. The brachypterous condition of the single known male is probably aberrant. Heie (1972) provides good descriptions of all known morphs. Types of this species are housed in the USNM (holotype, one paratype), BMNH (3 apterous vivipara paratypes), C.E Smith collection (two apterous viviparae, not seen), and in the collection of O.E. Heie (One apterous vivipara and one ovipara, not seen). Abstrusomyzus valuliae (Robinson, 1974), new combination (Figss.83 12% 16) Myzus valuliae Robinson 1974: 469. Ovatus valuliae: Eastop and Hille Ris Lambers 1976: 329; Smith and Parron 1978: 226; Remaudiere and Remaudiere 19972 135. Abstrusomyzus valuliae is known only from the material used by Robinson for the description of the species from Manitoba, VOLUME 101, NUMBER 1 Canada. It was collected several times dur- ing the summer of 1973 and once in the spring of 1974 on Fragaria vesca, wild strawberry. These specimens include many apterous and alate viviparae. Little is known about the biology of this species. It is most likely monoecious, but host plant range is unknown. When on Fra- garia, the aphids cause the leaves to curl tightly, and the aphids feed inside the curled leaves (Robinson 1974). This species can be distinguished from A. phloxae as described under that species. Among the three darkly pigmented species of Abstrusomyzus, this species can be most easily distinguished by the long middle pair of setae on the abdominal tergite VIII. It is most similar to A. leucocrini, both species frequently having lateral ocelli or traces thereof in the apterous vivipara. Robinson (1974) described this species thoroughly. The holotype of A. valuliae, which we did not see, was deposited in the Canadian National Collection, Ottawa. Paratypes were deposited in the Canadian National Collection, BMNH, and the USNM. Many paratypes and some additional material from the same collections were examined for this study. Status of Ovatus The removal of four species from Ovatus leaves only a single species of the genus in North America, O. crataegarius. This spe- cies can be separated from Myzus found in North America by its lack of spinules on the hind tibia of nymphs, the lack of a dor- sal pigmented abdominal patch in the alate viviparae, and the presence of many sec- ondary rhinaria on antennal segments III, IV, and V in alate viviparae. Ovatus cra- taegarius can be separated from another somewhat similar species, Phorodon hu- muli (Schrank), by its first tarsal chaetotaxy of 3,3,3, lack of pigmented abdominal patch in the alate viviparae, and its antennal tu- bercles, which are strongly convergent, but lack the extremely prominent finger-like process of Phorodon. The primary way that 55 Ovatus is separated from Hyalomyzus is the swollen siphunculi of the latter genus. An excellent diagnosis for the genus Ovatus, as it is understood here, is provided by Heie (1994) (i.e., Heie’s diagnosis does not ad- dress the characters of the four species here placed in Abstrusomyzus). ACKNOWLEDGMENTS The authors thank Paul A. Brown (The Natural History Museum, London), David J. Voegtlin (Illinois Natural History Survey, Champaign), Cheryl Barr (University of California, Berkeley), James LaBonte (Oregon State University, Corvallis), and C. K. Chan (Agriculture Canada, Vancouver) for the loan of specimens. The manuscript was improved by the comments of Natalia J. Vandenberg and Michael E. Schauff of the U.S. Department of Agriculture, Sys- tematic Entomology Laboratory, Victor E Eastop of The Natural History Museum, London, David J. Voegtlin of the Ilinois Natural History Survey, and Robert G. Foottit of Agriculture Canada, Ottawa. Identifications of the ants and aphelinids were provided by D. R. Smith and M. E. Schauff, respectively, of the U.S. Depart- ment of Agriculture, Systematic Entomol- ogy Laboratory. LITERATURE CITED Blackman, R. L. and V. F Eastop. 1994. Aphids on the World’s Trees. CAB International, Walling- ford, U.K. 987 pp. Bremer, K. 1994. Branch support and tree stability. Cladistics 10: 295-304. Davis, J. J. 1910. A list of the Aphididae of Illinois, with notes on some of the species. Journal of Eco- nomic Entomology 3: 482—499. . 1911. Williams’ “‘The Aphididae of Nebras- ka’: a critical review. University Studies (Nebras- ka) 11: 253-291 +8 pls. Eastop, V. E 1966. A taxonomic study of Australian Aphidoidea (Homoptera). Australian Journal of Zoology 14: 399-592. . 1973. Deductions from the present day host plants of aphids and related insects. Symposium of the Royal Entomological Society, London 6: 157-178. Eastop, V. FE and D. Hille Ris Lambers. 1976. Survey 56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of the World’s Aphids. Dr. W. Junk, b.v., The Hague. 573 pp. Gillette, C. P. and M. A. Palmer. 1929. Five New Aphi- didae from Colorado. Annals of the Entomologi- cal Society of America 22: 468-476 +2 pls. . 1934. The Aphidae of Colorado. Part III. An- nals of the Entomological Society of America 27: 133-255. Gould, S. J. 1977. Ontogeny and Phylogeny. Harvard University Press, Cambridge. 501 pp. Heie, O. E. 1972. Ovatus reticulatus, a new species of aphid from Oxalis in North Carolina (Homoptera: Aphididae). Proceedings of the Entomological So- ciety of Washington 74: 446—450. . 1994. The Aphidoidea (Hemiptera) of Fen- noscandia and Denmark. V. Family Aphididae: Part 2 of tribe Macrosiphini of subfamily Aphi- dinae. Fauna Entomologica Scandinavica 28. 242 PP. Hille Ris Lambers, D. 1966. Notes on California aphids, with descriptions of new genera and new species. Hilgardia 37: 569-623. Jensen, A. S. 1997. Redefinition of the aphid genus Sitobion Mordvilko (Hemiptera: Aphididae) based on cladistic analyses, with emphasis on North American species. Systematic Entomology 22: 333-344. Maddison, W. P. and D. R. Maddison. 1992. Mac- Clade: Analysis of Phylogeny and Character Evo- lution. Version 3.0. Sinauer Associates, Sunder- land, Massachusetts. Mason, P. W. 1940. A revision of the North American aphids of the genus Myzus. United States Depart- ment of Agriculture Miscellaneous Publication No. 371: 1-30. Miyazaki, M. 1987. Morphology of aphids. pp. 1—25. In Minks, A. K. and Harrewijn, P., eds., Aphids, Their Biology, Natural Enemies and Control 2A. Elsevier, New York. Nielsson, R. J. and D. H. Habeck. 1971. The genus Hyalomyzus (Homoptera: Aphididae), with the de- scription of a new species. Annals of the Ento- mological Society of America 64: 883-887. Palmer, M.A. 1952. Aphids of the Rocky Mountain Region, Vol. 5. The Thomas Say Foundation. 452 Pp. Remaudiere, G. and M. Remaudiére. 1997. Catalogue of the World’s Aphids. INRA, Paris. 473 pp. Robinson, A. G. 1974. Four new species of aphids (Homoptera: Aphididae) from Manitoba. Canadi- an Entomologist 106: 467—472. Sampson, W. W. 1939. California of aphids of the ge- nus Phorodon. The Pan-Pacific Entomologist 15: 173-175. Smith, C. FE and C. S. Parron. 1978. An Annotated List of Aphididae (Homoptera) of North America. North Carolina Agricultural Experiment Station. 428 pp. Swofford, D. L. 1993. PAUP: Phylogenetic Analysis Using Parsimony, Version 3.1.1. Computer pro- gram distributed by the Illinois Natural History Survey, Champaign, Illinois. Williams, T. A. 1911. The Aphididae of Nebraska. University Studies (Nebraska) 10: 85-175. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 57-63 THE LARVAL INSTARS OF THE WHEAT MIDGE, SITODIPLOSIS MOSELLANA (GEHIN) (DIPTERA: CECIDOMYIIDAE) RAYMOND J. GAGNE AND JOHN E DOANE (RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. De- partment of Agriculture, % National Museum of Natural History, MRC 168, Washington, DC 20560-0168, U.S.A. (email: rgagne @sel.barc.usda.gov); (JFD) Agriculture and Agri-Food Canada Research Centre, 107 Science Place, Saskatoon, Saskatchewan S7H OX2 Canada. Abstract.—The wheat midge, Sitodiplosis mosellana (Géhin), is shown to have three larval instars. Each instar is described and illustrated, the initial instar for the first time. An alleged synonym of the wheat midge, Cecidomyia amyotii Fitch, is removed from synonymy with the wheat midge and considered a dubious name. Key Words: The wheat midge, Sitodiplosis mosellana (Géhin), is one of the two most important cecidomyiid pests of wheat in North Amer- ica, the other being the Hessian fly, Maye- tiola destructor (Say). Originally from the Palearctic Region, both are now well estab- lished in North America. All three larval instars of the Hessian fly have been de- scribed in detail (Gagné and Hatchett 1989), but those of the wheat midge are less well known, and its first instar has not pre- viously been described. The general life history of the wheat midge or, in the United Kingdom, the or- ange wheat blossom midge, has been sum- marized by Reeher (1945) and Barnes (1956), and more information on certain aspects of its attack and feeding were treated by Mukerji et al. (1988) and Elliott and Mann (1996). Females lay eggs on emerging spikes of wheat before anthesis. Upon hatching, the larvae crawl to and settle upon the developing flower parts, where they feed and interfere with the proper development of the kernels. The full grown larva eventually drops from the wheat head, crawls into the soil, and wheat midge, Sitodiplosis, larva constructs a silk cocoon in which the lar- va overwinters. We show that the wheat midge has three larval instars, as do the Hessian fly and all other cecidomyiids that have been carefully studied (Gagné 1989). Upon hatching, the first instar of the wheat midge crawls to a feeding site on the de- veloping wheat grain, settles, and begins to feed. Within two to three days the larva molts to the second instar, which contin- ues to feed and grow until the third instar begins to develop. The second instar skin then becomes brittle and serves as a tem- porary cocoon for the third instar, which does not feed. The fully developed third instar can exit from the second instar skin immediately but may stay within this skin on the kernel for several weeks. Upon leaving the temporary cocoon, the third instar drops to the ground and burrows into the soil where it spins its cocoon. In most other cecidomyiids the second instar skin is shed as soon as the third in- star is fully developed and, in gall-making species, can be found crumpled in a com- pact mass at the caudal end of the third 58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON instar. The situation in the wheat midge is somewhat analogous to that of the Hes- sian fly in which the second instar skin also serves as a cocoon for the non-feed- ing third instar, with the difference that the third instar of the Hessian fly pupates there also. Because the Hessian fly feeds head groundwards between the culm and leaf sheath, the third instar has to reverse position from head to tail within the sec- ond instar skin. In this way the pupa of the Hessian fly is positioned so that adults are able to exit from the wheat sheaths. The third instar of the wheat midge is not so constrained, so does not need to re- verse its position within the second instar skin. These two cecidomyiids on grasses are not closely related, each belonging to a different supertribe, so this rare devel- opment of a brittle second instar skin as a puparium or larvarium has evidently evolved separately. A puparium is present also in another grass-infesting gall midge, the sorghum midge, Stenodiplosis sorgh- icola (Coquillett) (Solinas 1986), which belongs to the same supertribe but to a different tribe than the wheat midge. A puparium is also known for other gall midges that do not occur on grasses but is still rare. Examples are Thurauia aquatica Riibsaamen from sedges in Europe and an undescribed species that belongs to no known genus from maple seeds in Japan. The description and figures that follow will allow recognition of each of the three instars of the wheat midge. They are nec- essary to correct misinformation in Borkent (1989) that was based on mixed series of three species and in which it was asserted that the wheat midge had four instars. METHODS In 1997, collections of wheat heads (cul- tivar Roblin) were made by one of us (JFD) from a plot area (latitude 52.1951, longi- tude 106.1071) near St. Denis, a hamlet about 26 miles east of Saskatoon, Saskatch- ewan, Canada. Twenty-five primary heads were collected at intervals of 1 or 2 days from July 7 until July 18 and then at inter- vals of 3 to 4 days until July 31. The first eggs were found on the outside of the glumes on July 7 and the first newly hatched larvae on July 9. The larvae were removed from the heads and preserved in 70% alcohol for subsequent examination. Some specimens were mounted on micro- scope slides using the method outlined in Gagné (1989); earlier instars were mounted in Hoyer’s medium because of their small size and the risk of their being lost in the various steps involved in balsam mounting. Terminology for larval morphology follows that in Gagné (1989). DESCRIPTION OF LARVAL INSTARS OF SITODIPLOSIS MOSELLANA The first instar differs markedly from the remaining instars. It has only one pair of functional spiracles (Figs. 10, 12), which are situated on the eighth abdominal seg- ment and are relatively large in relation to body size compared to the eighth abdomi- nal spiracle of other instars (Fig. 13). Its cuticle is entirely smooth except for several horizontal rows of tiny spicules dorsally and ventrally near the anterior part of most segments (Figs. 11, 12). The second and third instars each have spiracles on the first thoracic and on the first through eighth ab- dominal segments. The cuticle of the sec- ond and third instars is rough, covered al- most entirely by raised scale-like bumps (Figs. 4—7). The third instar differs from the second in having a spatula, the clove- shaped dermal structure on the venter of the prothorax (Fig. 9). Because this structure begins to develop while the third instar is still encased in the second instar, the spatula is usually visible through the skin of older second instars. An additional difference be- tween the second and third instars is that the two caudalmost terminal papillae are more equal in size in the third instar than in the second (Fig. 14). A more detailed description follows: First instar (Figs. 10—12).—Body length 0.45—1.05 mm. Antenna three times as long VOLUME 101, NUMBER 1 59 . ~ mene Figs. 1-7. Sitodiplosis mosellana. 1—3, Brittle second instar skins from which third instar is emerging (1, 2) or has emerged (3). 4, Second instar (dorsolateral). 5, Same (lateral). 6, Third instar (dorsal). 7, Same (lateral). 60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON as wide. Two pairs of spiracles apparent, one on prothoracic segment, the other on eighth abdominal segment: prothoracic pair evident only as short projection, evidently not functional; posterior pair prominent, with three acute apical projections. Cuticle smooth except for several horizontal rows of tiny spicules dorsally and ventrally near anterior part of each segment except for prothorax. Pattern and number of papillae basic for supertribe Cecidomyiidi, their se- tae mostly very short, dorsal and pleural pa- pillae of eighth segment slightly longer than preceding segments, and terminal papillae modified as follows: one dorsal, subcaudal pair with short setae not surpassing length of those on preceding segments; one lateral pair with elongate setae several times lon- ger than subcaudal pair; two caudal pairs with stout, wide setae, the outer of these two pairs more than twice length of inner pair. Second instar (Figs. 4, 5, 8, 9, 14).— Body length 1.05—3.30 mm. Antenna about twice as long as wide. Spiracles present on prothoracic and first through eighth abdom- inal segments, eighth abdominal pair slight- ly larger than preceding spiracles, none with apical projections. Cuticle rugose, cov- ered almost completely with raised scale- like bumps, these much smaller and ar- ranged in several regular anteroventral hor- izontal rows on mesothoracic to eighth ab- dominal segments. Pattern of papillae sim- ilar to that of first instar but setae more con- spicuous. Terminal papillae modified as follows: two lateralmost papillae with setae, the more ventral pair with longer setae; two pairs with stout setae, the inner pair with setae noticeably thinner than outer pair. Third instar (Figs. 6-9, 13, 14).—Body length 2.80—3.20 mm. Antenna and spira- cles as for second instar. Cuticle also as for second instar except for presence of ventral prothoracic spatula and horizontal antero- ventral rows of bumps more numerous and more extensive. Pattern of papillae similar to that of second instar. Terminal papillae modified as for second instar except inner Figs. 8, 9. Sitodiplosis mosellana. 8, Third instar inside second instar skin (ventral). 9, Detail of same (a = spatula). pair of caudal, stout setae more nearly equal in size to outer pair. Remarks.—Borkent (1989) described and illustrated four alleged instars for the wheat midge, each with a spatula. The first instar of that paper is a third instar of an undetermined Clinodiplosis sp. Its posterior abdominal segment, as illustrated in his Fig. 2A, has three of the four pairs of terminal papillae corniform (short and stout) and only one pair setiform, characteristic of spe- cies of Clinodiplosis. The presence of a spatula indicates that the specimen is a third instar. Specimens of these larvae occasion- ally are found in association with wheat midge larvae. Representatives of this spe- cies in the USNM are curated with wheat midge larvae until adults are reared and the species can be identified further. The second instar of Borkent (1989) is a third instar of an undescribed species of Contarinia (broad sense). It has one pair of VOLUME 101, NUMBER 1 61 a —— dorsal papilla ahha } _.<\+— pleural papilla ) +—- spiracle =«——_———— terminal papillae Figs. 10-14. Sitodiplosis mosellana. 10, First instar (dorsal). 11, Same, detail of head and first two thoracic segments. 12, Same, detail of posterior segments. 13, Third instar, posterior part of eighth segment and terminal segment (dorsal). 14, Juxtaposed third instar (upper) and second instar (lower) terminal segments (dorsal). Bar lengths are in mm. 62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON corniform (short and stout) papillae and three pairs of setiform papillae, two of which are of similar size. The presence of a spatula indicates that this specimen is also a third instar. Specimens in the USNM of similar larvae found in association with the wheat midge also have a spatula and are definitely third instars. These are also tem- porarily curated with wheat midge larvae until adults are found that can be identified further. The third and fourth instars of Borkent (1989) are actually the second and third in- star, respectively, of the wheat midge. Both are described in that paper as having a spat- ula, but the second instar has none. As not- ed above, older specimens of the second in- star may appear to have a spatula due to the developing third instar inside (Figs. 8, 9). One can be certain of the train of instars of a particular species by observing nearly ful- ly developed preecdysal instars within the body of a previous instar, as we have done here: Borkent (1989) cited Borkent (in press), “Description of the larval instars of the Wheat Midge Sitodiplosis mosellana (Gé- hin) (Diptera: Cecidomyiidae). Can. J. Zool.’ No paper by Borkent on the subject of the wheat midge has appeared in the Ca- nadian Journal of Zoology. This occasion is taken to remove the name Cecidomyia amyotii Fitch from syn- onymy with the wheat midge. Felt (1925) listed C. amyotii as “‘probably a synonym of” the wheat midge and Foote (1965) cat- aloged it as a synonym of the wheat midge for the first time. According to Fitch’s (1861) original description of C. amyotii, based on three specimens caught at a light, the female antenna has ‘“‘eighteen (7?) joints, not separated by pedicels, the joints glob- ular.”” The female of the wheat midge has 12 antennal flagellomeres, each separated by a conspicuous pedicel or neck, so C. amyotii cannot be the wheat midge. Be- cause the types of C. amyotii are presum- ably lost and the species cannot be deter- mined with certainty, we consider this spe- cies a dubious name. ACKNOWLEDGMENTS We thank Nit Malikul for making the slide preparations; Cathy Anderson for computer assistance in making the plates; and, for their comments on drafts of the manuscript: Philip S. Barker and Robert J. Lamb of Agriculture Canada Research Cen- tre, Winnipeg, Manitoba; John W. Brown and Allen L. Norrbom of the Systematic Entomology Laboratory, USDA, Washing- ton, DC; Keith M. Harris, International In- stitute of Entomology, CAB, London, UK; and Odette Rohfritsch, Institut de Biologie Moléculaire des Plantes, Strasbourg, France. LITERATURE CITED Barnes, H. F. 1956. Gall Midges of Economic Impor- tance, Vol. 7: Gall Midges of Cereal Crops. Cros- by Lockwood & Son, London. 261 pp. and 16 pls. Borkent, A. 1989. A review of the wheat blossom midge, Sitodiplosis mosellana (Géhin) (Diptera: Cecidomyiidae) in Canada. Agriculture Canada Research Branch Technical Bulletin 1989—SE. Ot- tawa, Ontario. 18 pp, 6 pls. Elliott, R. H. and L. W. Mann. 1996. Susceptibility of red spring wheat, Triticum aestivum L. CV. Ka- tepwa, during heading and anthesis to damage by wheat midge, Sitodiplosis mosellana (Géhin) (Diptera: Cecidomyiidae). The Canadian Ento- mologist 128: 367-375. Felt, E. P. 1925. Key to gall midges (a resumé of stud- ies I-VII, Itonididae). New York State Museum Bulletin 257: 1—239, pls. 1-8. Fitch, A. 1861. Sixth report on the noxious and other insects of the State of New York. New York State Agricultural Society Transactions (1860) 20: 745— 868, pls. 1-2. Foote, R. H. 1965. Family Cecidomyiidae. /n 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. U.S. Department of Agriculture Handbook 276, 1696 pp. Gagné, R. J. 1989. The Plant-Feeding Gall Midges of North America. Cornell University Press, Ithaca, New York. xiii, 355 pp., 4 pls. Gagné, R. J. and J. H. Hatchett. 1989. Larval instars of the Hessian fly (Diptera: Cecidomyiidae). An- nals of the Entomological Society of America 82: 73-79. Mukerji, M. K., O. O. Olfert, and J. EF Doane. 1988. VOLUME 101, NUMBER 1 63 Development of sampling designs for egg and lar- Northwest. United States Department of Agricul- val populations of the wheat midge, Sitodiplosis ture Circular 732: 1-8. mosellana (Géhin) (Diptera: Cecidomyiidae), in Solinas, M. 1986. Allocontarinia, n.g., A. sorghicola wheat. The Canadian Entomologist 120: 497-505. (Coq.) n. comb. (Diptera, Cecidomyiidae). Ento- Reeher, M. M. 1945. The wheat midge in the Pacific mologica 21: 23-35. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 64-68 THE LARVA OF PELECINUS POLYTURATOR (DRURY) (HYMENOPTERA: PELECINIDAE) NorMAN E JOHNSON, LUCIANA MUSETTI, JAMES B. JOHNSON, AND KERRY KATOVICH (NFJ, LM) Department of Entomology, The Ohio State University, 1315 Kinnear Road, Columbus, OH 43212-1192, U.S.A. (e-mail: johnson.2@osu.edu;luciana @iris.biosci. ohio-state.edu); (JBJ) Division of Entomology, Department of Plant, Soil, and Entomo- logical Sciences, University of Idaho, Moscow, ID 83844-2339, U.S.A. (e-mail: djohnson @novell.uidaho.edu); (KK) Department of Entomology, 237 Russell Laborato- ries, University of Wisconsin, Madison, WI 53706, U.S.A. Abstract.—The first instar larva and exuviae of the last instar larva of Pelecinus po- lyturator (Drury) are described, illustrated, and compared with published descriptions of other larvae of Proctotrupoidea. Key Words: The application of characters from larval morphology to the study of relationships within the Hymenoptera is very uneven. There are some good examples of the use- fulness of immatures as an additional source of data (e.g., Evans 1987), but for many superfamilies little or nothing is known. Progress in this area is hindered for several reasons. Most Apocrita are parasit- oids of other insects, and the hosts for a number of groups are very poorly known. Many species are internal parasitoids; thus it is difficult to obtain early instars and of- ten the larvae are highly simplified in struc- ture. Within the Hymenoptera a remarkable ar- ray of larval body plans is found. Clausen (1940) outlined fourteen types among the parasitic Hymenoptera alone. Some of these are highly simplified, “‘embryonic,”’ forms, while others possess well-developed, ex- aggerated mandibles and caudal appendag- es. Later instars usually converge upon a generalized, hymenopteriform type. Little phylogenetic pattern has been found amidst this variety; at present we are confronted larval morphology, Pelecinidae, Proctotrupoidea with a diversity of forms without any un- derlying organizing principle. The superfamily Proctotrupoidea s. str. is comprised of ten extant families of internal parasitoids. The hosts for three of these (Renyxidae, Austroniidae, and Peradeni- idae) are unknown, and only a bare mini- mum of information is available for the Monomachidae and Roproniidae. The only families for which immature stages have been described are the Diapriidae (a large group of nearly 2,000 recognized species), Proctotrupidae (331 described species), and Heloridae (a relict group of 10 extant spe- cies). In total, the larvae of only six species from this complex have been described. We report here on the larvae of another family, the Pelecinidae, a small group (only one species currently recognized) of uncertain affinities (Rasnitsyn 1980, Dowton et al. 1997). MATERIALS AND METHODS Five parasitoid larvae were dissected from larvae of Scarabaeidae (Coleoptera), and preserved in ethanol. The specimens VOLUME 101, NUMBER 1 were found in the posterior two-thirds of the abdomen of the host. Three final instar exuviae were found attached to scarab re- mains from which Pelecinus had pupated. Specimens are stored in the collections of JBJ; the Ohio State University; the Insect Research Collection, University of Wiscon- sin; and El Colegio de la Frontera Sur, San Crist6bal de las Casas, Chiapas. I[llustra- tions were made using a camera lucida of whole specimens under alcohol and exuviae in temporary slide mounts in glycerine jel- ly. Pelecinus polyturator (Drury) Material examined.—USA. Michigan, Newaygo Co., 18 April 1974, host in soil of oak forest, ex Phyllophaga, one first in- star; Branch Co., 23 May 1974, hosts in soil of oak-hickory forest, four first instars, three from large 5 cm long larvae, probably Phyllophaga, one from small 2.5 cm larva, possibly Serica sp. Wisconsin, Marquette Co., 11 August 1992, in sandy soil of forest meadow, ex larva of Phyllophaga drakei (Kirby) one final instar exuviae; Jackson Co., 4 June 1992, ex P. drakei, one final instar exuviae; Oconto Co., 28 May 1996, ex Phyllophaga rugosa (Melsheimer), one final instar exuviae. MEXICO. Chiapas, Te- nejapa, Balun Canal, 2,300 m, 14 February 1997, ex Phyllophaga obsoleta (Blanchard) third instar, one final instar exuviae. First instar (Figs. 1—4).— Length 3.3—5.3 mm; mandibulate larva (Clausen 1940); head capsule well-developed, covering dor- sal and lateral sides of head, margins darkly pigmented, sclerotization extending beyond margins, gradually disappearing posterior- ly; epicranial suture (Fig. 2, es) well-devel- oped; no indication of eyes; antenna (Figs. 1—3, a) indicated by small paired submedial papilla; clypeolabral area (Fig. 3, c/) largely membranous, supported by ovoid sclero- tized ring, dorsal portion of this ring some- times incomplete; labrum with two medial tubercles (Fig. 1, /f); mandible (Figs. 1, 3, md) strongly developed, falcate, bearing a small subapical tooth; maxilla (Fig. 1, 3, 65 mx) supported anteriorly by narrow stipital sclerite, otherwise lobelike, membranous; maxillary palp, labium, and labial palp un- differentiated; head supported internally by extensive, strongly pigmented tentorium (Fig. 4, tn) in shape of central plate with anterior extensions continuous with labral sclerite, lateral arms surrounding base of mandibles, and broad posterior bilobed plate in labial region, a central ovoid fora- men visible, anterior to this with more strongly pigmented triangular prominence, anterior apex of triangle produced into small costa extending into labrum; no pro- legs visible; body with indeterminate num- ber of segments, without setae, apex of ab- domen acute; no spiracles visible. Final instar (Fig. 5).—Head capsule with posterior sclerotized, pigmented band, oth- erwise largely membranous; mandible (md) very small, weakly articulated with head; antenna, labrum, maxilla, maxillary palp in- distinguishable; labium (/b) visible as me- dial triangular raised surface behind man- dibles, with large circular field correspond- ing to each labial palp (/p), a small central area presumably representing opening of la- bial gland (/g) between palpi; mouthparts unsupported by sclerotized pleurostoma or hypostoma; body with 7 pairs of spiracles visible; tracheae well-developed. DISCUSSION The exuviae of the last instar larvae are associated with pharate adult Pelecinus po- lyturator, and their identity is unequivocal. The early instar larvae, however, are strik- ingly divergent in structure from the exu- viae. Our determination of them was based on the fact that they were internal parasit- oids dissected from larvae of Phyllophaga Harris (Coleoptera: Scarabaeidae), the only recorded host in the United States and Can- ada. The specimens also were collected in an area in which Pelecinus was very abun- dant. Muesebeck (1979) recorded Tiphia berbereti Allen, T. tegulina Malloch, T. transversa Say, T. vulgaris Robertson, and T. intermedia Malloch (Tiphiidae); Myzin- 66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0.5 Figs. 1-5. Pelecinus polyturator, larva. 1—4, Head of first instar. 1, Lateral view. 2, Dorsal view. 3, Frontal view, specimen with mandibles closed. 4, Frontal view, specimen with open mandibles exposing tentorium. 5, Mouthparts from final instar exuviae, right mandible detached. Abbreviations: a = antenna; cl = clypeolabral area; es = epicranial suture; lb = labium; lg = opening of labial gland; Ip = labial palp; It = labral tubercle; md = mandible; mx = maxilla; tn = tentorium. Scale in mm. VOLUME 101, NUMBER 1 um quinquecinctum (Fabricius) (Tiphidae); and Ophion nigrovarius Provancher (Ichneumonidae) as parasitoids of Phyllo- phaga. Woodruff and Beck (1989) listed a second species of Ophion as well as a num- ber of additional species of tiphiids and sco- liids. We ruled out the aculeates because they are external parasitoids. Ichneumo- noids usually are characterized by the pos- session of a hypostomal spur (Short 1978), a structure that was not observed in these specimens. Determination of the number of larval in- stars of internal parasitoids requires large numbers of observations of cohorts of known age in order to detect structural changes associated with molting. This has not been done yet for any species of proc- totrupoid, and no one has yet been able to rear Pelecinus through its life cycle. We could not determine the age of the observed larvae directly or infer their age from pub- lished observations of related species. Clau- sen (1940) stated that the characteristics that set apart mandibulate larvae are lost at the first molt. This was confirmed by Clan- cy (1946) in his studies of Helorus, another proctotrupoid. Therefore, we concluded that the larvae dissected from the hosts must be late first instars. Very little information on the immature Stages of proctotrupoids exists to form a context in which to discuss the structural features of Pelecinus. Larvae have been de- scribed and illustrated for Helorus anom- alipes (Panzer) (Heloridae; Clancy 1946), an unidentified species of Basalys West- wood (Diapriidae; Simmonds 1952), Basa- lys tritoma Thomson (Diapriidae; Wright et al. 1946), Coptera silvestrii (Kieffer) (Dia- priidae; Pemberton and Willard 1918), Par- acodrus apterogynus (Haliday) (Proctotru- pidae; Zolk 1924), Phaenoserphus viator (Haliday) (Proctotrupidae; Eastham 1929), and Brachyserphus parvulus (Nees ab Esenbeck) (Proctotrupidae; Osborne 1960). Large, sickle-shaped mandibles have been reported in the first instar for all these spe- cies. A sclerotized head capsule was de- 67 scribed in H. anomalipes, B. tritoma, and C. silvestril. Phaenoserphus viator lacks a complete head capsule, but does have a sclerotized ring surrounding the mouth- parts. Distinct antennal lobes are found in the helorid, Basalys spp. and the proctotru- pids, larger and more prominent than the structures found in Pelecinus. We observed no prolegs on any of the first instar larvae; these structures have been reported for H. anomalipes, P. viator, B. parvulus, and an unidentified proctotrupid (Clausen 1940; presumably Nothoserphus scymni Ash- mead). Two of our specimens have paired, nipple-like protuberances beneath the pos- terior portion of the head capsule (visible in Fig. 1). Because of their position, we hesitate to call these prolegs or to homol- ogize them with the labial palpi. The num- ber of observed spiracles reported varies from three (B. tritoma, C. silvestrii) to ten pairs (P. viator). The most striking feature we observed in the first-instar larva was the large tentorial endoskeleton. A similar structure was very briefly described in P. viator by Eastham (1929), suggesting that it may not have been as apparent or strongly pigmented as in Pelecinus. The tentorium is not men- tioned in the other descriptions. The larval specimens were dissected from hosts in the spring (18 April, 1974; 23 May, 1974) in Michigan. Therefore, it ap- pears that the species overwinters as late first instars within the Phyllophaga larvae. No more than a single larva was found in any one host. Three specimens were recov- ered from large (5 cm) hosts, presumably the final instar of the beetle. A fourth was found in a much smaller larva, either a much younger specimen or a different ge- nus, perhaps Serica MacLeay (Coleoptera: Scarabaeidae). Host size may contribute to the large variation in size of adult Peleci- nuS. ACKNOWLEDGMENTS Thanks to Lorena Ruiz-Montoya (San Cristobal de las Casas, Mexico) and Daniel 68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON K. Young (Madison, WI) for the loan of specimens. This material is based in part upon work supported by the National Sci- ence Foundation under Grant No. DEB- 9521648. LITERATURE CITED Clancy, D. W. 1946. The insect parasites of the Chry- sopidae (Neuroptera). University of California Publications in Entomology 7: 403-496. Clausen, C. P. 1940. Entomophagous Insects. Mc- Graw-Hill Book Company, Inc., New York. 688 PPp- Dowton, M., A. D. Austin, N. Dillon, and E. Bar- towsky. 1997. Molecular phylogeny of the apo- critan wasps: the Proctotrupomorpha and Eva- niomorpha. Systematic Entomology 22: 245-255. Eastham, L. E. S. 1929. The post-embryonic devel- opment of Phaenoserphus viator Hal. (Proctotry- poidea), a parasite of the larva of Prerostichus ni- ger (Carabidae), with notes on the anatomy of the larva. Parasitology 21: 1-21. Evans, H. E. 1987. Order Hymenoptera, pp. 597—710. In Stehr, EF W., ed., Immature Insects, Vol. 1. Ken- dall/Hunt Publishing Company, Dubuque, Iowa. 754 pp. Muesebeck, C. EK W. 1979. Pelecinoidea, pp. 1119- 1120: Jn, Krombein; K. V:, PD: Hurd, Jr, D. R. Smith, and B. D. Burks, eds. Catalog of Hyme- noptera in America north of Mexico. Smithsonian Institution Press, Washington, DC. Vol. 1, 1198 PP: Osborne, P. 1960. Observations on the natural enemies of Meligethes aeneus (F) and M. viridescens (FE) [Coleoptera: Nitidulidae]. Parasitology 50: 91— 110. Pemberton, C. E. and H. F Willard. 1918. A contri- bution to the biology of fruit-fly parasites in Ha- waii. Journal of Agricultural Research 15: 419— 466. Rasnitsyn, A. P. 1980. [The origin and evolution of the Hymenoptera.] Trudy Paleontologicheskogo Insti- tuta 174: 1-190. Simmonds, F. J. 1952. Parasites of the frit-fly, Osci- nella frit (L.), in eastern North America. Bulletin of Entomological Research 43: 503-542. Short, J. R. T. 1978. The final larval instars of the Ichneumonidae. Memoirs of the American Ento- mological Institute No. 25, 508 pp. Woodruff, R. E. and B. M. Beck. 1989. The scarab beetles of Florida (Coleoptera: Scarabaeidae). Part II. The May or June beetles (genus Phyllophaga). Arthropods of Florida and Neighboring Land Ar- eas 13. 226 pp. Wright, D. W., Q. A. Geering, and D. G. Ashby. 1946. The insect parasites of the carrot fly, Psila rosae, Fab. Bulletin of Entomological Research 37: 507— 29: Zolk, K. 1924. Paracodrus apterogynus Halid. biolo- gia kohta. Zur Biologie von Paracodrus aptero- gynus Halid. Tartu Ulikooli Entomoloogia-Katse- jaama Teadaanded 5: 3-10. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 69-74 NEW GENUS AND NEW SPECIES OF AMORBINI (HETEROPTERA: COREIDAE) FROM AUSTRALIA HARRY BRAILOVSKY AND GERASIMOS CASSIS (HB) Departamento de Zoologia, Instituto de Biologia UNAM, Apdo Postal No. 70- 153, México, 04510 D.E, México (e-mail: coreidae @ servidor.unam.mx); (GC) Australian Museum, 6’8 College Street, RO. Box A285, Sydney South, N.S.W. 2000, Australia. Abstract.—Kormiirania, n. gen., and two new species, K. magna and K. parva, col- lected in Eastern Australia are described in the tribe Amorbini (Coreidae). Habitus illus- trations, drawings of the male and female genitalia, and a key are provided. Key Words: Australia The tribe Amorbini (Hemiptera: Heter- optera: Coreidae) is represented by 7 genera in the Australian region: Acroelytron Mayr (1 species), Amorbus Dallas (15 species), Cneius Stal (1 species), Gelonus Stal (1 species), Kurnaina Distant (1 species), Tambourina Distant (1 species), and a new genus (2 species) (Brailovsky and Mon- teith, in press). The present paper adds one new genus and two new species of Amorbini from eastern Australia. For several years, the spe- cies discussed here have remained unnamed and undescribed in the hope that additional specimens would be found. However, it seems desirable to draw the attention of en- tomologists to these species in the hope that additional specimens will be collected and some information obtained on their ecology and food requirements. Two striking fea- tures of this new genus are the upturned juga forming a stout long horn or conical tubercle, and the mandibular plate expand- ed on a remarkable stout conical tubercle. The following abbreviations are used for the institutions cited in this paper: BPBM (Bernice P. Bishop Museum, Honolulu, Ha- wail); SAMA (South Australian Museum, Insecta, Heteroptera, Coreidae, Amorbini, new genus, new species, Eastern Adelaide); UNAM (Instituto de Biologia, Universidad Nacional Aut6noma de Méxi- CO). All measurements are given in millimeters. Kormiirania Brailovsky and Cassis, new genus Diagnosis.—This new genus resembles Gelonus Stal (1865) in having the head wider than long, tylus unarmed and extend- ing anteriorly to the juga, antenniferous tu- bercles unarmed, tibiae sulcate, not foliate, and abdominal sternite VII of the female with plica and fissura. It is easily distin- guished because it is the only known genus in the tribe Amorbini with the upturned juga forming a stout long horn or conical tubercle, and the mandibular plate striking- ly expanded on a large conical tubercle. Kormijirania has a stout antennal seg- ment I, short, and barely crested, antennal segments IJ and III almost cylindrical, bare- ly flattened, buccula anteriorly with a clear spine-like projection, pronotum slightly wider than long, and fore and middle fem- ora ventrally with small granules or tiny spine-like projections, never with a large and laminate subdistal spine. In Ge/onus the 70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON juga are flattened, the mandibular plate un- armed, antennal segments I to III uniformly cylindrical, not crested or flattened, buccula anteriorly rounded, pronotum clearly wider than long, and fore and middle femora ven- trally provided with a large and unique sub- distal laminate spine. A new genus (Brailovsky and Monteith, in press), is related to Gelonus and Kormi- jirania and is recognized by the laterally compressed tylus, which is projected up- ward as an acute projection, with femora unarmed, juga flat, mandibular plate un- armed, and antennal segments I to III cylin- drical. Description.—Macropterous, body stout, moderate sized. Head: Width across eyes greater than length, quadrate, dorsally flat, non declivent, barely pro- duced beyond antenniferous tubercles, with deep circular pit close to base of ty- lus; tylus unarmed, slightly deflexed; juga shorter than tylus, upturned to form a stout long horn or conical tubercle; anten- niferous tubercle unarmed, prominently produced, wide, separated by distance greater than their own width; sides of head in front of eyes almost straight; an- tennal segment I robust, thickest, barely crested, constricted basally; segments II and III stout, cylindrical, barely flattened; segment IV fusiform; antennal segment II longest, IV shortest, and III longer than I; ocelli conspicuous, closer to eyes, and lo- cated on an hypothetical line with lower margin slightly above lower margin of eyes; preocellar pit deep; eyes protruding; postocular tubercle markedly produced; buccula rounded, short, not projecting be- yond anterior third of antenniferous tu- bercles, slightly raised, anteriorly with a sharp spine-like projection, and posteri- orly closed; rostrum reaching posterior third of mesosternum or anterior third of metasternum; rostral segment III shortest, I longest, and II longer than IV; mandib- ular plate expanded on a strong conical tubercle or acute tooth; ventrally with a deep longitudinal groove along midline to receive first and anterior third of rostral segment II; mandibular plate with mark- edly stout conical tubercle. Thorax: Pronotum wider than long, tra- peziform, non declivent; collar wide; fron- tal angles produced forward as rounded teeth; humeral angles rounded, not exposed; anterolateral margins slightly emarginate, obliquely straight, finely serrate; posterolat- eral margins sinuate, entire; posterior bor- der concave, entire; callar region indistinct, transversely flat, separated along midline by an obscure longitudinal groove; posterior lobe with transverse ridge, distinctly raised; prosternum mesally with a slight depres- sion; mesosternum with deep longitudinal groove to receive rostrum; anterior third of mesosternum without lateral expansions; anterior lobe of metathoracic peritreme el- evated, irregularly reniform, posterior lobe sharp, small. Legs: Short; femora slightly incrassate; fore and middle femora densely granulate, with small spine-like projections both dor- sally and ventrally; hind femur slightly granulate, ventrally armed with two short subdistal spines or very tiny tubercles; tib- iae terete, sulcate. Scutellum: Triangular, longer than wide; apex truncated or subacute; disc flat. Hemelytra: Macropterous, almost reach- ing apex of last abdominal segment; costal margin emarginate; apical margin straight; apical angle short, barely reaching middle third of hemelytral membrane. Abdomen: Connexival segments reflexed above margin of hemelytron at rest; upper margin weakly serrate; posterior angles of connexival segments simple, not spinose; abdominal spiracles II to VII submarginal, closer to middle third. Male genitalia: Genital capsule: Simple; posteroventral border slightly concave, with posterolateral angles broadly produced (Figsmiy 2): Female genitalia: Abdominal sternite VII with plica and fissura; plica triangular, slightly elevated, apically subacute, and reaching anterior third of sternite VII. VOLUME 101, NUMBER 1 1mm Figs. 1—4. q/\ 1, 2. Male genital capsule of Kormijirania parva. |, Caudal view. 2, Lateral view. 3, 4, Female genital plates of K. magna. 3, Caudal view. 4, Lateral view. Genital segments: Gonocoxae I enlarged dorso-ventrally, in caudal view closed, in lateral view slightly convex, with upper Margin sinuate; paratergite VIII triangu- lar, with spiracle visible; paratergite IX square, larger than paratergite VIII (Figs. 3, 4). Integument: Body surface rather dull, al- most glabrous. Head, antennal segments I to III, pronotum, clavus, corium, legs, con- nexival segments, propleuron, mesopleu- ron, metapleuron, and pleural abdominal sterna densely granulate; prosternum, me- sosternum, metasternum, abdominal sterna, and genital plates almost smooth; posterior lobe of pronotum, clavus and corium, densely punctate. Etymology.—We are pleased to name this new genus for Dr. Nicholas A. Kor- milev, distinguished hemipterist. Type species.—Kormijirania parva Brai- lovsky and Cassis, new species. Kormijirania magna Brailovsky and Cassis, new species (Figs. 3-5) Description.—Measurements: Female: Head length 1.39; width across eyes 2.04; interocular space 1.30; interocellar space 0.46; preocular distance 1.05; length of an- tennal segments: I, 1.79; I, 2.13; III and IV mutilated. Pronotum: Total length 2.54, width across frontal angles 1.89; width across humeral angles 2.97. Scutellar length 1.51; width 1.24. Total body length 12.10. Female: Dorsal coloration: Head, anten- nal segments I and I (III and IV mutilated), anterior lobe of pronotal disk, clavus and corium dirty yellow with red-brown to chestnut-orange punctures and_ tubercles; posterior lobe of pronotal disc dirty yellow with black to red-brown punctures; scutel- lum dark brown to black; apical angle of corium almost black; hemelytral membrane dirty white, with veins, basal angle, and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON WZ Dorsal view of Kormijirania magna (@). Figs 5: VOLUME 101, NUMBER 1 Fig. 6. Dorsal view of Kormijirania parva (<¢). scattered spots pale brown; connexival seg- ments III to VIII light orange yellow, with upper margin of posterior third, and tuber- cles brown; connexival segment IX pale or- ange yellow, with black quadrate spot on middle third. Ventral coloration: Including rostral segments I to IV (apex of IV black), and legs dirty yellow with red-brown tu- bercles; abdominal sterna, and genital plates shiny yellow, with pale orange dis- coidal spots. Structure: Genital plates as in Figs. 3, 4. 74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Male: Unknown Type material—Holotype: ¢°, Australia, Queensland, Mackay (without additional data) (BPBM). Etymology.—Named for its large size; from the Latin word magnus. Kormiirania parva Brailovsky and Cassis, new species (Figs: 1; 2;/6) Description.—Measurements: Male: Head length 1.20; width across eyes 1.86; interocular space 1.17; interocellar space 0.43; preocular distance 0.96; length of an- tennal segments: I, 1.24; II, 1.76; Il, 1.39; IV, 1.14. Pronotum: Total length 1.70; width across frontal angles 1.51; width across humeral angles 1.96. Scutellar length 0.98; width 0.80. Total body length 9.00. Female: Head length 1.35; width across eyes 1.92; interocular space 1.20; interocel- lar space 0.43; preocular distance 0.96; length of antennal segments: I, 1.36; II, 1.87; II, 1.54; IV, mutilated. Pronotum: To- tal length 2.10; width across frontal angles 1.70; width across humeral angles 2.25. Scutellar length 1.24; width 1.02. Total body length 10.00. Male: Dorsal coloration: Yellow with punctures orange hazel; ocellar tubercle red brown; antennal segments I to III yellow with red brown tubercles; segment IV with anterior half whitish yellow, and posterior half orange hazel; humeral angles red brown; apex of scutellum whitish yellow; hemelytral membrane dirty white, with veins, basal angle, and scattered spots pale brown; connexival segments II to VII yel- low with upper margin of posterior third or entirely the posterior third reddish brown; abdominal segments I to VI light orange yellow and VII yellow with H-shaped dark spot. Ventral coloration: Included rostral segments I to IV (apex of IV black), buc- cula, and anterior and posterior lobe of metathoracic peritreme light yellow with some red-brown tubercles; genital capsule dark brown with three irregular yellow spots; legs yellow with red to pink tuber- cles. Structure: Genital capsule as in Figs. 1-2. Female: Similar to male. Type material.—Holotype: 3, Australia, Cairns District, Col. A. M. Lea (without ad- ditional data) (SAMA). Paratype: 1 &, same data as holotype (SAMA). Both spec- imens glued on the same card. Etymology.—Named for its small size; from the Latin word parva, rather small. KEY TO SPECIES OF KORMIJIRANIA 1. Femora dirty yellow without red to pink tuber- cles (Fig. 5); antennal segment I longer than 1.70, and II longer than 2.00 Stoeneeee magna Brailovsky and Cassis, new species — Femora yellow with red to pink tubercles (Fig. 6); antennal segment I shorter than 1.40, and Ik shorterthan 9085S 5 ae see | rene Panacea parva Brailovsky and Cassis, new species ACKNOWLEDGMENTS We thank Gordon E Gross (SAMA) and Gordon M. Nishida (BPBM) for the loan of specimens, and Ernesto Barrera (UNAM), Albino Luna (UNAM), and Jesus Contreras for preparation of dorsal view illustrations and male and female genitalia. LITERATURE CITED Brailovsky, H. and G. B. Monteith. In press. A new genus of Amorbini (Heteroptera: Coreidae) from Australia, with two new species. Memoirs of the Queensland Museum. Stal, C. 1865. Hemiptera Africana II. Stockholm: Nordstedtiana. 200 pp. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 75-85 LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF TRUPANEA ARIZONENSIS MALLOCH (DIPTERA: TEPHRITIDAE) ON TRIXIS CALIFORNICA KELLOGG VAR. CALIFORNICA (ASTERACEAE) IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN AND JEFFREY A. TEERINK Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e- mail: rgoeden @ucrac 1 .ucr.edu) Abstract.—Trupanea arizonensis Malloch is a monophagous, univoltine fruit fly (Dip- tera: Tephritidae) infesting flower heads of Trixis californica Kellogg var. californica (Asteraceae) in southern California. The egg, first- and third-instar larvae, and puparia are described and figured for the first ttme. As with each of the eight other 7rupanea species previously studied, the lateral spiracular complex of the third instar is unique to T. arizonensis, comprising three verruciform sensilla on the meso- and metathorax and two verruciform sensilla on each abdominal segment. The third instar of 7. arizonensis also differs from those of other Trupanea species previously studied in that both the meso- and metathorax are circumscribed by verruciform sensilla. The life cycle is of the aggre- gative type; whereby, the adults are long-lived and comprise the over-summering and over-wintering stage that returns with winter rainfall to aggregate on regrowing host plants for mating, and for oviposition during spring in the preblossom flower heads. The first instar tunneled into a single floral tube or ovule of a single immature floret, and each second and third instar continued its development by feeding principally on sap conducted to the excavated distal end of the same, then slightly stunted ovule. Pupariation occurs in the open flower heads, within which an average of 13% of the achenes were destroyed among heads containing puparia. Pteromalus sp. (Hymenoptera: Pteromalidae) was reared from individual puparia of 7. arizonensis as a solitary, larval-pupal endoparasitoid. Key Words: Insecta, Trupanea, Asteraceae, nonfrugivorous Tephritidae, biology, tax- onomy of immature stages, flower-head feeding, monophagy, host-plant range, parasitoid Trupanea arizonensis Malloch (Diptera: Tephritidae) is a rarely collected, monoph- agous or nearly monophagous species be- longing to one of the larger and more wide- spread genera of nonfrugivorous fruit flies in North America and California (Foote and Blanc 1963, Foote et al. 1993). But, being of little or no economic importance, most species of Trupanea remained little known (Foote 1960, Foote et al. 1993) until de- tailed life histories were published for nine species from southern California (Cavender and Goeden 1982; Goeden 1987, 1988; Goeden and Teerink 1997, 1998; Goeden et al. 1998a, b; Headrick and Goeden 1991; Knio et al. 1996b), along with descriptions of the immature stages of eight of these species (Cavender and Goeden 1982, Goe- den and Teerink 1997, 1998; Goeden et al. 1998a, b; Headrick and Goeden 1991, Knio et al. 1996a, Teerink and Goeden 1998). This paper describes the life history and 76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON most immature stages of a tenth species, T. arizonensis. MATERIALS AND METHODS This study was based in large part on dis- sections of subsamples of flower heads of Trixis californica Kellogg var. californica (Asteraceae) infested by 7. arizonensis from samples collected during 1991—1995 in the low-elevation Colorado (northern Sonoran) Desert and high-elevation, eastern Mojave Desert in southern California in the manner described by Goeden (1985, 1992). The principal study sites in the Colorado Desert were Valliceto Valley at the mouth of Smugglers Canyon, 440-m_ elevation, San Diego Co.; the Edmund C. Jaeger Na- ture Preserve at 847 m, Desert Center, Riv- erside Co.; and Chino Canyon, 1 km NW of Palm Springs, at 270 m, Riverside Co. One-liter samples of excised, immature and mature flower heads containing eggs, lar- vae, and puparia were transported in cold- chests in an air-conditioned vehicle to the laboratory and stored under refrigeration for subsequent dissection, photography, de- scription, and measurement. Nine eggs, 18 first- and 16 third-instar larvae, and six pu- paria dissected from flower heads were pre- served in 70% EtOH for scanning electron microscopy (SEM). Additional puparia were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity chambers at room temperature for adult and parasitoid emergence. Specimens for SEM were hydrated to distilled water in a decreasing series of acidulated EtOH. They were osmicated for 24 h, dehydrated through an increasing series of acidulated EtOH and two, I-h immersions in Hexa- methlydisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, and studied with a Philips XL30-FEG scanning electron microscope in the Insti- tute of Geophysics and Planetary Physics, University of California, Riverside. Most adults reared from isolated puparia were individually caged in 850-ml, clear- plastic, screened-top cages with a cotton wick and basal water reservoir and provi- sioned with a strip of paper toweling im- pregnated with yeast hydrolyzate and su- crose. These cages were used for longevity studies in the insectary of the Department of Entomology, University of California, Riverside, at 25 + 1°C, and 14/10 (L/D) photoperiod. Plant names used in this paper follow Hickman (1993) and Bremer (1994); te- phritid names and adult terminology follow Foote et al. (1993). Terminology and tele- graphic format used to describe the imma- ture stages follow Knio et al. (1996a), Goe- den and Teerink (1997, 1998), Goeden et al. (1998a, b), and Teerink and Goeden (1998), and our earlier works cited therein. Means + SE are used throughout this paper. Voucher specimens of 7. arizonensis and its parasitoids reside in the research collections of RDG; preserved specimens of eggs, lar- vae and puparia are stored in a separate col- lection of immature Tephritidae acquired by JAT and now maintained by RDG. RESULTS AND DISCUSSION TAXONOMY Adult.—Trupanea arizonensis was first described by Malloch (1942) as Trypanea arizonensis. Malloch (1942), Foote (1960), Foote and Blanc (1963), and Foote et al. (1993) pictured the wing pattern, which Foote (1960) described as not sexually di- morphic, but essentially alike in every im- portant character in the female and male. However, the variable nature of this wing pattern, especially among males, initially caused Goeden and Ricker (1989) to mis- identify specimens reared from Trixis cali- fornica as Trupanea actinobola (Loew), an- other variable species (Foote and Blanc 1963, Foote et al. 1993, Goeden et al. 1998b). Goeden (1992) corrected this error and re-identified the flies from 7. califor- nica as T. arizonensis; moreover, Goeden et al. (1998b) further supported this identifi- cation based on the different tribal- and sep- arate, subtribal-level, host-plant affiliations VOLUME 101, NUMBER 1 of the three biotypes of 7. actinobola in southern California, which exclude 7rixis californica in the tribe Mutisieae (see be- low). Nine (10.6%) of 85 ¢, but none of 65 2 of Trupanea arizonensis in the research col- lection of RDG reared from flower heads of Trixis californica have only one dark ray, not two rays, contrary to Foote et al. (1993), extending into cell dm from vein CuA,. Moreover, the single, remaining, what is otherwise called the distal ray (Foote et al. 1993), is broken in cell dm and does not reach vein CuA, in 39 ¢ (45.9%) and 18 2 (27.7%). There is no dark spot on vein CuA, or on the wing margin in line with the broken distal ray in an additional 10 6 (11.8%) and 3 & (4.6%), or with such a dark spot on CuA, or on the wing margin in line with the broken distal ray in an ad- ditional 2 6 (2.4%) and 3 2 (4.6%), re- spectively. The proximal ray usually ex- tends, though sometimes very limitedly, into cell dm in 76 6 (89%) and all 65 Q, and there is usually (not always, Foote et al. 1993) a dark spot on vein CuA, in line with it; however, in 3 2 (4.6%) the broken proximal ray is reduced to a spot in the middle of cell dm in line with a spot on vein CuA,, and another 2 6 (2.3%) and 1 2 (1.5%) lack a dark spot in line with it. Therefore, all females and most males of 7. arizonensis can still be distinguished from other Trupanea spp. by use of the separate keys for the two sexes in Foote et al. (1993) if one is aware of the above variations and perceives two dark rays, however abbrevi- ated, extending into cell dm, neither of which necessarily crosses vein CuA,, much less continues to the hind wing margin. Males with only one dark ray extending into cell dm from vein CuA,, like the nine noted above, will run to 7. actinobola, as they did earlier for Goeden and Ricker (1989). Couplet 8 (pp. 421 and 424 in Foote et al. 1993) distinguishing 7. maculigera Foote from 7. arizonensis apparently re- mains valid; although the angles formed by the proximal and distal rays, not the dis- HU SDP eee | a May 3.0 _148x 31.1 Spot Magn rae spot Magr wo /-_-——————4_ 1078S. 0 290bx 1.2 ‘ Fig. 1. Egg of Trupanea arizonensis: (A) habitus, pedicel to left; (B) pedicel. tances between them along vein CuA,, should be compared with specimens of T. arizonensis having proximal or distal rays or both broken in cell dm. Immature stages.—The immature stages of T. arizonensis heretofore have neither been described nor illustrated. Egg: Seventy eggs of T. arizonensis dis- sected from heads of 7. californica were white, opaque, smooth; elongate-ellipsoi- dal, 0.70 + 0.006 (range, 0.54—0.85) mm long, 0.15 + 0.002 (range, 0.12—0.23) mm wide, smoothly rounded at tapered basal end (Fig. 1A), pedicel 0.02 mm long, with single row of subcircular aeropyles (Fig. 1B). The egg of T. arizonensis is narrow like the eggs of T. actinobola (Goeden et al. 1998b) and 7. pseudovicina (Goeden and Teerink 1998), but is much longer than both of these species. The egg body in 7. ari- 78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 100 jim Fig. 2. First instar of Trupanea arizonensis: (A) habitus, anterior end to left; (B) gnathocephalon, anterior view, 1, dorsal sensory organ, 2, terminal sensory organ, 3, pit sensory organ, 4, lateral sensory organ, 5, stomal sense organ; (C) gnathocephalon, ventral view, 1, anterior sensory lobe, 2, mouth hook, 3, median oral lobe, 4, 9} labial lobe, 5, pit sensillum; (D) caudal segment, 1, intermediate sensory complex. zonensis tapers smoothly into the pedicel, more so than in the other Trupanea species previously studied (Goeden and Teerink 1997, 1998; Goeden et al. 1998a, b; Knio et al. 1996a; Teerink and Goeden 1998). The pedicel is similar to these other con- geners in having a single row of aeropyles (Goeden and Teerink 1997, 1998; Goeden et al. 1998a, b; Teerink and Goeden 1998), with the exception of 7. bisetosa (Coquil- lett), which has one or two rows (Knio et al. 1996a). First instar: White, elongate-cylindrical, rounded anteriorly and posteriorly (Fig. 2A), minute acanthae circumscribe inter- segmental lines (Fig. 2A); gnathocephalon smooth (roughness in Fig. 2 is an artifact), lacking rugose pads (Fig. 2C); dorsal sen- stelex sensillum, , rima, 3, interspiracular process, 4, sory organ a dome-shaped papilla (Fig. 2B- 1); anterior sensory lobe (Fig. 2C-1) bears terminal sensory organ (Fig. 2B-2), pit sen- sory organ (Fig. 2B-3), lateral sensory or- gan (Fig. 2B-4), and supralateral sensory organ; stomal sense organ indistinct (Fig. 2B-5); mouth hooks bidentate (Fig. 2C-2); median oral lobe laterally flattened (Fig. 2C-3), labial lobe (Fig. 2C-4) attached to median oral lobe; pit sensillum laterad of anterior sensory lobe (Fig. 2C-5); prothorax lacking rugose pads, few minute acanthae ventrad of mouth lumen; stelex sensilla cir- cumscribe caudal segment in 2-dorsal, 4- ventral arrangement (Fig. 2D-1); posterior spiracular plates bear two ovoid rimae (Fig. 2D-2), and four, spatulate interspiracular processes (Fig. 2D-3); intermediate sensory VOLUME 101, NUMBER 1 complex consists of a medusoid and stelex sensillum (Fig. 2D-4). The first instar of 7. arizonensis is very similar to other congeners previously stud- ied in habitus and sensory structures (Goe- den and Teerink 1998; Goeden et al. 1998a, b; Knio et al. 1996a; Teerink and Goeden 1998). The lateral spiracular complex was not seen. There are slight differences in the interspiracular processes. In 7. jonesi, the interspiracular processes are greatly re- duced (Goeden et al. 1998a), in T. arizo- nensis and T. conjuncta the processes are single and spatulate (Teerink and Goeden 1998), and in 7. pseudovicina Hering, T. actinobola, T. bisetosa and T. nigricornis (Coquillett), the interspiracular processes are divided, each with 1—4 branches (Goe- den and Teerink 1997; Goeden et al. 1998b; Knio et al. 1996a). Third instar: White, elongate-cylindri- cal, tapering anteriorly, rounded posteriorly, minute acanthae circumscribe intersegmen- tal lines in bands increasing in width pos- teriorly (Fig. 3A); gnathocephalon conical, rugose pads dorsal and lateral to mouth lu- men (Fig. 3B-1); dorsal sensory organ a dome-shaped papilla (Fig. 3B-2, 3C-1); an- terior sensory lobe (Fig. 3B-3) bears the ter- minal sensory organ (Fig. 3C-2), pit sen- sory organ (Fig. 3C-3), lateral sensory organ (Fig. 3C-4), and supralateral sensory organ (Fig. 3C-5); stomal sense organ ven- trolaterad of anterior sensory lobe (Fig. 3B- 4, 3C-6); mouth hooks tridentate (Fig. 3B- 5, 3D-1); median oral lobe tapered anteri- orly (Fig. 3D-2); six pit sensilla circum- scribe gnathocephalon posterior to rugose pads (Fig. 3B-6); minute acanthae circum- scribe anterior margin of prothorax (Fig. 3E-1); rugose pads (Fig. 3E-2) and a single row of verruciform sensilla (Fig. 3E-3) cir- cumscribe prothorax posteriorad of minute acanthae; additional verruciform sensilla on dorsal half of prothorax (Fig. 3E-4); ante- rior thoracic spiracle on posterior margin of prothorax bears three ovoid papillae (Fig. 3E-5), meso- and metathorax circumscribed by verruciform sensilla; meso- and meta- 79 thoracic lateral spiracular complexes consist of a spiracle (Fig. 3F-1), and three verru- ciform sensilla (Fig. 3F-2); abdominal lat- eral spiracular complex consists of a spira- cle (Fig. 3G-1) and two verruciform sensilla (Fig. 3G-2); caudal segment circumscribed by minute acanthae (Fig. 3H-1); posterior spiracular plates bear three ovoid rimae, ca. 0.03 mm in length (Fig. 3H-2), and four interspiracular processes, each with 3-5 branches, longest measuring 0.02 mm (Fig. 3H-3); intermediate sensory complex con- sists of a stelex sensillum and a medusoid sensillum (not shown). The third instar of 7. arizonensis is sim- ilar in general habitus to 7. pseudovicina in being elongate-cylindrical, maybe even more attenuated (Goeden and Teerink 1998). As with the third instars of each of the eight other Trupanea species previously studied, the lateral spiracular complex is unique to 7. arizonensis, with three verru- ciform sensilla in the meso- and metathorax and two verruciform sensilla in the abdom- inal segments (Goeden and Teerink 1997, 1998; Goeden et al. 1998a, b; Headrick and Goeden 1991; Knio et al. 1996a; Teerink and Goeden 1998). There are other differ- ences between individual species. Trupanea arizonensis differs from T. actinobola in lacking elongated integumental petals; 7. arizonensis has fewer rugose pads circum- scribing the prothorax, and the prothorax is completely circumscribed by minute acan- thae (Goeden et al. 1998b). Also, T. arizo- nensis differs from the other Trupanea spe- cies previously studied, in that the meso- and metathorax are circumscribed by ver- ruciform sensilla; whereas, only the meso- thorax in 7. nigricornis was circumscribed by verruciform sensilla (Goeden and Teer- ink 1997, 1998; Goeden et al. 1998a, b; Headrick and Goeden 1991; Knio et al. 1996a; Teerink and Goeden 1998). Trupa- nea arizonensis differs from its sympha- gous congener in Trixis californica, T. con- juncta, in not being finely punctate nor bar- rel-shaped, and TJ. arizonensis also has 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Spot Magn WD SOT mm 2) 2 Acc.V Spot’Magn igs Ry fAct.V\) Spot Magn { WD Sd 205m a 10.0 KV"3.0 _3070x 37.6 _ ace 0.0 kV3'0_ 1047x376 = Acc.V Spot Magn 3 100 kV 30_ 293x es r Acc SpéhMagn© WD°&—————+1; j 001m H 10:0 KV.3.0.. 362x 17:2 Fig. 3. Third instar of Trupanea arizonensis: (A) habitus, anterior end to left: (B) gnathocephalon, antero- lateral view, 1, rugose pads, 2, dorsal sensory organ, 3, anterior sensory lobe, 4, stomal sense organ, 5, mouth hook, 6, pit sensillum; (C) anterior sensory lobe, 1, dorsal sensory organ, 2, terminal sensory organ, 3, pit sensory organ, 4, lateral sensory organ, 5, supralateral sensory organ, 6, stomal sense organ; (D) gnathocephalon, ventral view, 1, mouth hook, 2, median oral lobe; (E) gnathocephalon, prothorax, lateral view, 1, minute acanthae, VOLUME 101, NUMBER 1 fewer papillae in the anterior thoracic spi- racle (Teerink and Goeden 1998). Puparium: Black, elongate-cylindrical, minute acanthae circumscribe intersegmen- tal lines, (Fig. 4A); anterior end bears the invagination scar (Fig. 4B-1), and anterior thoracic spiracles (Fig. 4B-2); caudal seg- ment circumscribed by minute acanthae (Fig. 4C-1); posterior spiracular plates bear three ovoid rimae (Fig. 4C-2) and four in- terspiracular processes, each with 3-5 branches (Fig. 4C-3). Sixty-four puparia av- eraged 2.68 + 0.02 (range, 2.28—3.24) mm in length; 0.92 + 0.01 (range, 0.76—1.16) mm in width. DISTRIBUTION AND Hosts The distribution of 7. arizonensis mapped in North America north of Mexico by Foote et al. (1993) is confined to several, mostly U.S. southern border locations in Arizona, California, and Texas. Goeden and Ricker (1989) reported 7. arizonensis as T. actinobola from Trixis californica, and as mentioned above, Goe- den (1992) corrected this misidentification and thus provided the first and sole host- plant record for 7. arizonensis. This host- plant belongs to the subtribe Nassauviinae in the tribe Mutisieae of the Asteraceae (Hickman 1993, Bremer 1994); this tribe rarely occurs in North America (Bremer 1994), and is represented by only three spe- cies in California (Munz 1974), the second of which, Acourtia microcephala de- Candolle, does not host 7. arizonensis (Goeden and Headrick 1991). The third plant species is the rare, unsampled, xero- phytic shrub, Hecastocleis shockleyi A. Gray (Hickman 1993, Bremer 1994). In comparison, 7. actinobola, apparently is confined to three subtribes of the different tribe Astereae in California (Goeden and — 81 Teerink 1998b). Accordingly, T. arizonen- sis may be either a true monophage (one host-plant species) on Trixis californica, which occurs as far east as Texas in North America north of Mexico (Hickman 1993), or a near-monophage (one host-plant genus) on one or more of the 50 other congeners in the southwestern United States, Central, and South America and West Indies (Bre- mer 1994). Like several other tephritid spe- cies that we have studied, e.g., Trupanea conjuncta (Goeden 1987), T. pseudovicina Hering (Goeden and Teerink 1998), and To- moplagia cressoni Aczél (Goeden and Headrick 1991), Trupanea arizonensis rep- resents a native southern California tephri- tid closely associated with a native host- plant, which is primarily distributed in Mexico and southward, where these tephri- tids remain little known. BIOLOGY Egg.—In 38 closed, preblossom, imma- ture flower heads, all 130 eggs were insert- ed pedicel-last between the tips of the phyl- laries, perpendicular to the receptacle, and among or within the florets and pappus (Fig. 5A). The diameters of the receptacles of 32 of these flower heads containing eggs averaged 2:6 2,0.) (@ange, 1.3—3.7) mm, and the 38 infested heads contained an av- erage of 3.4 + 0.3 (range, 1-9) eggs ovi- posited, mostly singly, or side-by-side in pairs (Figs. 5A, B), but also in groups of up to five, by one or more females. Larva.—Upon eclosion, first instars tun- neled into a single floral tube or ovule of an immature floret (Fig. 5C). An average of 2.1 + 0.2 (range, 1—4) first instars was found feeding within 22 closed, preblossom flower heads. The receptacles of these heads averaged 3.1 + 0.2 (range, 2.2—4.6) mm in diameter with an average of 18 + 2 2, rugose pads, 3, verruciform sensillum, 4, verruciform sensillum, 5, anterior thoracic spiracle; (F) metathorax, 1, spiracle, 2, verruciform sensilla; (G) fifth abdominal segment, 1, spiracle, 2, verruciform sensilla; (H) caudal segment, 1, minute acanthae, 2, rima, 3, interspiracular process. 82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON WDA eed 33.3 500 jim cc.V Spot Magn 0.0kV 3.0° 38x Fig. 4. Puparium of Trupanea arizonensis: (A) habitus, anterior end to right; (B) anterior end, 1, in- vagination scar, 2, anterior thoracic spiracle; (C) caudal segment, 1, minute acanthae, 2, rima, 3, interspiracular process. (range, 15—24) florets, of which an average of only 1.6 + 0.2 (range, 1-3) florets, or 9% (range, 4—17%), were damaged. No re- ceptacles within these 14 infested flower heads was pitted by first-instar feeding. Second instars fed solitarily at the distal ends of individual ovules or soft achenes within different florets of separate preblos- som and open flower heads, respectively. Their mouthparts were directed towards the receptacles. Receptacles of 16 flower heads containing second instars were not pitted and averaged 3.4 + 0.2 (range, 2.3—4.8) mm in diameter. These flower heads con- tained an average of 1.6 + 0.2 (range, 1-3) second instars that had damaged an average of 1.8 + 0.2 (range, 1—4) ovules, or 11% (range, 6—22%) of an average total of 17 + 0.8 (range, 12—20) ovules per flower head. Most third instars also confined their feeding to the same single, separate ovule or soft achene at the centers, and less com- monly at the margins, of preblossom or open flower heads (Fig. 5D). Third instars fed with their long axes oriented perpendic- ular to and mouthparts directed towards the receptacles, and on the distal parts of the ovules or soft achenes, well above the re- ceptacles (Fig. 5D). In 28 flower heads av- eraging 4.0 + 0.1 (range, 3.1—5.0) mm in diameter and containing an average of 1.5 + 0.2 (range, 1—5) third instars, an average of 1.6 + 0.2 (range, 1-5) ovules were dam- aged, or 9% (range, 4—25%). Thus, each larva confined its feeding to the distal parts of a single floret, at most including corolla tube and contents, pappus, and upper part of the ovule (Fig. 5D); the ovule continued to grow and functioned as a nutrient sink (Harris and Shorthouse 1996) augmented by the larva which fed mainly on sap drawn to the cuplike depression in the distal end of the ovule (Fig. 5D). This mode of feed- ing is the least damaging to host-plant re- production, as well as the most exquisitely evolved manner of sap-feeding yet reported among florivorous Tephritidae (Headrick and Goeden 1998). This manner of larval feeding, first reported by Headrick and Goeden (1990b), apparently is facultative, undetected, or absent in some florivorous species, e.g., Paracantha gentilis Hering (Headrick and Goeden 1990a, b), Trupanea nigricornis and T. bisetosa (Knio et al. VOLUME 101, NUMBER | Figes: 83 Se said igo 3p, ia RE a \ Witbac Sy a _ cam § st Life stages of Trupanea arizonensis in or on Trixis californica: (A) one pair of eggs (arrow) inserted among immature florets in closed, preblossom flower head; (B) two pairs of eggs inserted among florets in closed, preblossom flower head; (C) first instar tunneling in single floret (arrow); (D) two third instars feeding on separate soft achenes; (E) single puparium atop stunted achene in flower head; (F) adult female at rest on flower head. Lines = | mm. (1996b); although it also has been reported in some gallicolous species, e.g., Aciurina thoracica Curran (Headrick and Goeden 1993), and is facilitated by specialized mouthparts in larval Tephritinae (Headrick and Goeden 1990a, 1993, 1998). Upon completing feeding, the larvae ori- ented their anterior ends away from the re- ceptacles, retracted their mouthparts, and pupariated (Fig. 5E). Pupa.—Flower heads containing puparia (Fig. 5E) contained the greatest amounts of damage produced by the seed-feeding lar- vae of T. arizonensis. The receptacles of 41 open and postblossom flower heads con- taining puparia averaged 4.4 + 0.1 (range, 2.9—6.2) mm in diameter and bone an av- erage total of 19.2 + 0.6 (12-26) soft achenes, of which an average of 2.2 + 0.3 (range, 1—9) soft achenes or 13% (range, 4— 58%) were damaged. Again, no receptacles were fed upon. These heads contained an average of 1.8 + 0.2 (range, 1—7) puparia. Most puparia of 7. arizonensis were found near the centers of the flower heads, and all had their anterior ends facing away from 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the receptacles, and their long axes were perpendicular to the receptacles (Fig. 5E). Adult.—Adults (Fig. 5F) emerged from mature, postblossom flower heads, and were long-lived under insectary conditions, as 15 unmated males lived an average of 89 + 14 (range, 11—194) days, and six virgin females averaged 126 + 7 (range, 110—150) days. Like several other, monophagous and nearly monophagous congeners studied (Goeden 1988, Goeden and Teerink 1997, 1998), the longevities of these flies were among the longer averages and maxima for adults that we have recorded for native spe- cies of nonfrugivorous Tephritidae from southern California. Such longevities are commensurate with the aggregative type of life cycle ascribed below to this tephritid. The premating and mating behaviors of T. arizonensis were not studied in the field, and again, like most congeners that we have studied, adults would not mate in petri dish arenas otherwise so useful with many other, noncongeneric, nonfrugivorous species (Headrick and Goeden 1994). Seasonal history.—The life cycle of T. arizonensis in southern California appears to follow an aggregative pattern in which the long-lived adults in reproductive dia- pause Over-summer in riparian habitats and mountain meadows. They return to lower elevations in the fall, and following the on- set of winter rainfall, aggregate on Trixis californica in the low-elevation Colorado Desert in winter (January—February) to mate and later to oviposit (March—April) (Headrick and Goeden 1994). A single an- nual generation is produced each year and most of the life span of 7. arizonensis is spent as an adult. Natural enemies.—Preromalus sp. (Hy- menoptera: Pteromalidae) was reared from individual puparia of 7. arizonensis as a solitary, larval-pupal endoparasitoid; how- ever, at least some of the other species of parasitoids reported from Tomoplagia cres- soni Aczél (Goeden and Headrick 1991) and Trupanea conjuncta (Goeden 1987), which co-occur in symphagy in the heads of Trixis californica (Goeden and Ricker 1989, Goeden 1997), probably also parasit- ize T. arizonensis. ACKNOWLEDGMENTS We thank Andrew C. Sanders, Curator of the Herbarium, Department of Botany and Plant Sciences, University of California, Riverside, for identification of plants men- tioned in this paper. The parasitoids were identified by Harry E. Andersen, Huntington Beach, California. We also are grateful to Kristine Gilbert for technical as- sistance, and to D. H. Headrick for his help- ful comments on an earlier draft of this pa- per. LITERATURE CITED Bremer, K. 1994. Asteraceae Cladistics & Classifica- tion. Timber Press, Inc. Portland, Oregon. Cavender, G. L. and R. D. Goeden. 1982. Life history of Trupanea bisetosa (Diptera: Tephritidae) on wild sunflower in southern California. Annals of the Entomological Society of America 75: 400— 406. Foote, R. H. 1960. A revision of the genus Trupanea in America north of Mexico. United States De- partment of Agriculture Technical Bulletin 1214. 29 pp. Foote, R. H. and FL. Blanc. 1963. The fruit flies or Tephritidae of California. Bulletin of the Califor- nia Insect Survey 7, 117 pp. Foote, R. H., EF L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York. Goeden, R. D. 1985. Host-plant relations of Trupanea spp. (Diptera: Tephritidae) in southern California. Proceedings of the Entomological Society of Washington 87: 564-571. 1987. Life history of Trupanea conjuncta (Adams) on Trixis californica Kellogg in southern California (Diptera: Tephritidae). Pan-Pacific En- tomologist 63: 284-291. 1988. Life history of Trupanea imperfecta (Coquillett) on Bebbia juncea (Bentham) Greene in the Colorado Desert of southern California (Diptera: Tephritidae). Pan-Pacific Entomologist 64: 345-351. . 1992. Analysis of known and new host re- cords for Trupanea from California (Diptera: Te- phritidae). Proceedings of the Entomological So- ciety of Washington 94: 107-118. . 1997. Symphagy among florivorous fruit flies (Diptera: Tephritidae) in southern California, pp. VOLUME 101, NUMBER 1 39-52 In Dettner, K., G. Bauer, and W. VOkI, eds., Vertical Food Web Interactions: Evolutionary Pat- terns and Driving Forces. Ecological Studies 130, Springer-Verlag, Heidelberg, Germany. Goeden, R. D. and D. H. Headrick. 1991. Notes on the biology, hosts, and immature stages of To- moplagia cressoni (Aczél) in southern California (Diptera: Tephritidae). Proceedings of the Ento- mological Society of Washington 73: 549-558. Goeden, R. D. and D. W. Ricker. 1989. Phytophagous insect faunas of the desert shrubs, Bebbia juncea and Trixis californica in southern California. An- nals of the Entomological Society of America 82: 325-321. Goeden, R. D. and J. A. Teerink. 1997. Life history and description of immature stages of Trupanea signata Foote (Diptera: Tephritidae) on Gnaphal- ium luteo-album L. in southern California. Pro- ceedings of the Entomological Society of Wash- ington 99: 748-755. . 1998. Life history and description of imma- ture stages of Trupanea pseudovicina Hering (Diptera: Tephritidae) on Porophyllum gracile Bentham (Asteraceae) in southern California. Pro- ceedings of the Entomological Society of Wash- ington 100: 361-372. Goeden, R. D., J. A. Teerink, and D. H. Headrick. 1998a. Life history and description of immature stages of Trupanea jonesi Curran (Diptera: Te- phritidae) on native Asteraceae in southern Cali- fornia. Proceedings of the Entomological Society of Washington 100: 126-140. . 1998b. Life history and description of im- mature stages of Trupanea actinobola (Loew) (Diptera: Tephritidae) on Acamptopappus sphaer- ocephalus (Harvey and Gray) Gray (Asteraceae) in southern California. Proceedings of the Ento- mological Society of Washington 100: 674-688. Harris, P. and J. D. Shorthouse. 1996. Effectiveness of gall inducers in weed biological control. Canadian Entomologist 128: 1021-1055. Headrick, D. H. and R. D. Goeden. 1990a. Description of the immature stages of Paracantha gentilis (Diptera: Tephritidae). Annals of the Entmological Society of America 83: 220-229. 85 . 1990b. Resource utilization by larvae of Par- acantha gentilis (Diptera: Tephritidae) in capitula of Cirsium californicum and C. proteanum (As- teraceae) in southern California. Proceedings of the Entomological Society of Washington 92: 512-520. . 1991. Life history of Trupanea californica Malloch (Diptera: Tephritidae) on Gnaphalium spp. in southern California. Proceedings of the En- tomological Society of Washington 93: 559-570. . 1993. Life history and description of imma- ture stages of Aciurina thoracica Curran (Diptera: Tephritidae) on Baccharis sarothroides Gray in southern California. Proceedings of the Entomo- logical Society of Washington 95: 210—222. 1994. Reproductive behavior of California fruit flies and the classification and evolution of Tephritidae (Diptera) mating systems. Studia Dip- terologica 1(2): 194-252. . 1998. The biology of nonfrugivorous tephritid fruit flies. Annual Review of Entomology 43: 217-241. Hickman, J. C. (ed.) 1993. The Jepson Manual. Uni- versity of California Press. Berkeley and Los An- geles. Knio, K. M., R. D. Goeden, and D. H. Headrick. 1996a. Descriptions of immature stages of Tru- panea nigricornis and T. bisetosa (Diptera: Te- phritidae) from southern California. Annals of the Entomological Society of America 89: 1-11. . 1996b. Comparative biologies of the cryptic, sympatric species, 7rupanea nigricornis and T. bisetosa (Diptera: Tephritidae) in southern Cali- fornia. Annals of the Entomological Society of America 89: 252-260. Malloch, J. R. 1942. Notes on two genera of American flies of the family Trypetidae. Proceedings of the United States National Museum 92(3123): 1-20. Munz, P. A. 1974. A Flora of Southern California. Uni- versity of California Press, Berkeley and Los An- geles. Teerink, J. A. and R. D. Goeden. 1998. Description of immature stages of Trupanea conjuncta Adams (Diptera: Tephritidae). Proceedings of the Ento- mological Society of Washington 100: 431—438. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 86-93 TWO NEW SPECIES OF AGARODES BANKS (TRICHOPTERA: SERICOSTOMATIDAE) FROM SOUTHEASTERN UNITED STATES A. C. KETH AND S. C. HARRIS Department of Biology, Clarion University, Clarion, PA 16214, U.S.A. (e-mail: harris @ mail.clarion.edu) Abstract.—Two new species, Agarodes logani, from the eastern panhandle of Florida, and Agarodes tuskaloosa, from west-central Alabama, are described, illustrated, and com- pared to congeners. The genus now totals twelve species, all restricted to eastern North America. Key Words: Trichoptera, Sericostomatidae The genus Agarodes Banks is restricted to eastern North America and ranges from southern Canada to the southern United States (Ross and Scott 1974). Agarodes lar- vae prefer smaller, spring-fed streams with a medium current and sandy substrate but they have been occasionally collected in larger streams fed by surface water and in the sandy, depositional areas of lakes. Ten species of Agarodes were previously recognized (Harris 1987) with all but A. distincta Ulmer and A. grisea Banks re- stricted to southeastern United States (Ross and Scott 1974). In Alabama, five species occur, A. crassicornis Walker, A. libalis Ross and Scott, A. alabamensis Harris on the Coastal Plain; A. grisea Banks in north- ern Alabama; and A. stannardi Ross in a restricted, northwestern region of the state. To this list from Alabama an apparently rare, new species is added, A. tuskaloosa. This new species is currently known only from the type locality, on a small, spring- fed stream at the northern edge of the Coastal Plain. In Florida, three species oc- cur, A. crassicornis Walker and A. libalis Ross and Scott at scattered localities; and A. ziczac Ross and Scott known only from , Agarodes, new species, southeast the type locality in the central panhandle region. To this Florida list, A. /ogani, a new species, is added. This species is currently known only from the type locality, a small spring-run in a deep ravine, in the eastern panhandle region. Agarodes tuskaloosa and Agarodes logani are members of the sub- genus Agarodes Banks (Ross and Wallace 1974) based on the small antennal scape and slender mesal lobe of the maxillary palp. Both were collected with a black light. Type material is deposited at the National Museum of Natural History, Smithsonian Institution, Washington D.C. Terminology follows that of Schmid (1980) and Ross and Scott (1974). Agarodes logani Keth and Harris, new species (Figs. 1, 4) Diagnosis.—This species resembles A. stannardi but differs, primarily, in the elon- gate, strongly curved dorsomesal process of the mesal processes of the inferior append- age. The dorsomesal process (Fig. 1) of A. logani is about twice the length seen in A. stannardi (Fig. 2) and much more serrate ventrally. The female is very similar to A. VOLUME 101, NUMBER | Fig. 1. Agarodes logani, male genitalia. a) lateral, b) ventral, c) dorsal , d) phallus. 87 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 101, NUMBER 1 stannardi differing only in the pair of short processes extending from apex of the dor- sum of segment IX. These processes in A. stannardi are divergent and basally lobate in dorsal view (Fig. 3) and in A. logani nar- row and lacking basal lobes (Fig. 4). Male.—Length 9-11 mm. Body, legs, and head golden to dark brown. Antenna brown with 43 segments, scape small and rectangular. Labial palpus 0.9 mm long; maxillary palpus 0.5 mm in length, oblong, and bearing 2 small inner processes. Wings golden to dark brown; forewing with exten- sive peg-like setae in anal region and along M and Cu veins. Abdominal segment VIII annular. Segment IX inserted within seg- ments VII and VIII, reduced, and dorsally fused with base of segment X. Segment X elongate and narrow in dorsal view, divided apically; in lateral view sinuate dorsally and lightly sclerotized, tapering to a point and curving slightly dorsad at apex. Genitalia as in Fig. | with preanal appendage narrow and elongate extending nearly half length of tergum X, curving dorsad. Inferior ap- pendage uniform over entire length in lat- eral view; in ventral view united basally, slender anteriorly, widening midway, and narrowing posteriorly. Mesal processes bi- partite; dorsomesal process over *%4 length of inferior appendage, slender and narrow- ing to acute apex with large, serrate, ven- trad projections from % of length to apex and curving markedly ventrad in lateral view. Ventromesal process thin and nearly equal in length to dorsomesal process, di- vergent, curving markedly mesad at tip, slightly serrate distally at midlength and dorsally at apex. Phallus long and cylindri- cal, widening at membranous apex. Female.—Length, color, and general structure as in male. Genitalia as in Fig. 4 with cerci long and contiguous, mesally merging with a short lobe. Apex of tergum IX having a pair of short processes pro- jecting posteriorly from dorsum; processes narrow and fused basally, lacking basola- teral lobes. Genital chamber narrow with margins folded, sclerotized, and curving 89 mesad in ventral view, a pair of sclerous, scoop-like lobes diverging from base in dorsal view. Immatures.—Unknown Etymology.—Latin: of Logan, primary author’s son Holotype.—<¢, Florida, Gadsden County, headwaters of Quincy Creek, 7 km. north Quincy at Florida A&M Research and Ex- tension Center, N30°39'27", W84°36'50", 19 April 1994, Pescador and Rasmussen. Paratypes.—Florida, same as above, | 6; same, but N30°39'19", W84°36'51”, 6 Oc- tober 1993, Jones, Pecador, and Rasmussen, Nye Distribution.—A garodes logani is known only from the type locality. More speci- mens have been found in the same area in- cluding Agarodes larvae which, upon rear- ing, may be found to be A. logani. Agarodes tuskaloosa Keth and Harris, new species (Fig. 5) Diagnosis.—This species resembles A. stannardi and the preceding new species, but it differs from A. stannardi in the much thinner inner process of the mesal process of the inferior appendage and from A. /o- gani in the presence of a short basomesal process (Fig. 5). The dorsal and ventrome- sal processes of A. tuskaloosa are like those of A. stannardi but twice the length. Male.—Length 11-13 mm. Body, legs and head golden to reddish brown. Antenna dark brown with 43 segments, scape small and trapezoidal. Labial palpus 1.0 mm long; maxillary palpus 0.6 mm in length, oblong. Wings light brown. Abdominal segment VIII annular. Segment IX inserted within segments VII and VIII, reduced, and dor- sally fused with base of segment X. Seg- ment X elongate and narrow in dorsal view, split apically; narrowing to elongate, acute apex in lateral view. Genitalia as in Fig. 5 with preanal appendage narrow, elongate, 4 length of tergum X, uniform over entire length in lateral view. Inferior appendage narrow basally, broadening toward apex in 90 Fig. 3. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Agarodes stannardi, female genitalia. a) lateral, b) ventral, c) dorsal. VOLUME 101, NUMBER 1 9] Fig. 4. Agarodes logani, female genitalia. a) lateral, b) ventral, c) dorsal. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Agarodes tuskaloosa, male genitalia. a) lateral, b) ventral, c) dorsal, d) phallus. VOLUME 101, NUMBER 1 lateral view; in ventral view slender and united basally, broadening % of length, and tapering to acute apex. Mesal processes tri- partite; basomesal process short and acute with slightly serrate apex dorsally, dorso- mesal process %4 length of inferior append- age and strongly curved in lateral view, apex slightly serrate ventrally; inner process thin, uniform, and elongate. Phallus long and cylindrical, widening slightly at mem- branous apex. Female.—Unknown Immatures.—Unknown Etymology.—Native American spelling: of Tuscaloosa region Holotype.—d6, Alabama, Tuscaloosa County, Big Sandy Creek, 7.2 km. south of Coaling, on unmarked county road, 15 May 1991, Harris. Paratype.—Alabama, locality and date as holotype, 1 dé. Distribution.—A garodes tuskaloosa is known only from the type locality. Exten- sive collecting at the type locality and else- where in Big Sandy Creek has yielded no additional specimens. 93 ACKNOWLEDGMENTS The Department of Biology of Clarion University of Pennsylvania provided equip- ment and facilities for this study and is thankfully recognized. Specimens for com- parison were provided by Dr. Brian Armi- tage at the Ohio Biological Survey and by Dr. John Morse at Clemson University. Thanks also to Jerome Jones, Manuel Pes- cador, and Andrew Rasmussen, of Florida A & M University, for specimen collections in northern Florida. LITERATURE CITED Harris, S. C. 1987. A new species of Agarodes (Tri- choptera: Sericostomatidae) from southeastern United States. Proceedings of the Entomological Society of Washington 89(1): 74-76. Ross, H. H. and D. C. Scott. 1974. A review of the caddisfly genus Agarodes with descriptions of new species (Trichoptera: Sericostomatidae). Jour- nal of the Georgia Entomological Society 9: 147— 155: Ross, H. H. and J. B. Wallace. 1974. The North Amer- ican genera of the family Sericostomatidae (Tri- choptera). Journal of the Georgia Entomological Society 9: 42-48. Schmid, F 1980. Genera des Trichoptéres du Canada et des états adjacents, pt. 7: 296 pp. Jn les insectes et arachnides du Canada. Agriculture Canada, Ot- tawa. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 94-98 THE HIGHER CLASSIFICATION OF THE 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.—Characters are taken from the literature and their derived states determined. The following higher classification of the Alydidae best agrees with the data: family Alydidae; subfamilies Alydinae and Micrelytrinae, the Alydinae with tribes Alydini and Daclerini; the Micrelytrinae with tribes Micrelytrini and Leptocorisini; the latter with subtribes Leptocorisidi and Noliphidi. Key Words: clerini Until 1965 the family Alydidae had been variously treated as a subfamily of Corei- dae, or as a family in its own right. How- ever, when treated as a family the habit of subfamilial treatment persisted, and the lower categories were considered tribes, not subfamilies (see Schaefer [1965] for the taxonomic history). In 1965 I presented ev- idence for family rank of the group, and recognized two subfamilies, Alydinae and Micrelytrinae, the latter with two tribes, Mi- crelytrini and Leptocorisini. In the same year, Ahmad (1965) pub- lished his revision of Leptocorisa and its relatives. He treated this group as a subfam- ily, of status equal to Alydinae (sensu mihi) and Micrelytrinae (Micrelytrini mei). He discussed these three subfamilies very brief- ly, but did not give arguments for treating them as of equal rank. Ahmad and I had not seen each other’s papers, and therefore neither of us could consider the higher-rank treatment of the other. Since 1965, Ahmad’s view has prevailed, with a lapse in 1979, when Ahmad et al. (1979) treated Leptocorisini and Micrelytri- ni as tribes in the subfamily Leptocorisinae. Alydidae, higher classification, Leptocorisini, Micrelytrinae, Noliphini, Da- Nevertheless, most post-1965 authors treat- ed the three groups as subfamilies, perhaps because Ahmad’s (1965) study of the Lep- tocorisinae (sensu suo) was more detailed than mine of the entire Alydidae. Indeed, I myself subsequently treated the three groups as subfamilies (see, for example, Schaefer 1972, 1980, Schaefer et al. 1989, but see Schaefer 1996); most notably, the three subfamilies are recognized in the cat- alog of Nearctic Heteroptera (Froeschner 1988). (Note: the change by Henry and Froeschner [1992] from Leptocorisinae to Leptocorinae is surely incorrect: the generic name is Leptocorisa, not Leptocoris [which is a genus in Rhopalidae].) In 1993, Li and Zheng published a study of alydid phylogeny. In it they concluded “that Schaefer’s (1965) division of this family into two subfamilies—Alydinae and Micrelytrinae—is reasonable.”’ Their ‘‘cladograms do not support Ahmad’s (1965) and some other authors’ contention that the Alydidae be divided into three sub- families—Alydinae, Leptocorisinae, and Micrelytrinae’” (quoted from the English abstract of Li and Zheng [1993]). Since the VOLUME 101, NUMBER 1 appearance of this paper, Prof. Zheng has most generously provided me with an En- glish translation of it. Li and Zheng (1993) discuss and illus- trate many characters in some detail, thus adding substantially to the already rich lit- erature on Alydidae (see references in Table 1). Accordingly, I have extracted characters from this literature and have attempted to determine their derived states, in an attempt to establish better the Alydidae’s higher classification. METHODS I extracted from the literature on the Al- ydidae characters whose states could be tabulated and whose polarities for the most part could be determined. All these char- acters are ones important in the higher sys- tematics of Coreoidea and, indeed, of Pen- tatomomorpha (the heteropteran infraorder to which Coreoidea belongs; see Henry 1997). Sample size is of course a problem; all genera and many species of Alydinae are described in Schaffner’s dissertation (1964), but only for the Leptocorisini are all mem- bers of the group treated and their mor- phology described, in Ahmad’s (1965) monograph. In particular, the Micrelytrini need revision, with attention given to mor- phological features of systematic impor- tance in other alydids. In polarizing the character states I take the Coreidae as the outgroup. Henry (1997) presents convincing evidence that this fam- ily is the sibling group of Alydidae. Other arguments for my polarizing occur in the references and in footnotes to Table 1. RESULTS AND DISCUSSION Twenty-eight characters and their states are in Table | (where I anticipate my con- clusions by treating Micrelytrini and Lep- tocorisini as tribes and Alydinae as a sub- family). Of these characters, I polarized 23; three of the remainder I could not polarize (Distribution, Host plants, Rostral seg- ments); one (Scent gland auricle) is ambig- uous; and it is not clear to me if bifid (Mi- 95 crelytrini) and trifid (Leptocorisini) medial projections of the genital capsule are sepa- rate advances over the (primitive) alydine condition, or whether the trifid condition is a further advance over the bifid (in which case, this character would group Micrely- trini and Leptocorisini together). Thirteen of the characters are autapo- morphies of one of the three groups. These autapomorphies are not the only ones defin- ing these groups, of course, because I was not seeking autapomorphies in the litera- ture. Nevertheless, the fact that Alydinae has eight autapomorphies, and the other two groups have fewer (Micrelytrini: 4; Lepto- corisini: 1), supports the subfamily status of Alydinae. Within the Alydinae, Ahmad ei al. (1979) created a tribe, Daclerini, for the published genus Daclera and for another, unpublished genus. Although they present the new tribe as “MS,” it is briefly described in their key and therefore appears to be valid. I have not seen specimens either of Daclera or of the undescribed genus. However, Li and Zheng (1993) write that Daclera has many apo- morphies not shared with other Alydinae. Therefore I treat Daclerini as a tribe in Al- ydinae, pending further study. Within the subfamily Micrelytrinae, Mi- crelytrini and Leptocorisini share six apo- morphies (seven, if the median projection condition is synapomorphic; see above), more than either group shares with Alydi- nae (Micrelytrini and Alydinae: 2; Lepto- corisini and Alydinae: 2). Three of these four synapomorphies shared by Alydinae and either Micrelytrini or Leptocorisini are head characters. The states of these char- acters vary considerably in the Coreoidea (Schaefer 1965), and their common posses- sion in Alydinae and one of the other tribes may therefore be homoplasious. If one ac- cepts that eight autapomorphies is a reason- able argument for subfamily status for Al- ydinae, then six synapomorphies should support subfamily status for Micrelytrini plus Leptocorisini. Moreover, four of these six synapomorphies are characters of the 96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Differences and similarities among Micrelytrini, Leptocorisini, and Alydinae. Apomorphic states in boldface. Micrelytrini Leptocorisini Alydinae Reference Distribution tropical, subtropical tropical, subtropical tropical, subtropical; Schaffner 1964, Ah- Host plants Head Midcephalic sulcus Head constricted basally Head with ‘‘col- lar Ocelli on tubercle Paraclypei well de- veloped Rostral segments Thorax Hind femur Hind tibia Forewing media Forewing costa Scent gland peri- treme: lateral and anterior auricles fused Abdomen Trichobothria (Sth sternum) Male genital capsule Ventral rim External opening Dorsal wall Median projection Cuplike sclerite with lateral pro- jections Segment 10 Paramere Graminae (7?) deep yes no yes sometimes 3 44 = twice 3 not armed straight, untoothed coriaceous basally in membrane not fused to radius and media variable (no: Schae- fen d965=tyes3 1 and Zheng 1993) in a triangle with spine posterior sclerotized bifid no sclerotized dorsally apex not tuberculate Graminae deep rarely yes rarely not armed straight, untoothed coriaceous basally in membrane not fused to radius and media no in a triangle without spine® dorsal sclerotized trifid no sclerotized dorsally apex not tuberculate some temperate Leguminosae shallow or absent Pe BN ices tal ali ee) rarely twice 3 spined or with stiff setae* usually curved and with ventral tooth? not coriaceous fused? yes in a line without spine dorsal or postero- dorsal membranous single yes membranous dorsal- ly apex tuberculate mad 1965 Schaefer 1979 Li and Zheng 1993 Li and Zheng 1993 Schaefer 1965 Li and Zheng 1993 Li and Zheng 1993, Ahmad 1965 Ahmad 1965, Schae- fer, unpubl. Schaffner 1964, Ah- mad 1965 Schaffner 1964, Li and Zheng 1993 Li and Zheng 1993 Schaefer 1965, Li and Zheng 1993 Schaefer 1965, Li and Zheng 1993 Schaefer 1965, 1975 Schaefer 1980 Schaefer 1980 Schaefer 1980, Schaefer et al. 1989 Schaefer 1980, Schaefer et al. 1989 Schaefer et al. 1989 Schaefer 1980 Ahmad 1965, Li and Zheng 1993 VOLUME 101, NUMBER 1 Table 1. Continued. Micrelytrini Leptocorisini Alydinae Reference Male aedeagus Vesica not slender, coiled' Conjunctiva laterally with pair of asymmetrical appendages dorsally with pair of apically di- rected append- ages Phallosoma ages Female genitalia 9th paratergite divided 2nd valvula Ring sclerites Ring-sclerite sacs 2 pairs 1 pair® | pair 1 median sac not slender, coiled! laterally with pair of asymmetrical appendages dorsally with pair of apically di- rected append- not divided partly membranous sclerotized Schaefer 1965, Ah- mad 1965, Li and Zheng 1993 Schaefer 1965, Ah- mad 1965, Li and Zheng 1993 Schaefer 1965, Ah- slender, straight! without these ap- pendages without these ap- pendages mad 1965, Li and Zheng 1993 not divided Schaefer 1965 sclerotized Schaefer 1965 | pair Schaefer 1965 | pair Schaefer 1965 «Curved and armed femora and tibia are uncommon in Coreoidea (except some Meropachydinae and some male Coreinae). > Fusion of wing veins appears ipso facto to be more advanced than nonfusion. © The posterior abdominal trichobothria in Coreoidea are usually in a triangle (except Rhopalidae) (Schaefer 1975). 4 These features vary independently of one another (Schaefer 1980), and therefore may be treated as sepa- rate characters. © Spine is present in at least one leptocorisine (discussion in Schaefer 1980, p. 126). ' According to Li and Zheng (1993), the apomorphic state of Alydinae differs from the apomorphic state of Micrelytrinae. * Ahmad (1965) writes that Leptocorisini have O-4 pairs of ‘‘intervalvular sclerites’’; I believe these are not the same as the sacs associated with the ring sclerites. male genitalia, a character complex always useful in heteropteran higher classification; common possession here is unlikely to be homoplasious In addition, other characters support the uniting of these two groups as a subfamily; these characters and the argu- ments based upon them, are not easily tab- ulated: see Schaefer (1965). Li and Zheng (1993) comment upon the antlike fascies of Alydinae, treating it as an autapomorphy of the group. It is true that immature alydines are antlike, but so are the adults of several Micrelytrini. In fact, there are two groups of Micrelytrini, one of somewhat or quite elongate insects, and the other of smaller often antlike insects (Schaefer 1996). Members of both groups occur in both the New and Old World trop- ics. By chance, Li and Zheng (1993) took as their representatives of Micrelytrini Mar- cius and Paramarcius, both members of the somewhat elongate (and nonantlike) group. Of great interest would be a study of the phylogenetic relationships among these four groups (New World and Old World antlike and elongate micrelytrines), be- tween the antlike micrelytrines and the Al- ydinae (whose nymphs are antlike), and be- tween the elongate micrelytrines and the Leptocorisini (most of whose genera are Old World tropical; the Noliphidi are some- what elongate and the Leptocorisidi are very elongate). I discussed some of these relationships earlier (Schaefer 1972). CLASSIFICATION As a result of this work, I suggest the following classification, in which the tribes of Ahmad’s (1965) Leptocorisinae are re- duced to subtribes. 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Family Alydidae Amyot et Serville 1843 Subfamily Alydinae Amyot et Serville 1843 Tribe Alydini Tribe Daclerini Subfamily Micrelytrinae Stal 1867 Tribe Micrelytrini Stal 1867 Tribe Leptocorisini Stal 1870 Subtribe Leptocorisidi Stal 1870 Subtribe Noliphidi Ahmad 1965 Note: Based on their cladistic analysis, Li and Zheng (1993) suggest that Acestra, a genus placed uneasily in the Micrelytrini (see discussion in Li and Zheng [1993]), be removed from Micrelytrini and raised to tribal rank in the Micrelytrinae; they do not do this formally. Also, as I mentioned above, Li and Zheng (1993) found autapo- morphies in Daclera, the only described ge- nus now in Daclerini. These two genera should be studied more closely, as should the Micrelytrini as a whole. ACKNOWLEDGMENTS I am deeply grateful to Zheng Le-yi for translating the Li and Zheng (1993) paper for me. Without his kindness and generos- ity, my paper had been difficult to under- take and impossible to complete satisfac- torily. I thank also Ann Harlan for word- processing this paper and for being a fa- vorite daughter. REFERENCES CITED Ahmad, I. 1965. The Leptocorisinae (Heteroptera: Al- ydidae) of the world. Bulletin of the British Mu- seum (Natural History) Entomology Supplement 5: 1-156. Ahmad, I., N. Abbas, M. U. Shadab, and A. A. Khan. 1979. Generic and supergeneric keys with refer- ence to a checklist of alydid and stenocephalid fauna of Pakistan, Azad Kashmir and Bangladesh (Heteroptera: Coreoidea) with notes on their dis- tribution and food plants. Supplement of the En- tomological Society of Karachi, Pakistan 4(2): 1— 19. Amyot, C. J. B. and A. Serville. 1843. Historie natu- relle des Insectes—Hémipteres. Librairies Ency- clopédique de Roret, Paris. Froeschner, R. C. 1988. Family Alydidae Amyot and Serville, 1843. The broad-headed bugs, pp. 4-11. In Henry, T.J. and R. C. Froeschner, eds., Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E.J. Brill, Leiden, The Netherlands. Henry, T. J. 1997. Phylogenetic analysis of family groups within the infraorder Pentatomomorpha (Hemiptera: Heteroptera), with emphasis on the Lygaeoidea. Annals of the Entomological Society of America 90: 275-301. Henry, T. J. and R. C. Froeschner. 1992. Corrections and additions to the “‘Catalog of the Heteroptera, or true bugs, of Canada and the continental United States.”’ Proceedings of the Entomological Society of Washington 94: 263-272. Li, Xin-Zheng and Zheng Le-yi. 1993. Preliminary study on the phylogeny of Alydidae. Acta Zoo- taxonomia Sinica 18: 330-343 (Chinese, English summary.) [Note: I have used an English trans- lation prepared by Zheng Le-yi.] Schaefer, C. W. 1965. The morphology and higher classification of the Coreoidea (Hemiptera-Heter- optera). Part III. The families Rhopalidae, Alydi- dae, and Coreidae. Miscellaneous Publications of the Entomological Society of America 5(1): 1—76. 1972. Clades and grades in the Alydidae. Journal of the Kansas Entomological Society 45: 135-141. . 1975. Heteropteran trichobothria (Hemiptera: Heteroptera). International Journal of Insect Mor- phology and Embryology 4: 193-264. . 1979. The host plants of the Alydinae, with a note on heteroptypic feeding aggregations (He- miptera: Coreoidea: Alydidae). Journal of the Kansas Entomological Society 53: 115-122. . 1980. The genital capsule of the Alydidae (Hemiptera: Heteroptera: Coreoidea). Internation- al Journal of Insect Morphology and Embryology 9: 107-128. . 1996. A new species of Cydamus, with a key to the species of the genus (Hemiptera: Alydidae). Annals of the Entomological Society of America 89: 37-40. Schaefer, C. W., J. C. Schaffner, and I. Ahmad. 1989. The Alydus-group, with notes on the alydine gen- ital capsule (Hemiptera: Heteroptera: Alydidae). Annals of the Entomological Society of America 82: 500-507. Schaffner, J. C. 1964. A taxonomic revision of certain genera of the tribe Alydini (Heteroptera: Corei- dae), Ph.D. Dissertation, Iowa State University, Ames, Iowa. Stal, C. 1867. Bidrag till hemipterernas systematik. Ofversigt af Kongliga Svenska Vetenskaps-Aka- demiens Forhandlingar 24 (7): 491—560. 1870. Enumaeratio hemipterorum. Pt. 1. Svenska Vetenskaps-Akademiens Handlingar 9(1): 1-232. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 99-105 OBSERVATIONS ON TWELVE FAMILIES OF HOMOPTERA IN MACAU, SOUTHEASTERN CHINA, FROM 1989 TO THE PRESENT EMMETT R. EASTON AND WING-WAH PUN (ERE) University of Macau, P.O. Box 3001, Macau; (WWP) Departamento de Servi¢os Agrarios de Camara Municipal das Ilhas, Coloane, Macau Abstract.—The insect species found in present day Macao are largely those feeding upon small numbers of remnant agricultural plantings or on those plants representing the subtropical evergreen broad leaf forest or remnants of the tropical monsoon rainforest as agricultural crops are only grown on a small scale. Records of 37 species of Homoptera of the Aleyrodidae, Aphididae, Cicadellidae, Cicadidae, Coccidae, Delphacidae, Diaspi- didae, Flatidae, Fulgoridae, Margarodidae, Pseudococcidae and Psyllidae are listed for the Portuguese territory of Macao of which 27 are new records. Key Words: In the early part of the present century a number of homopteran species were listed as occuring in Macao and Hong Kong as well as part of the Guangdong Province of mainland China all of which was known at that time as South China (Kirkaldy 1909). Rice was considered the main agricultural crop and farming was the chief livelihood of most of the people in south east China. The land area that John C.W. Kershaw (Kir- kaldy 1909) collected insects on was con- sidered an island at that time but it is now connected to the mainland of China and to- day is considered the peninsula. The ma- jority of the collections made in this study, however were taken from the islands of Tai- pa and Coloane which are connected to the peninsula either by two bridges (Taipa) or a causeway (Coloane) which did not exist in the early part of the century. Many of the insect species noted to occur in Macao at the beginning of the century may now be non-existant. We were unable to find evi- dence, for example, of the planthoppers de- scribed by Frederick Muir (1913) because of modernization and construction of hotels Macao, Homoptera, faunal list and apartment complexes that have re- placed agricultural farmlands. Easton and Pun (1997) discussed the species of true bugs in the region and here we discuss the Homoptera. MATERIALS AND METHODS Insects were sampled routinely from the walls of window-lit buildings (Taipa island) that are often illuminated at night. Other in- sect groups less sensitive to light, such as the cicadas other than those in the genera Cryptotympana, and Huechys, as well as the Flatidae, Fulgoridae and the scale in- sects, were sampled with an insect net from trees, shrubs and grasses in forested areas on the islands of both Coloane and Taipa as well as from the peninsular area connected to the Guangdong region of China. The names of the plant hosts follow the Macau Catalogue of plants and their addenda (1991). Voucher specimens of the Homop- tera named here are housed in the Ento- mology Museum of the Agrarian Services on Coloane (Seac Pai van Park) under the curatorship of the second author. 100 List OF SPECIES HOMOPTERA Aleyrodidae Aleurocanthus spiniferus (Quaintance), orange spiny whitefly. No date, collector unknown, NEW RECORD, Coloane Island, Macao, ex. Ficus rumphi, Rosa chinensis. Mound and Halsey (1978) list it from India, Sri Lanka, Taiwan, Thailand, Malaysia, Philippines, Sumatra, Japan, and Hong Kong, while Peng and Liu (1992) report it from the Fujian, Guangxi, Henan, Hunan, Jiangsi, and Zhejiang provinces of China. Trialeurodes vaporariorum (Westwood), greenhouse whitefly. No date, collector un- known, NEW RECORD. Coloane Island, Macao, ex. Brassica oberacea, Citrus med- ica and Hybiscus rosa-sinensis. Mound and Halsey (1978) list it from Malaya, India, Sri Lanka, Hawaii, New Guinea, New Zealand and most of the provinces of mainland Chi- na. Aphididae Aphis gossypii Glover, cotton or melon aphid. 5 Dec 1992, PWW leg. NEW RE- CORD, Coloane Island, ex. Al/thaea rosea, Capsicum frutescens, Chrysanthemum mor- ifolium, Citrullus lanatus, Colocassia es- culenta, Cucurbita moschata, Dianthus car- yophyllus, Hibiscus rosa-sinensis, Lilium Japonicum, Litchi chinensis, Phaseolus ra- diatus, Psidium guajava, Punica granatum and egg plant, Solanum melongena. It has been reported in Hong Kong (Lee and Win- ney 1981) and is widely distributed in Chi- na (Peng and Liu 1992). Aphis nerii Boyer de Fonscolombe, ner- ium or Oleander aphid. 5 Dec 1992, PWW leg, NEW RECORD, Coloane Island, Ma- cau, ex. Asclepius curassavica, Nerium in- dicum. Raychaudhuri (1980) lists it throughout India and Japan, Java, Korea, and Taiwan. In Hong Kong, Lee and Win- ney (1981) report it common during winter months when it feeds on the shoots and leaves of oleander. Peng and Liu (1992) list PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON it from Guangdong, Guangxi, Hunan, and Jiangsu provinces of China. Formosaphis micheliae Takahashi. 21 March 1995, PWW leg. NEW RECORD, Coloane Island, Macao, ex. white jade or- chid tree, Michelia alba and M. champaca. Blackman and Eastrop (1994) record it from Japan and Taiwan and Lee and Win- ney (1981) found it in Hong Kong. Myzus persicae (Sulzer), green peach aphid. No date or collector, NEW RE- CORD, Coloane Island, Macao, ex. Bras- sica chinensis, B. oberacea and Prunus per- sica. This is considered a cosmopolitan temperate species and it is more common during winter months in Hong Kong with populations declining after April (Hill et al. 1982). It is widely distributed in China. Neophyllaphis podocarpi Takahashi, buddhist pine aphid. 9 Feb 1995, PWW leg. NEW RECORD, Coloane Island, Macao, ex. Podocarpus macrophyllus. Distribution includes Malaysia, Taiwan, Hong Kong, Ja- pan and the Guanxi, Hunan, Jilin, and Zhe- jiang provinces of China (Peng and Liu 1992): Shivaphis celti B. Das. 17 March 1994, PWW leg. NEW RECORD, Coloane Is- land, Macao, ex. Celtis sinensis. Distribu- tion includes Korea, Japan, India, Taiwan, Hong Kong and the Fujian, Guangdong, Guangxi, Guizhou, Hebei, Hunan, Jiangsu, Liaoning, Shandong, Sichuan, and Yunnan provinces of China (Peng and Liu 1992). Tinocallis kahawaluokalani (Kirkaldy), crepe myrtle aphid. 27 Oct 1993, PWW leg. NEW RECORD, Coloane Island, Macao. ex. Lagerstroemia indica. This species is also found in India, Taiwan, Japan, and Ha- wali (Raychaudhuri 1980). Cicadellidae Lodiana brevis (Walker), yellow-banded leafhopper. 9 Dec 1993. Ng Wai Man leg. NEW RECORD, Coloane Island, Macao, ex. Cinnamomum camphora, Ficus micro- carpa and Euphoria longan. In Hong Kong Lee and Winney (1981) reported it from Citrus sp. Distribution includes the Guang- VOLUME 101, NUMBER 1 dong, Guizhou, and Yunnan provinces of China as well as India, Taiwan, Malaysia, Thailand, and Japan (Datta 1988). Nephotettix virescens (Distant), green rice leafhopper. No date or collector, NEW RECORD, Coloane Island, Macao, ex. Cit- rus sp., Bambusa sp., and Saccharum offi- cinarum. There are records of it from rice in the Tai Lung farm insect collection in Hong Kong and Hill (1975) lists it as a pest of rice in India and China. Petalocephala chlorocephala Walker. 4 Sept 1993, Ng Wai Man leg, NEW RE- CORD, Coloane Island, Macao, no host data. Tettigoniella spectra (Distant), rice white leafhopper. 15 Sept 1992, Ng Wai Man leg, NEW RECORD, Coloane Island, Macao, ex. Morus alba, Oryza sativa, Saccharum officinarium. Distribution includes Taiwan, India, and Japan as well as the Guangdong and Hunan provinces of China (Peng and Liu 1992). Another species of leafhopper, Dryado- morpha pallida Kirkaldy has not been col- lected in our study but Webb (1981) re- ported it from Macao and Hong Kong as well as the neighboring areas of India, Bangladesh, Taiwan, Japan, Laos, and the Philippines. Cicadidae Cryptotympana atrata (FE), large brown cicada. 20 June 1994. Cheong Chi Kong leg, Coloane island, Macao, ex. Melia aze- darach, Morus alba, Tectonis grandis. Emergence of adults occurs from late May through early July in general but emergence on Taipa Island in 1998 was earlier (May 15). Distribution includes Hong Kong, the provinces of Guangdong, Hebei, and Zhe- jiang of China (Wu 1935) as well as Ma- laysia, Japan, and Taiwan. Cryptotympana mandarina Distant. May 1997, 15 May 1998, ERE leg, Taipa Island, Macao. Kershaw (1903) is believed to be the first to record it from Macao while Kir- kaldy (1909) reported it from Hong Kong. Nymphal skins have been observed on the 101 trunks of various trees such as Acacia con- fusa and Casuarina equisetifolia in the mu- nicipal cemetary on Taipa island (Easton 1992) as well as the foxglove tree, Paulow- nia fortunei on Guia Hill of Macao penin- sula in May, 1997 and on Hibiscus tiliaceus near the Monte Forte, Fortress, May 10, 1998. Emergence holes near the base of the trees indicated where the nymphs left the soil. Watery fluid has been observed emit- ting from the anal region of resting adults on Taipa Island suggesting that they had re- cently emerged from the ground. As feed- ing in adults has not been documented, the watery fluids may have accumulated while the nymphs were underground and release may be necessary before sound production can take place. Distribution also includes the Guangdong and Hainan provinces of China and Taiwan. Chremistica ochracea (Walker). 23 May 1995, 10 May 1998, ERE leg, Taipa Island, Macao. Hayashi (1977) recorded this spe- cies from Macao, and Hill et al. (1982) il- lustrated it as the green clearwing cicada under the genus Dundubia in Hong Kong. Distribution also includes Taiwan and the Guangdong region of China. Gaeana maculata Walker, yellow-spotted black cicada. 14 April 1994, Ng Wai Man leg, Coloane Island; 7 May 1998, ERE leg, Guia Hill, Macao peninsula. In the Seac Pai Van agricultural park on Coloane Island, nymphs were observed emerging as early as March 25 (Easton 1992). It has not been observed on Taipa island suggesting certain food plants may be necessary for nymphal development. In Hong Kong, Cheung and Marshall (1973) reported it from Schinus terebinthiofolius, Christmas berry tree, and Paulownia tormetosa and in Macao the se- nior author has observed it among the fo- liage of Pawlownia fortunei on the Guia Hill. Wu (1935) recorded it from Myanmar (Burma), Assam area of India, Vietnam, and Guangxi Province of China. Huechys sanguineus (DeGeer), red-nosed cicada. 31 August 1992, Ng Wai Man leg, Coloane Island, Macao. Kirkaldy (1909) 102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON first reported this species from Macao and Hong Kong. From 1989-1997 emergence from the ground has always been later in the year than with the former species and from September through December (Easton 1991). In 1998 however an emergence of 15—20 individuals was observed in an ar- boretum on Coloane island on May 10. Specific food plants have not been associ- ated with nymphal feeding. Mogannia hebes (Walker), grass cicada. 9 June 1994, ERE leg, NEW RECORD, Taipa Island on shrub on university campus; June 1996, May 1998, ERE leg, Seac Pai Van Agricultural Park, Coloane Island. Kir- kaldy (1909) was probably the first to re- cord it from Hong Kong, and Hayashi (1976) described its feeding on the stems of Miscanthus grass where females oviposit into stems and leaf midribs from April to June. Distribution includes Korea, India, Myanmar, and the Chejiang, Fujian, Guang- dong, Guangxi, Jiangsi, Hunan, Sichuan, and Yunnan provinces of China. Mogannia nasalis (White). 25 May 1966, ERE leg, Taipa Island, Macao on tree fo- liage along hiking trail in thickly forested hillside. This species was first recorded in Macao and Hong Kong by Kirkaldy (1909). It is also found in the Assam area of India. Platypleura hilpa Walker, spotted brown cicada. 2 August 1992, PWW leg, Coloane Island, Macao. Nymphs were observed emerging at night near the trunks of Casu- arina equisetifolia trees near the municipal cemetary on Taipa Island (Easton 1992). Kirkaldy (1909) reported it from Hong Kong and Macao, and it is also found in the Guangdong and Hunan provinces of China (Peng and Liu 1992). Coccidae Ceroplastes ceriferus (Fabricius), indian wax scale. 18 Dec 1992. PWW leg, NEW RECORD, Coloane Island, Macao, ex. Cin- namomum camphora, Melastoma sanguti- neum, Michelia figo, Morus alba. Tang (1991) listed it from Australia and Hawaii and it is also found in Japan, India, Sri Lan- ka and provinces of China south of the Yangtze River. Saissetia coffeae (Walker), helmet or hemispherical scale. 19 Jan 1988, PWW leg, NEW RECORD, Coloane Island, Ma- cao, ex. Fukien tea, Carmona microphylla and sago palm, Cycas revoluta. Hill (1975) listed citrus, guava and mango as alternate hosts but mainly it is found on coffee plants and widespread in the tropics including southeastern Asia. Delphacidae Nilaparvata lugens (Stal), brown plant- hopper of tice. 22. Oct, 1997-7 JERE wes. NEW RECORD, Taipa Island on window ledges and outside on floors of university campus buildings illuminated by lights. Ac- cording to Hill (1975) and Wada et al. (1987), N. lugens invades Japan with the monsoon winds every year from China, so the large numbers (1,000+) observed in Macao over a brief period of 1—2 days sug- gests that the insects migrated from the neighboring Guangdong Province since rice is not grown commercially locally in Ma- cao as a crop. In Malaysia and Indonesia, it is a major pest of rice and produces a browning effect on the plants known as ‘““*hopperburn.”’ Lee and Winney (1981) also report it from Gladiolus gandavensis in Hong Kong. Yang (1989) gives its distri- bution as Australia, New Guinea, India, Ko- rea, Taiwan, Philippines, Vietnam, and the Pacific islands of Fiji, Guam, Yap, and Pa- lau, while Kuoh et al. (1983) list it from Anhui, Fujian, Gansu, Guangdong, Gu- angxi, Guizhou, Hebei, Henan, Honan, Hu- nan, Jiangsu, Jiangxi, Jilin, Liaoning, Shan- dong, Shanxi, Yunnan, and Zhejiang Prov- inces in China. Other species of Delphacidae reported earlier by Muir (1913), such as Belocera sinensis Muir, Phyllodinus macaoensis Muir, and Tropidocephala saccharivorella Matsumura, were not collected in this study and may no longer be found locally due to VOLUME 101, NUMBER 1 the urbanization which has replaced agri- cultural crops. Diaspididae Aulacaspis rosarum Borchsinius, Asiatic rose scale. No date or collector, NEW RE- CORD, Coloane Island, Macao, ex. Rosa chinensis. Distribution includes the Fujian, Guangdong, Guangxi, Hunan, Jiangsu, Ji- angxi, Shandong, Sichuan, Yunnan, and Zhejiang provinces of China (Peng and Liu 1:99): Aulacaspis yabunikkei Kuwana. 21 April 1988, PWW leg, NEW RECORD, Coloane Island, Macao, ex. Cinnamomum campho- ra. In Hong Kong, Lee and Winney (1981) also record it from pond spice, Litsea glu- tinosa. Distribution includes Japan and the Guangdong, Guizhou, Hunan, Sichuan, Yunnan, and Zhejiang Provinces of China (Peng and Liu 1992). Hemiberlesia pitysophila Takagi. 7 June 1988, PWW leg, NEW RECORD, Coloane Island, Macao, ex. Pinus mansoniana. Lepidosaphes laterochitinosa Green. 20 May 1997, PWW leg, NEW RECORD, Co- loane Island, Macao, ex. Osmanthus fra- grans, Kwai-Fah. In Hong Kong there are records of it from this host as well as Schef- flera octophylla, \vy tree, in the Tai Lung Farm insect collection. Parlatoria pergandii Comstock, chaff Scales 23."reb) 1993: Yau HC. leg, NEW RECORD, Coloane Island, Macao, ex. Jas- minium sambac and Citrus sp. Distribution in the world is widespread, but in Asia it has been recorded from Australia, New Zealand, Japan, India, Philippines, Taiwan, and from the Anhui, Fujian, Guangdong, Guangxi, Hainan, Hebei, Henan, Hubei, Hunan, Jiangsi, Jiangsu, Liaoning, Qinghai, Shanxi, Shaanxi, Shanghai, Sichuan, Yun- nan, and Zhejiang Provinces of China (Peng and Liu 1992). Pseudaulacaspis cockerelli (Cooley), oyster or oleander scale. 7 May 1988, PWW leg, NEW RECORD, Coloane Is- land, Macao, ex. Michelia alba, M. figo and coconut palm, Cocos nucifera. In Hong 103 Kong, there are also records from bamboo palm, Chrysalidocarpus lutescens (Lee and Winney 1981). Its distribution includes Thailand, Taiwan, and the Guangdong, Hu- bei, Hunan, Jiangsu, Jiangxi, Shandong, Sichuan, Yunnan, and Zhejiang provinces of China (Peng and Liu 1992). Flatidae Lawana imitata (Melichar), white moth bug. 18 July 1991, PWW leg, Coloane Is- land, Macao, ex. Bauhinia sp., Euphoria longan, Hibiscus tiliaceus, Jasminium mes- nyi, Litsea monopetala, Murraya panicula- ta, and Pittosporum tobira. This insect is very common in lychee fruit tree orchards near Shenzhen in the Guangdong region of China. Easton (1992) reported it from the Seac Pai Van Agricultural Park in Macau. Its distribution also includes the Hainan, Hunan, Guizhou, Guangxi, and Yunnan provinces of China, Japan (Peng and Liu 1992) as well as Hong Kong (Lee and Win- ney 198): Seliza lignaria (Walker). 9 May 1997, ERE leg, NEW RECORD, Taipa Island, Macao, ex. Miscanthus grass along hiking trail in recently burned vegetation. Fennah (1956) listed it from Hong Kong and the Guangdong province of China, and it is also believed to occur in the Anhui, Fujian, Guizhou, Hunan, Sichuan, Yunnan, and Zhejiang provinces of China and India (Peng and Liu 1992). Medler (1992) reported Salurnis margi- nella (Guérin-Meneville) and Geisha dis- tinctissima (Walker) as occuring in Macao in the early part of this century, but we have been unable to document their presence in this study. Fulgoridae Fulgora candelaria (L.), lantern fly or lantern bug. 2 May 1994, Cheong Pak Fai leg, Coloane Island; 10 Dec 1990, ERE leg, Guia Hill near lighthouse, Macao Peninsu- la, ex. Litchi chinensis, Euphoria longan, Morus alba, Sapium sebiferum. Kershaw 104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and Kirkaldy (1910) were probably the first to describe, illustrate, and discuss the life history stages in the area known as South China which includes Hong Kong and Ma- cao. Even though four host plants are given above, Kershaw and Kirkaldy (1910) felt that E. Jongan and mango trees were the most important for the reproduction of this species. Its distribution also includes India (Assam), Cambodia and the Hainan, Hu- nan, Guangdong, Guangxi, and Sichuan provinces of China. Zanna chinensis (Distant). 31 Aug 1990, ERE leg, Taipa Island, Macao, ex. Casua- rina equisetifolia, horsetail tree. There are also records of this species in Hong Kong from the Tai Lung Farm insect collection. Wu (1935) recorded it from the Naga Hills area between India and Myanmar (Burma). Margarodidae Icerya purchasi Maskell, cottony cushion scale. 10 Feb 1994, PWW leg, Coloane Is- land; 12 May 1997, ERE leg, Taipa Island, NEW RECORD, ex. Cassia surattensis, the sunshine tree, Acacia confusa, Acalypha wilkesiana, Casuarina equisetifolia, Hyper- icum chinense, Chinese St. Johnswort, and Pentas lanceolata. In Hong Kong, Lee and Winney (1981) reported this species from Rosa spp. and Citrus limonia, Chinese lem- on. It has a widespread distribution includ- ing Japan, Korea, Philippines, Indonesia, Malaysia, and Sri Lanka in southeastern Asia, including the Gansu, Guizhou, Hei- longjiang, Hunan, Ningxia, Qinghai, and Xinjiang provinces of China (Peng and Liu 1992). Pseudococcidae Maconellicoccus hirsutus (Green), Asian hibiscus mealybug. 16 June 1988, PWW leg, NEW RECORD, Coloane Island, Ma- cao, ex. Hibiscus rosa-sinensis. In Hong Kong, it has been reported from Cuban bast, H. tiliaceus. Williams (1996) listed it from Bangladesh, Brunnei, Myanmar (Bur- ma), Cambodia, India, Indonesia, Laos, Malaysia, Nepal, Pakistan, Philippines, Sri Lanka, Thailand, and China in southeastern Asia. Psyllidae Macrohomotoma striata Crawford, fig shoot'psyllid’) 29% Decemi993) PWiW. wiles: NEW RECORD, Coloane Island, Macao, ex. Ficus microcarpa and Ficus retusa. In Hong Kong, Hill and Cheung (1978) de- scribed small waxy colonies inhabiting api- cal and lateral shoots of twigs and branches. Hollis and Broomfield (1989) reported it from India and Hodkinson, (1986) listed it from Ryukyu Is. In the family Tropiduchidae Lee and Winney (1981) reported Kallitaxila ma- caoana (Muir) in Hong Kong, but we have not collected it in this study even though the original type locality was in Macao (Muir 1913). ACKNOWLEDGMENTS The authors thank Clive S.K. Lau and Stephen Lai Yue-hong, agricultural officers, Hong Kong Government, for allowing one of us (ERE) access to their insect collection on the Tai Lung Farm near Sheung Shui, New Territories. We are also grateful for species determinations of the Flatidae by John T. Medler of the B.P. Bishop Museum, Honolulu, Hawaii, and to J.P. Duffels, Zo6l- ogisch Museum, Instituut voor Systematiek en Populatiebiologie, Amsterdam, The Netherlands, for determination of the cicad- idae. The delphacid planthopper was iden- tified by Yang Jeng-Tze of the Department of Entomology, National Chung Hsing Uni- versity, Taichung, Taiwan. LITERATURE CITED Blackman, R. L. and V. FE Eastrop. 1994. Aphid’s on the World’s Trees, An Identification and Infor- mation Guide, CAB International, Natural History Museum, London, 986 pp, 16 pls. Cheung, W. W. K. and A. T. Marshall. 1973. Water and ion regulation in cicadas in relation to xylem feeding. Journal of Insect Physiology 19: 1801— 1816. Datta, B. 1988. On Oriental cicadellidae (Homoptera: Insecta). Records of the Zoological Society of In- VOLUME 101, NUMBER 1 dia. Miscellaneous Publications, Occasional Paper No. 90. 256 pp. Easton, E. R. 1991. Annotated list of the insects of Macau observed during 1989. Entomological News 102(2): 105-111. Easton, E. R. 1992. 1990 additions to the annotated list of the insects of Macau. Entomological News 103(1): 30-36. Easton, E. R. and W. W. Pun. 1997. Observations on some Hemiptera/Heteroptera of Macau, southeast Asia. Proceedings of the Entomological Society of Washington 99(3): 574-582. Fennah, R. G. 1956. Fulgoroidea from southern China. Proceedings of the California Academy of Scien- ces 28(13): 441-527. Hayashi, M. 1976. On the species of the genus Mo- gannia Amyot et Serville 1853 (Homoptera, Ci- cadidae) of the Ryukyus and Taiwan. Kontyt 44(1): 27-42. Hayashi, M. 1977. New records of cicadas (Homop- tera, Cicadidae) from Hong Kong. Kontyti 45(1): 91. Hill, D. S. 1975. Agricultural Insect Pests of the Trop- ics and Their Control. Cambridge University Press, Cambridge, U.K., 746 pp. Hill, D. S. and W. W. K. Cheung. 1978. Hong Kong Insects. Urban Council, Hong Kong, 128 pp. Hill, D. S., P. Hore, and I. W. B. Thornton. 1982. In- sects of Hong Kong. Hong Kong University Press, Hong Kong, 503 pp. Hodkinson, I. D. 1986. The Psyllids (Homoptera: Psyl- loidea) of the Oriental zoogeographical region: An annotated checklist. Journal of Natural History 20(2): 299-357. Hollis, D. and P. S. Broomfield. 1989. Ficus-feeding psyllids (Homoptera) with special reference to the Homotomidae. Bulletin of the British Museum of Natural History (Entomology) 58(2): 131-183. Kershaw, J. C. W. 1903. A naturalist’s notes from Chi- na. Field Naturalist Quarterly 2: 233-235. Kershaw, J. C. W. and G. W. Kirkaldy. 1910. A mem- oir on the anatomy and life-history of the homop- terous insect Pyrops candelaria (or candle-fly) Zoologische Jahrbiicher 29(2): 105-124. Kirkaldy, G. W. 1909. A list of the Hemiptera of ori- ental China. Part I. Annales de la Société Ento- mologique de Beligique 53: 177-183. Kuoh, C.-L., Ding, J.-H., Tian Li-X., and C. L. Hwang. 1983. Homoptera: Delphacidae. Economic Insect fauna of China. Fascicle 27. Science Press, Bei- jing 165 pp, 13 pls. (In Chinese.) Lee, L. H. Y. and R. Winney. 1981. Checklist of ag- 105 ricultural insects of Hong Kong 1981. Agriculture and Fisheries Department Bulletin Number 2, Hong Kong, 164 pp. Macau, Catalogue of the Plants of Macau. 1991. Ca- mara Municipal das Ilhas, Instituto de Investiga- ¢ao Cientifica Tropical, Lisboa. 177 pp. Medler, J. T. 1992. Revision of the tribe Phyllphantini in the Oriental Region with descriptions of new genera and new species (Homoptera:Flatidae) Ori- ental Insects 26: 1-38. Mound, L. A. and S. H. Halsey. 1978. Whitefly of the World, A systematic catalogue of the Aleyrodidae (Homoptera) with host plant and natural enemy data. British Museum (Natural History) and John Wiley and Sons, Chichester. 340 pp. Muir, FE 1913. On some new fulgoroidea. Proceedings of the Hawaiian Entomological Society 2(5): 237— 269. Peng, J.-W. and Liu, Y.-Q. 1992. Editors in Chief. Ico- nography of Forest Insects in Hunan, China. In- stitute of Forestry Sciences, 1,473 pp. (In Chi- nese.) Raychaudhuri, D. N. 1980. Aphids of North-east India and Bhutan. The Zoological Society, Calcutta, In- dia. 520 pp. Tang, E-D. 1991. The Coccidae of China. Shanxi Unit- ed Universites Press, Taiyuan, China, 377 pp, 84 pls. (In Chinese.) Wada, T., H. Seino, Y. Ogawa, and T. Nakasuga. 1987. Evidence of autumn overseas migration in the rice planthoppers, Nilaparvata lugens and Sogatella furcifera: analysis of light trap catches and asso- ciated weather patterns. Ecological Entomology 12: 321-330. Webb, M. D. 1981. The Asian, Australasian and Pa- cific Paraboloponinae (Homoptera: Cicadellidae), a taxonomic revision with key to all the known genera of the subfamily. Bulletin of the British Museum of Natural History (Entomology) 43(2): 1-76. Williams, D. J. 1996. A brief account of the hibiscus mealybug, Maconellicoccus hirsutus (Hemiptera: Pseudococcidae), a pest of agriculture and horti- culture, with descriptions of two related species from southern Asia. Bulletin of Entomological Research 86: 617-628. Wu, C. FE 1935. Catalogus Insectorum Sinensium (Cat- alogue of Chinese Insects) Il. The Fan Memorial Institute of Biology, Beijing, China, 634 pp. Yang, C. T. 1989. Delphacidae of Taiwan (II) (Ho- moptera: Fulgoroidea) National Science Council Special Publication No. 6, Republic of China, 334 pp. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 106-122 BIOLOGY OF ANDRENA (SCRAPTEROPSIS) FENNINGERI VIERECK (HYMENOPTERA: ANDRENIDAE), HARBINGER OF SPRING SUZANNE W. T. BATRA Bee Research Laboratory, PSI, Agricultural Research Service, U.S. Department of Ag- riculture, Bldg. 476, BARC-East, Beltsville, MD 20705 Abstract.—Andrena fenningeri Viereck is the first native bee species to fly each spring at Beltsville, MD; males appear as early as February 9, sometimes before their floral hosts begin to bloom. A permanent aggregation of nests in red clay was studied for 10 years. These univoltine, solitary bees break diapause and move at 4°C from their natal cells toward the soil surface in midwinter, ready to emerge and mate as soon as the topmost soil warms. They thermoregulate by aggregating their nests in the warmest available microclimate, and by basking on cold, sunny days to achieve the minimum 11°C required for flight. The most important host is Acer rubrum; Prunus, Pyrus, and Salix are also visited. Male swarming behavior, phenology, nest structure, and associates (including 5 Nomada spp., Myopa sp., and the behavior of 3 unusual species of Eustalomyia [Antho- myiidae]) are discussed. This bee may be manageable as an orchard pollinator, if suitable microhabitat and supplemental hosts are provided. Key Words: conopids, Acer The Holarctic bee genus Andrena in- cludes about 700 Eurasian and 500 North American species; both species and individ- uals are dominant components of the bee fauna during springtime; they are important pollinators of crops and wild plants (see ref- erences in Batra 1990). The behavior and ecology of few species have been studied. Most species of Andrena are solitary, uni- voltine, polylectic bees; many of them share hosts, geographic ranges, and times of adult activity; the reasons why there are so many species of Andrena remain unknown. This report concerns the nesting behavior of an Eastern, cold-adapted, polylectic, sol- itary species, which is the first native bee species to fly each spring at the Beltsville Agricultural Research Center (Prince George’s Co., Maryland). Adult activity of- ten begins while snow and ice remain on bees, nests, thermoregulation, phenology, fruit pollination, Eustalomyia, the frozen ground in shady places, and no plants are yet in bloom. Andrena fenningeri Viereck (det. W. E. LaBerge) is in the North American subgenus Scrapteropsis, which includes 18 vernal species (LaBerge 1971). The nesting behavior of only one of them, A. (Scrapteropsis) alleghaniensis Viereck, has previously been studied (Batra 1990). This solitary bee resembles A. fenningeri in its preference for Acer as a nectar and pol- len source and in the location of its nest aggregation so as to maximize insolation. I investigated the behavior of A. fennin- geri at intervals over a period of 10 years (1987 to 1997), at an aggregation of nests in a sunny spot at the north edge of a large field (the ‘“‘Rose Garden,” off Entomology Rd.). Aggregations of nests of Andrena may persist for decades (Chambers 1968, Schénitzer and Klinksik 1990, 1992, Rid- VOLUME 101, NUMBER 1 dick 1992). If the factors that permit or en- courage permanent aggregations could be determined, this basic information may prove useful in conserving existing natural aggregations, and also for actively manag- ing Andrena bees to pollinate fruit crops. For these reasons, the unusually early adult activity of A. fenningeri was investigated. MATERIALS AND METHODS I made observations early in each season and during fair weather, when bees were flying. A total of 214 hours were spent, most of them during the unusually warm and early spring of 1990 (76 hours), and the cool, late spring of 1992 (60 hours). En- trances to nests with tumuli were marked with small, numbered aluminum tags, in- serted 2 cm north of each nest. Meteorolog- ical data were recorded during visits. Soil temperatures at sites 1-6 were taken by in- serting calibrated bimetallic dial probe ther- mometers to the appropriate depths. Nests were excavated by shovel and trowel after pouring plaster into them, which rendered tunnels easily visible. Adult bees and clep- toparasites were netted, then pinned, or pre- served in FAA for dissection under a mi- croscope. Pollen was stained with lacto- phenol-cotton blue and examined micro- scopically. I indicate means after ranges (in parentheses). Voucher specimens will be deposited in the National Museum of Nat- ural History, Smithsonian Institution, Washington, D. C. NESTS AND THEIR CONTENTS The nests of A. fenningeri in level ground (Fig. 1) were vertical tunnels that penetrat- ed the uppermost humus-and-root-filled zone of the clay soil, to a zone of dense, poorly drained, red marine clay, where the cells were made. Nests ended at the top of a zone of gritty “hardpan.’’ From late fall through early spring, the red clay was very moist, often muddy; after the tree canopy had leafed out in May (after adult bees ceased activity), the upper part of the clay dried to an adobe-like, hard consistency 107 (soil penetrometer reading 3.5—4.5 kg/cm), providing protection to the growing brood and dormant adults in the cells. Evaporation and transpiration by plants during the hot and relatively dry summer months dried the clay. Rainy, cool weather, and the cessation of most transpiration after November per- mitted the soil to moisten and soften. Adult bees emerged from their cells in the soft- ened soil, and crawled toward the surface of the soil during mid-winter. They waited just below the soil surface by late winter, ready to fly during the first warm days of spring. A total of 22 entire nests was examined © 10°19872 350 1989>141n1 991); Because the lateral tunnels that led to cells were backfilled by the solitary mother bees with soil and obliterated after oviposition, many cells that were found could not be traced to their nests’ tunnels (Fig. 1) The nests’ tun- nels reached depths of 16—25 (20.8) cm, and were 4.5—5.5 mm in diameter. Nest en- trances were irregular, 3.5—5.5 mm in di- ameter, and when new, surrounded by a usually circular tumulus of loose, dry, soil particles 2.0—5.0 (3.3) cm in diameter and 0.5—1.0 cm high. The tumulus may not be rebuilt when it had disappeared due to rain. Nest entrances were often closed by loose soil particles when the bees were not for- aging. The entrances to about 20% of the nests, which had been initiated beneath fall- en leaves or in dense turf grasses, were dif- ficult to find. Nests had up to 5 brood cells (1.5) each. They were at the ends of lateral tunnels, which were 1|.0—6.0 (2.5) cm long and 3.5— 5.0 mm in diameter. Cells were made at depths of 13.0—26.0 (21.4) cm. The 39 cells were nearly horizontal, of the usual ovoid shape of Andrena cells, and coated inter- nally with thin, shining, transparent water- proof linings, secreted by Dufour’s glands. Cells were 10.0—12.5 (11.0) mm long, 5.0— 6.0 (5.5) mm in maximum diameter, taper- ing to the cell’s entrance, which was 2.5— 4.0 (3.3) mm in diameter. After oviposition, this entrance was sealed with compressed 108 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Two nests of A. fenningeri in red marine clay soil(stippled area). Nests extended to a zone of denser soil (hardpan, hatched area), at a depth of 23 cm. The lateral tunnels that led to completed cells were filled with soil by the mother bees. Nest A, closed at the surface, was without an adult bee. It had 3 cells with eggs on pollen balls (examined May 4, 1989). Nest B was open, but without a tumulus. Two sealed cells contained eggs on pollen balls; one of these cells (C) had droplets of moisture on its thin, smooth waterproof lining. Another, open cell (D) had a dead female bee with a large, live conopid maggot filling her abdomen. She died in the position taken by bees when they oviposit (nest examined May 7, 1987). VOLUME 101, NUMBER 1 soil pellets, laid down in two concentric rings around a central depression. The Du- four’s-gland secretion in many genera of bees varies in composition. It polymerizes, forming a thin, smooth solid that water- proofs subterranean cells; it can also func- tion as a pheromone, and as food for larvae. In Andrena, it is composed mainly of iso- prenoid esters (see review in Ayasse et al. 1990). Small, regularly-spaced, tasteless (to me) droplets, probably of water, were some- times found on the hydrophobic cell lin- ings, near the entrances of both empty and provisioned cells (Fig. IC). Perhaps the shape of cells influences the condensation pattern, keeping water from condensing on the hygroscopic provisions, where it would permit spoilage. Yeasts in nectar and am- bient fungi ferment and spoil the wetted provisions of many subterranean-nesting bees, causing significant mortality of their larvae (Batra 1970). The soil that seals en- trances to cells and laterals of A. fenningeri is without a visibly water-repellent lining. It is porous, permitting some circulation of the water-saturated subsoil air. The survival of this bee, living inside cells for many months, and the preservation of its hygro- scopic provisions inside humid cells, may depend on such a condensation-site-con- trolling feature of its brood cells. Pollen balls (provisions) of A. fenningeri that had been made in early spring were olive-green, and composed solely of pollen of Acer rubrum L., mixed with nectar. Those made after Acer bloom were various shades of yellow, and made of up to 3 spe- cies of pollen. Pollen balls were firm, smooth, spheroid, somewhat flat on top, and 3.7—4.0 mm high and 4.0—4.8 mm wide (Fig. 1C). Occasionally, laterals and cells with large pollen balls, but no eggs or lar- vae, had been filled with earth. Eggs were strongly arched, white, 2.0—2.7 mm long and 0.5 mm thick. Larvae, ranging from 2.5—4.0 mm long, fed on top of the provi- sions; they later lay on their backs beneath their pollen balls, and reached 8 mm long 109 when all their provisions had been eaten. Larvae transformed to prepupae in late May, after defecation. No cocoons were made. The times of pupation and transfor- mation to adults were not studied. The time of adult emergence each spring, foraging, oviposition, and larval develop- ment varied with weather conditions each year. Although individual females were not marked, many of their nests had been. The appearance of new tumuli late in spring where there had been no nests, the mid-sea- son closure of nests that had been tagged in early spring, and the few cells per nest, in- dicated that some female A. fenningeri make more than one nest each, as do some other species of Andrena. At the end of the nesting season, during the first week of May, the old females became disoriented and exhibited displacement activity, such as random digging, as is seen in Andrena nycthemera Imhoff (Sch6nitzer and Klink- sik 1990). FLORAL Hosts Andrena fenningeri is polylectic (La Berge 1971, Hurd 1979), visiting Acer, Prunus, Pyrus, Salix, and several other hosts. The first host to bloom each spring and the first to be visited for nectar and pol- len at Beltsville was Acer rubrum. This is a dominant tree, occupying about 20% of the forest canopy near the nests. It is an important food resource for a wide variety of insects, available just as the insects emerge from hibernation (Batra 1985). The first provisions made by A. fenningeri were composed of the olive-green pollen from A. rubrum. In this respect, A. fenningeri re- sembles a related species, A. (Scrapterop- sis) alleghaniensis Viereck, which provi- sions its cells with maple pollen in New York (Batra 1990). Several species of ma- ple trees provide nectar and pollen for var- ious other Andrena bees in Europe (Cham- bers 1968) and North America (LaBerge 1971; more references in Batra 1990). After the red maples finished blooming, A. fen- ningeri visited flowers of Prunus, Pyrus 110 and forbs, growing near the aggregation. Thus, the provisions that were made later in spring were yellow. The time of emergence of adult A. fen- ningeri in some years coincided with the beginning of bloom of A. rubrum and Salix sp. In other years, the bees emerged a few days before any host plants had started to bloom. They mated, and began nesting, without having eaten anything. For exam- ple, 2 of 3 females that were starting to ex- cavate nests on February 28, 1992, were in- seminated, but all 3 had empty crops; 14 of 15 males in a mating swarm on that day had empty crops (one had eaten some ma- ple pollen); all of these 18 early bees had large fat bodies in their abdomens (meta- somas) that fueled their flights. By March 2, 1992, 5 males in a mating swarm had empty crops and small fat bodies; 7 females collected while flying over the aggregation still had large fat bodies, but they also had filled their crops exclusively with A. ru- brum pollen, and eggs were developing in their ovaries. The beginning of flowering by A. rubrum varies by up to 2 months with weather conditions in early spring; it begins with male flowers and ends about 2-3 weeks later, with female flowers, the trees being usually dioecious and dichogamous. Bloom began on the following dates at Beltsville: April 2, 1978; March 19, 1982; March 6, 1983 (Batra 1985); February 1, 1989; February 12, 1990; February 21, 1991; February 28, 1992; March 24, 1993 and March 13, 1994. THERMOREGULATION BY AGGREGATION Andrena fenningeri is unusual in its use of dense red clay as a nesting substrate, thus resembling A. macra (Riddick 1990, 1992). It nested in clay, even though a large area of exposed, sunlit sandy soil was within about 100 m. Most species of Andrena nest in more porous, well-drained soils, espe- cially sand (Miliczky and Osgood 1995, Batra 1990); hence they are called *‘Sand- bienen”” in German (Gebhardt and Rohr 1987): PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON On April 21, 1987, when I discovered the aggregation, nests occupied an area of 5 X 8 m. Most nests (53, with tumuli) were in a 3 X 3 m area with exposed soil; it was a buck (deer) scrape, which was renewed ev- ery autumn (Fig. 2, site 1). There were 18 nests/m’, with a minimum internest distance of 1.0 cm. In the part of the aggregation that was in soil covered with short turf (site 2), mests were fewer, spaced up to 1 m apart. On March 8, 1990, the aggregation measured 5 X 34 m; there were up to 54 nests/m? in exposed soil (site 1), with in- ternest distances of 4—18 cm (x 9.5 cm; N 31). In the turf (site 2), nests were 1—4 m apart. On March 5, 1992, the aggregation measured 2 X 25 m; there were 100 nests/ m? at site | and 30 nests/m? at site 2. The aggregation had about 2,900 nests in 1992. By March 1997, the 2 small pine trees near the aggregation had grown, shading it. the bees had moved most nests 2—3 m to the west, where it was sunnier. The winters and early springs of 1997 and 1998 were un- usually rainy. Fewer than 100 nests of A. fenningeri remained at site 1 by late March, 1998 (1 nest/m’ maximum density). Andrena fenningeri nested in the dense aggregation in only one small part of the large field (Fig. 2, Sites 1 and 2). Aggre- gated nesting by solitary bees and wasps is common, and some aggregations may per- sist at a site for over 50 years. There are several, nonexclusive explanations for this phenomenon, including substrate limitation; improved efficiency in foraging; protection from predators and parasites; attraction to others of the same species; an opportunity to save time and energy by re-using existing nests; and philopatry, or re-nesting near the insects’ natal nests (see review in Rosen- heim 1990). Because A. fenningeri was ac- tive during early spring when the weather was often cold and rainy, the location of its aggregation in the warmest and sunniest part of the field appeared to be most advan- tageous; this maximized the number of hours available to the bees for foraging and reproduction. Floral resources were abun- VOLUME 101, NUMBER 1 111 Fig. 2. points North. Regularly mowed areas of short turf lawn are white (if sunny), striped (if partly shaded) or hatched (if always shaded). Stippled areas indicate locations without nests, being boggy, or fenced and shaded by dense tall grass and shrubs. Temperatures were recorded at locations (Sites) 1—6, all in red marine clay. The main portion of the nest aggregation was in a sunny patch of bare soil at Site 1, and it extended into turf-covered soil (Site 2). There were no nests below the short turf at Sites 3—6; Site 3 was shaded all day; Sites 4 and 5 Field where A. fenningeri nested (drawn from aerial photograph). It is 230 m wide, and the arrow were shaded in afternoons and mornings respectively; and Site 6 was sunlit all day. dant, thus not a limitation. The aggregation may have begun decades ago, when a mat- ed founding female successfully nested there, and her descendants returned to the area for nesting (philopatry). Because a closely related, cold-adapted species, A. (Scrapteropsis) alleghaniensis, placed its aggregation of nests in a southeasterly-fac- ing, sandy slope where they received max- imum insolation and warmth in early spring (Batra 1990), A. fenningeri was suspected of doing likewise, even though its aggre- gation was in level clay soil rather than in a sandy slope. In order to test this hypothesis, I made a series of temperature measurements at four depths (2, 8, 15, and 30 cm) at six sites (Sites 1-6, Fig. 2), using bimetallic-dial- probe thermometers. Most of the nest ag- gregation was at Site 1 (in exposed soil); some nests were in Site 2 (soil covered with short turf). A split-plot analysis was used on this ex- periment. The six sites were specifically se- lected for their shading and vegetation cov- erage conditions, and are therefore defined as fixed factors in this analysis. The sites are also defined as whole plots of the split- plot design, while the four levels of depth (2, 8, 15, 30 cm) at each site are the sub- 112 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Means and Standard Errors for the factorial combination of 4 depths and 6 sites. Depth 2 8 15 30 Site Mean? SEM Mean? SEM Mean? SEM Mean? SEM Means | 11.8 0.77 8.8 0.65 6.7 0.63 ell 0.56 8.3° 2 10.0 0.88 8.7 0.73 8.3 0.71 7.4% 0.62 8.6° 3 DES ee 0.82 Dem 0.68 2iOR 0.66 lene 0.59 265 4 Sole 0.89 One 0.73 6.7 0.71 6.1 0.63 6.9° 5 Size 0.89 6:9% 0.73 6.1 0.71 Sal 0.63 6.65 6 O19 0.88 7.8 OB 7.0 0.71 6.7 0.62 7.8¢ 8.44 6.99 6.24 5.6% * A least-squares-mean test examined possible difference between Site-1 (the one with the greatest nest con- centration) means and those of the other five sites. Within a column, a least-squares mean with one asterisk is significantly different from Site 1’s mean at 0.01 = P = 0.05. A mean with two asterisks is significantly different from Site 1’s mean at P < 0.01. > Least-square mean. © Mean of all temperature measurements for each site for all four depths. ‘Mean of all temperature measurements for each depth for all six sites. plot levels. Measurements of temperature done on different days were defined as hap- hazard (a random number table was not used). Data were recorded during early af- ternoon on 7 days from March 3 to March 26, 1992, during the time of peak nesting activity, and used as replication in the anal- ysis. A mixed-model analysis of variance was used to determine the fixed effects of site, depth, and their interaction. A hetero- geneous first-order autoregressive covari- ance structure was included in the ANOVA model to account for the possible associa- tions among depths. Least-significance dif- ference tests (0 = 0.05) were used to com- pare means of the fixed effects. Results are summarized in Table 1. Site | (the aggregation) was significantly warmer than site 3 at all depths (Table 1). It was warmer than Sites 4 and 5 at depths of 2 and 8 cm, but did not differ from them at 15 and 30 cm. Site 1 did not differ from Sites 2 and 6 at depths of 2, 8 and 15 cm, but, at 30 cm, it was slightly cooler than Site 2, and did not differ from Site 6. The right column of the Table shows the means of all measurements combined from the 4 depths at each site. It shows that the portion of the aggregation at Site 2 was slightly (but not significantly) warmer than the ma- jor portion in Site 1, and both of these parts of the aggregation were considerably warm- er than areas where the bees did not nest, especially Site 3. The soil-temperature analysis indicates that A. fenningeri used the warmest local area for the aggregation. Thus, the bees thermoregulated by choosing a warm nest- ing site (and also individually, by basking). Sites | and 2 are at the southern edge of a pine-oak forest. The level ground receives maximum insolation at such a location in March (Kimball and Hand 1922, Geiger 1965). The aggregation also benefitted from radiation that was reflected to it from near- by oak tree trunks (see Geiger 1965). Pine trees within the forest broke the cold north- west wind that prevailed on clear, sunny days in Maryland, and they retained heat that was radiated from the soil at night. The protective influence of a forest windbreak to the north of a field edge can raise air temperatures 5 cm above the ground all year, by 11°C above air temperatures in a field near the forest along the opposite, south edge (Wales 1972). Such microcli- matic differences are significant; for ex- ample, 6 species of vernal wild flowers on a south-facing slope bloomed on average 6 days earlier than the same species growing on a north-facing slope 50 m away; this dif- ference, correlated with cumulative differ- ences in air and soil temperatures, is equiv- alent to 176 km in latitude (Jackson 1966). VOLUME 101, NUMBER 1 There were slight differences within the aggregation between Sites 1 (exposed soil surface) and 2 (turf covered) that were not statistically significant, but could be detect- ed behaviorally when the temperature in the upper 2 cm of soil was near 11°C (marginal for flight initiation). For example, at noon on March 17, 1992, a sunny day after a frosty night, no bees were flying at either Sitelor Site; 2. At Site 1, temperatures were 11°C at 2 cm, 6.5°C at 8 cm, 4.5°C at 15cm, and 4.0°C at.30 cm depth. At 15: 30, bees were flying at Site 1 but were not flying at Site 2. At Site 1, soil temperatures by then were 13°C at 2 cm, 10.5°C at 8 cm, 8.0°C at 15 cm and 5.5°C at 30 cm depth. Temperatures then at Site 2 were too cool for flight, being only 10.5°C at 2 cm, 10.0°C at 8 cm, 9.5°C at 15 cm and 6.5°C at 30 cm depth. Although Site 2 was warmer than Site 1 at 15 and 30 cm, it was cooler at 8 cm, and much (2.5°C) cooler at 2 cm depth, where bees usually wait to warm up, before flying. A similar pattern was seen at 13:00 on March 25, 1992, a day of hazy sun after a frosty night: bees were flying at Site 1 but notat Site 2) (Site: 17) 145°C at 2:em, 90°C ates cmi6.0°C. at 15. cm,-and.5.5°C at 30 cm depth; Site 2: 11.0°C at 2.cm, 10.5°C at 8 cm, 8.5°C at 15 cm, and 7.5°C at 30 cm). Thus, the sun warmed the upper layers of bare soil at Site 1 more quickly than the same depths in turf-shaded Site 2, permit- ting flight from Site 1 on cool, sunny days. The insulating turf retained warmth over- night in the lower layers at Site 2, but this did not promote flight activity. Even such slight differences between sites in bare soil and those in turf may be significant for sur- vival among bees that must fly to forage during inclement weather. Andrena nycthemera also begins adult activity when the soil thaws (as early as February 23 in 1990). Many more bees nested in a sunny part of the aggregation than in a shaded portion, where the soil re- mained frozen longer, and bees in sunlit ar- eas began seasonal activity several days be- fore those that nested in shady portions of 113 the aggregation areas (SchOnitzer and Klinksik 1990). Slight microclimatic differ- ences also influence the nesting behvior of halictine bees (Batra 1997, Potts and Will- mer 1997). THERMOREGULATION BY INDIVIDUALS Species of Andrena that were investigat- ed by Stone and Willmer (1989) produce relatively little endothermic heat, compared to some other genera of bees; instead they depend on insolation and warmth from sub- strates to generate the minimum 8—12°C ambient temperature needed to begin flight. Many species of Andrena bask at nest en- trances and on vegetation before takeoff on sunny, cool days. They include A. fulva Miiller (Paxton 1991), A. erigeniae Robt., which basks on fallen leaves that are 6—9°C warmer than ambient temperatures (Bar- rows 1978), and A. nycthemera, which can begin flights after basking at 8—10°C on sunny days, but cannot fly on cloudy days until the ambient temperature reaches 15°C (Sch6nitzer and Klinksik 1990). Herrera (1995) found that A. bicolor F foraged on sunny days with an air temperature of at least 12—13°C; these small bees bask in warm microclimates to achieve the mini- mum internal thoracic temperature of 22°C needed to begin flight. Both sexes of A. fenningeri were able to begin flight on sunny days when the am- bient temperature of their microclimate was at least 11°C. Bees were often seen basking just inside nest entrances, on tumuli, be- neath or on fallen leaves, and on vegetation. The nearly black bodies of both sexes of A. fenningeri bear pale hairs, which are dens- est over the thorax (where warm-up of flight muscles is needed). In this way, the bees resemble the “‘heat-trap”’ structures of pussy willow flowers, which have been shown by Krog (1955) to absorb short- wave light (solar radiation), which passes through pale hairs into their black surfaces, and there becomes long wave radiation (heat), which is trapped in the dead airspace among the pale hairs. 114 Although A. fenningeri required a mini- mum, insolated temperature of 11°C to be- gin flying, they were active inside their nests at much lower temperatures. On Jan- uary 11, 1990, the soil at the aggregation (Site 1) had thawed after an unusually cold December 1989 (—20°C air temperature for several days, freezing the soil). I excavated twenty-seven cells on January 11, to study the phenology of the bees and determine how they can emerge so early each spring. The air temperature (at | m, in shade) was 10°C, and soil temperatures were 6°C at 2 cm, and 4°C at 10, 21 and 30 cm depths. Some bees had already emerged from their cells (4 6 and 3 ¢ ); 9 males and 11 fe- males were still in their cells (sex ratio near 1:1). Those in cells were resting on their backs. Bees that had emerged stood upright and had dug as far as 2 cm toward the sur- face; the earth that they had excavated was pushed backward, and packed into their na- tal brood cells, covering the fecal layer that had been deposited before their transfor- mation to prepupae late the previous May. When these cold (4°C) bees were disturbed, they vibrated their wings; when they were warmed slightly, they stood upright and be- gan to walk. These bees were stored in tis- sue culture wells at 4°C for 4 days. Some of them defecated (pale meconium), others chewed on moist soil that was placed in the wells with them. Thus, they were active at low temperatures. At the aggregation, air temperatures were above normal for January 1990, and by the 18th, the air temperature at 14:00 E.S.T. (1 m, shade) was 16°C, and soil temperatures were 13°C at 2 cm, 12°C at 10 cm (both above flight threshold), 10°C at 21 cm, and 8°C at 30 cm. During January, the bees worked their way toward the surface, and the first bees (males) were flying on Feb- ruary 9, a sunny day, when the air temper- ature at 14:00 (at 1 m, in shade) was 17°C, and soil temperatures at Site 1 were 11.5°C at 2 cm, 10°C at 10 cm, 8.5°C at 21 cm and 7.5°C at 30 cm. Acer rubrum, Draba and Salix had just begun to bloom (Fig. 3). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON These early males flew slowly, about 2— 7 cm above the aggregation (Site 1 only), frequently stopping to bask on dead oak leaves on the ground. Flying males dropped to the ground when clouds obscured the sun and when disturbed by the observer; they could then be captured by hand. On Feb- ruary 12, more nests were examined. One female had partly emerged from her cell at a depth of 17 cm, 3 others had emerged, and were in the soil at depths of 3, 9, and 11 cm, having moved toward the surface. No males remained in their cells; 2 were found in the soil at 3 and 6 cm depths. Soil temperatures were 11°C at 2 cm, 8°C at 20 cm and 6°C at 20 and 30 cm. Only males were seen again the next afternoon (sunny, 15°C), circling low over the aggregation in calm air, but they settled on the ground and crawled around when gusts of wind or clouds arrived. The males made intermittent circular flights over small areas (20—30 cm radius), which gradually moved across the aggregation. No territoriality or male ag- gression was seen, and the males were not seen in nearby trees, where they formed mating swarms later in February. Female A. fenningeri released a pungent citrus odor from their mandibular glands when han- dled, but males had little odor. Possibly the patrolling males can smell the mandibular gland pheromone, released as the females dig their way toward the surface. On February 28, 1991, male behavior was similar. Fresh tumuli indicated that some females had begun nesting, but none were flying. Many males were patrolling near the ground in a zigzag pattern, often entering and leaving nests. When shaded, they dropped to the ground, and could be picked up by hand; they could not fly even though they buzzed vigorously, an activity that should warm their flight muscles. They were able to resume flight when replaced in the sun for about a minute. These small males could warm up in the sun at marginal temperatures more quickly than could the larger females, which may explain the ab- sence of flying females. The air temperature VOLUME 101, NUMBER 1 r30C Fig. 3. Temperatures at the nest aggregation (Site 1) and phenology of A. fenningeri from January 11 through May 1, 1990, a warmer than normal spring. A, air temperatures at | m height in shade; U, underground soil temperatures at a depth of 30 cm. Bees began emerging from brood cells in January; then crawled up the tunnels of their natal nests in early February. Red maples (R bar) bloomed from February 9 to March 21. The first bees (males) emerged February 9; female bees began collecting pollen from red maple by February 22; when maples finished blooming, bee activity declined and females switched to other pollen sources, including fruit trees (F). By May 4, the forest canopy had leafed out, the clay soil at the aggregation had dried and hardened, and only a few senile bees remained. From March 6-15, a record heat wave, with southwest winds, prevailed (H), and the bees were unusually active; on March 16, a cold front brought rain, and snow (S) fell on March 20 and on March 24-25. (14:30, in shade at 1 m) was 13.5—14.0°C; the soil (Site 1) at 2 cm was 10°C; at 20 cm, 7.5°C and at 30 cm, 7°C. Males that had dropped to the ground to avoid wind often crawled beneath sunlit, dead, dry oak leaves, where temperatures were 12—15°C. In early March, in bright sun, temperatures under sunlit oak leaves reached 22°C when the ambient air temperature was 18°C. In early March 1992, some males that had been flying among trees in the mating swarm basked in the sun on dead leaves, but they crawled under the leaves when the wind gusted or clouds passed; some slept overnight in nest tunnels. When the air tem- perature at 1 m (shade) on a hazy day was at the 11°C threshold for flight, no female bees flew, but males were able to fly slowly for short distances between bouts of bask- ing in the weak sunlight. On a cloudy day, with air at 11°C, the temperature was only 10°C below a fallen leaf, and also at 2 cm in the soil; insolation was insufficient on cloudy days, thus, no bees flew. Because A. fenningeri made more nests in the portion of the aggregation that was 116 in bare soil (Site 1) than in the part that was in short (2—4 cm) turf (Site 2), and because bees emerged earlier each spring at Site 1 than at Site 2, soil temperatures at Site | were compared with those at Site 2. Al- though no statistically significant tempera- ture difference between Sites | and 2 was detected in March, 1992 (see Table 1), there were small differences between the sites that may have been sufficient to account for the slightly earlier, and slightly more, activ- ity at Site 1, especially early in the spring, when temperatures marginally permitted flight. Temperatures were measured at Sites 1 and 2 at the same time of day (early after- noon) on 13 days in March and April, 1990 and 1993. Site 2 was cooler than Site 1 by a mean of 2.6°C at 2 cm, and by 1.1°C at 8 cm; Site 2 was warmer than Site 1 by 0.4°C at 15 cm and by 1.4°C at 30 cm depth. Evidently, the upper levels of bare soil (Site 1) warm more rapidly each sunny day than the same levels under grass, but the insulating grass (Site 2) retains warmth at lower levels overnight. The difference between Sites 1 and 2 was most prominent during sunny weather, for example on March 8, 1990, a sunny day after a clear, cold night, at 2 cm depth (where bees wait before flights), Site 1 was 14°C (warm enough for bees to fly), but Site 2 at 2 cm depth was only 10°C, which was below the flight threshold. However, after a several days of rain on a cloudy day, there was little difference between Sites | and 2. The behavior of male A. fenningeri in mating swarms was also influenced by mi- croclimate. At first, males flew only a few centimeters above the warm ground, where most nests were aggregated. As the air tem- perature warmed, many males moved their patrolling to the tips of the branches of small (4—5-m-tall) pine trees near the ag- gregation, where they formed swarms. On February 28, 1992, males swarmed around the pine trees at an air temperature (1 m, shade) of 17.5°C. When clouds or haze ar- rived, the males suddenly ceased flying, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON alighted, and crawled between the dark green pine needles (which would retain heat); they resumed their flights when the sun reappeared. BEHAVIOR OF MALES The behavior of male Andrena bees has been described and reviewed by Barrows (1978), Gebbardt and RO6hr (1987), and Hallmen (1991). Males may emerge simul- taneously with, or earlier than, their females each year. They swarm conspicuously in the sunshine, circling and zigzagging above ag- gregations of nests, around flowers of host plants, and around tall landmarks, such as selected trees near aggregations. Males of most species are non-territorial, and they jointly patrol in search of females, without noticeable interactions among males. Males of A. nycthemera are unusually aggressive, biting each other and competitively digging in the ground, searching for emerging fe- males (Sh6nitzer and Klinksik 1990). De- spite the large numbers of both sexes that emerge and mate within a few days, and the numerous times males are seen pouncing on females and on various small dark objects, actual copulation is surprisingly rarely ob- served. I also did not observe the mating of A. fenningeri, in spite of many hours spent watching their behavior. Both sexes of A. fenningeri produce a lemon-like odor from their mandibular glands when captured. It is most distinct in females. The mandibular gland secretions of some Andrena bees contain complex mixtures of spiroacetals, monoterpenes, and other compounds (Bergstr6m and Tengo 1982). The secretion of A. fenningeri prob- ably includes geraniol and citronellol, which are major components in other spe- cies of Andrena (Bergstr6m and Tengéd 1982). Male bees use these secretions to mark the areas that they patrol in search of females (Bergstr6m and Tengéd 1982, Geb- hardt and RGhr 1987, Hallmen 1991); and female bees use both mandibular-gland and Dufour’s-gland secretions to mark and iden- VOLUME 101, NUMBER I tify their nests (Ayasse et al. 1990; Stein- mann 1990). During an unusually warm spring, the first male A. fenningeri emerged on Febru- ary 9, 1990, before the females. They emerged as late as March 7, in 1993, also before the females appeared. Some years, males flew before any host plants bloomed; in other years, the first bees appeared when hosts started blooming. Andrena nycthem- era males also may begin flying before any food is available (SchOnitzer and Klinksik 1990). During their first activity, males crawled, or patrolled, flying in the sunshine some 2—8 cm above the bare soil at Site 1, which radiated heat, warming the relatively calm layer of air near the ground. They flew upwind in small circles or in slow, waver- ing patterns, occasionally waiting on fallen leaves when clouds passed. They also fell to the ground and “froze” when alarmed (by my movements) and could be caught with the fingers. On cool days, newly emerged females also displayed such than- atosis, when disturbed, becoming immobile, dropping to the ground, with legs held stiff- ly, parallel and close to their bodies, a be- havior similar to that of elaterid beetles. Sometimes, such bees relaxed, and slowly crawled beneath dead leaves. This behavior was seen only in early spring, before host plants began to bloom. Perhaps it is an en- ergy-conserving defense mechanism, used at a time when the bees were subsisting on their stores of fat. Male A. fenningeri were seen entering and leaving exit holes and nest entrances. Some males slept in holes, but they did not dig for females. Occasion- ally, both males and females crawled on the surface of the aggregation, but mating was not seen there. Most females emerged each year after one or more warm, sunny days, and fresh tumuli indicated that nesting had begun. At this time, many male A. fenningeri ceased patrolling at the aggregation, and instead, they swarmed among the bare branches of an oak tree and around two small pines (P. virginiana Mill.) growing next to the ag- 117 gregation. Males of both Nomada sayi Robt. and N. perplexa Cress. swarmed to- gether with the males of A. fenningeri, both at the aggregation and around the trees; per- haps they have similar pheromones. Male A. fenningeri were seen to emerge from the ground and fly up to join these others around the trees. Males of A. fenningeri patrolled areas around the tips of the pine branches. They zigzagged along the downwind sides of the branches at a height of 2—4 m, occasionally alighting. They flew from one branch tip to the next, briefly hovering while facing the tip of each branch, before flying upwind to the next branch, where they zigzagged to- ward its tip. This was repeated, until they reached the upwind (and sunny) side of the tree, from which they drifted back on the breeze, to repeat the process of inspecting branch tips. Mating was not observed at these “‘swarm trees.”’ Similar upwind and zigzag patrolling, alternating with down- wind drifting, occurs in A. vaga (Hallmen 1991): On March 2, 1992, an attempt to attract males to females in a swarm was made. Four live females were tied with threads to the tips of pine branches where many males patrolled. They were conspicuous, but were bypassed by 16 males. Another 7 males hovered to briefly inspect them, and 4 males pounced on the females, but imme- diately released without mating. These fe- males orally released a distinct lemon-like odor. Other bees that were swarming around the pine trees were netted, yielding 10 males and 1 female, indicating that fe- males were also attracted to these “‘swarm trees’’ (see Hallmen 1991). The captured female and 5 males were dissected; the fe- male was inseminated and had fed on maple pollen, but the males had not eaten pollen or nectar, their crops being collapsed and their guts empty. A sample of 6 females that were flying over the aggregation on March 2 had all eaten maple pollen, and all were inseminated. Similarly, on February 28, 1992, 14 males that were swarming 118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON around the pine trees were dissected; 12 had empty crops and guts, | had eaten nectar, | had eaten some maple pollen; and most had large fat bodies. The females that were col- lected from nests with tumuli on that day varied: | had not mated or eaten, and still had rectal meconium; 2 were inseminated but had not eaten. The number of males seen around the swarm trees and above the aggregation each year slowly declined as spring advanced, and a week to 10 days af- ter emergence, males were no longer seen near the aggregation. By this time, host plants were blooming, and the males searched for females there. PARASITES AND ASSOCIATES As with other species of Andrena world- wide, A. fenningeri was associated with several other insects at the aggregation. These included the halictine bee, Dialictus versatus (Robertson), nesting at the western edge of the aggregation, and its cleptopar- asite, Sphecodes stygius Robertson. Five species of Nomada were active at the A. fenningeri aggregation, which I identified as N. bella Cresson, N. cressoni Robertson, N. parva Robertson, N. perplexa Cresson and N. sayi Robertson. Nomada perplexa was by far the most abundant of these clep- toparasitic anthophorid bees. They often en- tered nests, but I did not find any Nomada in brood cells. A conopid fly (probably My- opa sp.), bombyliid flies, blue and black fungi in brood cells, and three species in the genus Eustalomyia (Anthomyiidae) also were present. These anthomylids were at- tracted to flying A. fenningeri, but not to D. versatus, which had a different, more erratic flight pattern. The grouping of nests in a perennial ag- gregation permitted a permanent population of these natural enemies, but the bees had several defenses against them. For example, many nests were initiated beneath dead leaves, which concealed their entrances; tu- muli often were not rebuilt when they had been destroyed by rain, which made nest entrances inconspicuous; nest entrances were temporarily sealed with soil particles; returning foragers dodged cleptoparasites that followed them; foragers took circuitous routes, and entered nests abruptly, thus con- fusing and evading trailing parasitic bees and flies. Such evasive techniques have been seen in other species of solitary bees and wasps (Hager and Kurcezewski 1985, Meyer-Holzapfel 1986, Rosenheim 1990, Schonitzer and Klinksik 1990). The most abundant of all the parasites were three Eustalomyia species (det. W. Downes, Jr.). The only hosts previously known for this genus of anthomyiid flies are the solitary crabronine wasps, Ectemnius paucimaculatus (Packard) in North Ameri- ca (Krombein 1964), and E. cavifrons (Thomson) in Europe (Meyer-Holzapfel 1986). According to Downes (in litt.), the three species that are associated with A. fen- ningeri may be the first to be associated with bees, and probably are new records, because these flies appeared much earlier in spring than those species that are associated with crabronine wasps. The Eustalomyia species appeared in succession; of samples sent for identifica- tion, the first (species A, males) being ac- tive at the aggregation as early as March 9, 1992, 12 days after the first bees emerged that year. Females of species A were active at the aggregation March 15—22, 1990, and March 25, 1988; species-A males were found as late as April 27, 1988. Females of species B were collected at the aggregation on April 4, 1988, and females of species C were present on April, 27, 1988. The behavior and gross morphology of all three species of Eustalomyia were sim- ilar. It was not possible to distinguish among them in the field; thus, they will be discussed collectively here. Initially, they were assumed to be the anthomyiids, Leu- cophora obtusa (Zetterstedt), and L. mary- landica (Malloch), which are common at aggregations of three species of Colletes bees at Beltsville, and behave similarly to Eustalomyia. Leucophora obtusa also par- VOLUME 101, NUMBER 1 asitizes Andrena nycthemera (Sch6nitzer and Klinksik 1990). Cleptoparasitic anthomyiid and sarcoph- agid flies are often classified according to their behavior in relation to their hosts. Some are “‘satellite flies,” also called “‘sta- tion takers,” which perch near the nests of hosts, abruptly taking flight to pursue re- turning foragers to their nests. Others are ‘hole searchers,” flying about in search of hosts’ nests (Hager and Kurczewski 1985, Weislo 1986, Meyer-Holzapfel 1986). The host bees and wasps defend their nests against satellite flies by evasive maneuvers in flight. Intense activity by both hosts and parasites at dense aggregations also causes confusion of the parasites. Closure and con- cealment of nest entrances provide some protection from hole-searching flies and the cleptoparasitic bees, Nomada and Spheco- des (see review in Rosenheim 1990). A 1-m? area in the densest part of the aggregation (54 nests/m* on March 8) was watched for 74 minutes on a warm, sunny day (10:10—11:24, March 15, 1990), in or- der to observe the activities of cleptopar- asites, especially Eustalomyia. During this time, A. fenningeri foragers left nests 4 times and returned with yellow pollen 10 times (one trip lasted 14 minutes); two bees remained at nest entrances; usually, one No- mada parva continually patrolled above the nests; once one alighted briefly, to fan her wings at a nest entrance (probably to bring up odors from the nest). Most of the time, two Eustalomyia were perched in the area. The flies jumped up and briefly followed patrolling Nomada twice; they chased each other once; and one briefly followed a falling leaflet. One or two flies 10 times followed A. fenningeri, while both leaving and returning to their nests. Flies were seen entering A. fenningeri nests 3 times. In one instance, two flies followed a bee returning with pollen; one of them alighted and peered into the nest headfirst, then turned and slowly backed into the nest, until it disappeared. After 4 minutes, it re- appeared at the entrance, waited 10 sec- in Europe VS) onds, and flew away (the bee remained in- side). Another time, two flies followed a bee returning with pollen (she zigzagged in an attempt to lose them); one fly followed this bee into her nest headfirst, but left with- in a minute. Once, two flies followed a bee laden with pollen to her nest entrance, which was beneath a fallen leaf. On alight- ing near the nest, the flies shoved each other for 1—2 seconds, until one of them flew away. The “winner” then slowly backed tailfirst into the nest entrance, rested there for 2 minutes, then backed down into the tunnel, out of sight. After another 4 min- utes, the fly’s head reappeared at the en- trance, the fly walked out of the nest and rested for 2 minutes facing away from it, cleaned itself, and departed. The bees defended their nests in several ways. The entrances to many nests were closed with plugs of soil or obscured by loose tumuli. Entrances to other nests were camouflaged by, or hidden beneath, dead leaves, grass blades, and twigs. Bees that appeared to be ready to leave their nest en- trances on foraging trips were hesitant, backing down when other insects flew near- by; departing bees were briefly followed by flies, but they did not seem to take evasive paths. Ten bees, returning to their nests that were followed by Eustalomyia, performed elaborate evasive maneuvers before abrupt- ly diving into their nests’ entrances. In con- trast, two returning bees that were not fol- lowed by flies entered their nests directly. During evasive flights, returning bees pur- sued by flies zigzagged; one of them left the area for 2 minutes, returning to her nest again without the fly; another bee shook off two closely-following flies when she crawled beneath a leaf to the hidden en- trance of her nest; a third bee had nested in the shadow of a large piece of debris; the pursuing fly did not follow her, but alighted in a sunny area nearby. Another bee had nested in turf; this bee hit the grass blades and fell to the ground when she tried to zigzag to evade two pursuing flies. The placement of nests in shadows beneath de- 120 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bris may help prevent Eustalomyia attacks, but this may also be disadvantageous be- cause less warmth (insolation) would be present at such shady sites. Adult Eustalomyia spp. first appreared at the aggregation soon after the bees began to emerge. Nomada perplexa emerged at the same time as their hosts, and even shared mating aggregation sites with A. fen- ningeri. On cool, sunny days in early spring, Eustalomyia, Nomada, and _ their hosts basked on the warm soil at the aggre- gation. The initiation of adult activity by these insects in February through March varied by up to a month in different years, but the time of its termination each year in early May varied by only about a week. Only one small maggot was found (May 4, 1989), although 22 nests of A. fenningeri were examined, and Eustalomyia were abundant. It was on the side wall of a sealed cell that contained a normal egg on an in- tact pollen ball, in a marked nest. This nest was closed at the surface, with only this one cell, at the end of a backfilled lateral. This cell was being provisioned on April 21, and Eustalomyia had been very active at the ag- gregation. At that time, this nest had a tu- mulus; probably it was a new, secondary nest. There was no bee in this closed nest when it was examined on May 4. The genus Eustalomyia is ovoviviparous (Meyer-Hol- zapfel 1986). Those species associated with crabronine wasps oviposit at the host’s nest entrances; the fly larvae hatch immediately, and crawl to the wasps’ food stores (Meyer- Holzapfel 1986). In contrast, the species that are associated with A. fenningeri enter the bee’s nests, even while the adult bee is still inside, and they somehow manage to deposit their eggs (or perhaps larvae) inside cells that have been completed, but not yet sealed (the cell plug and long, backfilled lateral would probably prevent these deli- cate flies and their maggots from entering sealed cells). Because solitary bees imme- diately begin to seal their brood cells and backfill laterals on completing oviposition, which occurs after a period spent smooth- ing and perfecting the pollen ball (Batra 1970), it would be difficult for the flies to seize this opportunity. If flies oviposited in cells before pollen balls are completed, the gyrations and grooming of the pollen ball by the bees working in the confinement of their cells would damage fly eggs or larvae, and if detected, the bees may kill, eat, or remove them. On March 22, 1990, one Eu- stalomyia fly that had backed into a nest that was occupied by a bee, as if to oviposit, rapidly emerged while buzzing its wings, agitated, as if it had been attacked by the defensive bee. It then waited, 1 mm from the nest entrance and facing it, for the next 33 minutes. Perhaps the flies deposit their eggs in crevices in open laterals and the hatchling larvae slip into the open cells while the bees are temporarily quiescent, during the process of oviposition. Conopid flies also parasitize these bees. On May 7, 1987, two dead female A. fen- ningeri were found, one in each of two nests. In both instances, the bee, with swol- len abdomen, was poised over a completed pollen ball, as if to begin oviposition (Fig. 1D). One of them contained a brown pu- parium; a large maggot emerged from the pulsating abdomen of the other bee 30 min- utes after her collection. Evidently, the pressure of the maggots inside the bees mimicked the stimulus of eggs that were ready to lay, causing the bees to prepare normal-appearing pollen balls, and take the position for egg laying, as they died (there may have been additional stimuli). Both nests had other cells with eggs on pollen balls, suggesting that these bees had laid eggs soon before the rapidly-growing mag- gots consumed most of their abdominal contents. According to Smith (1966), An- drena bees are usually parasitized by spe- cies of conopid flies in the genus Myopa; host bees may actively fly and feed with a large maggot nearly filling the abdomen; death of the bee occurs shortly before the maggot’s pupation. VOLUME 101, NUMBER 1 CONCLUSION Andrena fenningeri is a species of soli- tary, vernal, univoltine bee that successfully exploits ecological resources that become available as soon as the ground thaws. In order to be prepared for the earliest bloom, this bee nests in permanent aggregations in the warmest available microhabitat. Over- wintered adults begin to move out of their natal brood cells toward the surface in mid- winter. On emergence from nests, they mate, often before food is available, and soon females begin to excavate new nests in the aggregation. When the temperature is marginal for flight, these black bees bask in the sun, to gain sufficient warmth for their activities. This species first feeds on early-bloom- ing red maple and willow, but later shifts to pear and peach as they begin to bloom. It may be possible to manage this species for orchard pollination, if suitable nesting sites and early-season hosts are provided. Areas of sunny, exposed, level clay soil along the north edges of orchards could be prepared and kept free of vegetation. They should be backed along their north sides by windbreaks, such as forests, walls, or solid fences. Red maples and willows should grow within 100 m. The initial population of bees could be obtained by transplanting cores of soil that contain nests from an ex- isting aggregation, as is done to move soil- dwelling alkali bees (Batra 1970). Once a population of bees is established near the orchard, it would become permanent and maintenance-free, except for the need to re- move (scrape) all vegetation from it each winter to permit maximum insolation in early spring. ACKNOWLEDGMENTS I thank Dr. L. W. Douglass and P. Pena- fiel of the Statistical Consulting Services, USDA, Beltsville, for the statistical analy- ses of soil temperatures. Dr. W. E. La Berge of the Illinois Natural History Survey kind- ly identified A. fenningeri. Dr. W. L. 12] Downes, Jr. of Grand Rapids, Michigan, identified the Eustalomyia and provided in- formation about this uncommon genus of flies. Reviews by Drs. E. M. Barrows of Georgetown University and B. B. Norden of the National Museum of Natural History improved this publication. LITERATURE CITED Ayasse, M., R. Leys, P. Pamilo and J. Tengo 1990. Kinship in communally nesting Andrena (Hyme- noptera: Andrenidae) bees, indicated by the com- position of Dufour’s gland secretion. Biochemical Systematics and Ecology 18: 453-460. Barrows, E. M. 1978. Male behavior in Andrena eri- geniae (Hymenoptera: Andrenidae) with compar- ative notes. Journal of the Kansas Entomological Society 51: 798-806. Batra, S. W. T. 1970. Behavior of the alkali bee, Nomia melanderi, within the nest. Annals of the Ento- mological Society of America 63: 400—406. . 1985. Red maple (Acer rubrum L.), an im- portant early spring food resource for honey bees and other insects. Journal of the Kansas Entomo- logical Society 58: 169-172. . 1990. Bionomics of a vernal solitary bee, An- drena (Scrapteropsis) alleghaniensis Viereck, in the Adirondacks of New York (Hymenoptera: An- drenidae). Journal of the Kansas Entomological Society 63: 260-266. . 1997. Bionomics of Lasioglossum (Evylaeus) matianense (Bltithgen) (Hymenoptera: Halicti- dae), the predominant pollinating bee in orchards at high altitude in the Great Himalaya of Garhwal, U.P. India. Proceedings of the Entomological So- ciety of Washington 99: 162-170. Bergstrom, G. and J. Teng6. 1982. Multicomponent mandibular gland secretions in three species of Andrena bees (Hymenoptera, Apoidea). Zeit- schrift fiir Naturforschung 37c: 1124-1129. Chambers, V. H. 1968. Pollens collected by species of Andrena (Hymenoptera: Apidae). Proceedings of the Royal Entomological Society, London (A) 43: 155-160. Gebhardt, M. and G. Rohr. 1987. Zur Bionomie der Sandbienen Andrena clarkella (Kirby), A. ciner- aria (L.), A. fuscipes (Kirby) und ihrer Kuckucks- bienen (Hymenoptera: Apoidea). Drosera 87: 89— 114. Geiger, R. 1965. The Climate Near the Ground. Har- vard University Press, Cambridge. 611 pp. Hager, B. J. and EK Kurczewski. 1985. Cleptoparasitism of Ammophila harti (Fernald) (Hymenoptera: Sphecidae) by Senotainia vigilans Allen, with ob- servations on Phrosinella aurifacies Downes (Diptera: Sarcophagidae). Psyche 92: 451-462. Hallmen, M. 1991. Einige Beobachtungen zum Flug- 122 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON verhalten von Drohnen an einer Kolonie der So- litarbiene Andrena vaga Panzer (Hymenoptera: Andrenidae). Nachrichten des Entomologischen Vereins Apollo, Frankfurt/Main N. F 12: 107— 120. Herrera, C. M. 1995. Floral biology, microclimate, and pollination by ectothermic bees in an early-bloom- ing herb. Ecology 76: 218-228. Hurd, P. D., Jr. 1979. Apoidea, pp. 1741-2209. In Krombein, K. V., D. R. Smith, and B. D. Burks, eds., Catalog of Hymenoptera in America North of Mexico, Vol. 2. Smithsonian Institution Press, Washington, D. C. Jackson, M. T. 1966. Effects of microclimate on spring flowering phenology. Ecology 47: 407—415. Kimball, H. H. and I. EK Hand. 1922. Daylight illumi- nation on horizontal, vertical, and sloping surfac- es. Monthly Weather Review 50: 615-627. Krog, J. 1955. Notes on temperature measurements in- dicative of special organization in Arctic and Sub- arctic plants for utilization of radiated heat from the Sun. Physiologia Plantarum 8: 836-839. Krombein, K. V. 1964. Natural history of Plummer’s Island, Maryland. XVII. The hibiscus wasp, an abundant rarity, and its associates (Hymenoptera: Sphecidae). Proceedings of the Biological Society of Washington 77: 72-112. La Berge, W. E. 1971. A revision of the bees of the genus Andrena of the Western Hemisphere. Part IV. Scrapteropsis, Xiphandrena and Rhaphan- drena. Transactions of the American Entomolog- ical Society 97: 441-520. Meyer-Holzapfel, M. 1986. Vergleichende Betrachtung des Verhaltens brutparasitischer Fliegen aus zwei Familien bei solitaren Aculeaten Hymenopteren (Methodische Kritik). Milleilungen der Schwei- zerischen Entomologischen Gesellschaft 59: 95— 110. Miliczky, E. R. and E. A. Osgood. 1995. Bionomics of Andrena (Melandrena) vicina Smith in Maine and Washington, with new parasite records for A. (M.) regularis Malloch and a review of Melan- drena biology. Journal of the Kansas Entomolog- ical Society 68: 51-66. Paxton, R. J. 1991. Profile of a solitary bee: Andrena fulva. Bee World 72: 11-18. Potts, S. G. And P. Willmer. 1997. Abiotic and biotic factors influencing nest-site selection by Halictus rubicundus, a ground-nesting halictine bee. Eco- logical Entomology 22: 319-328. Riddick, E. W. 1990. Andrena macra Mitchell (Hy- menoptera: Andrenidae) overwinter and delay spring emergence in Virginia. Proceedings of the Entomological Society of Washington 92: 771— M2. . 1992. Nest distribution of the solitary bee An- drena macra Mitchell (Hymenoptera: Andreni- dae), with observations on nest structure. Pro- ceedings of the Entomological Society of Wash- ington 94: 568-575. Rosenheim, J. A. 1990. Density-dependent parasitism and the evolution of aggregated nesting in the sol- itary Hymenoptera. Annals of the Entomological Society of America 83: 277-286. Schnitzer, K. and C. Klinksik. 1990. The ethology of the solitary bee Andrena nycthemera Imhoff 1866 (Hymenoptera, Apoidea). Entomofauna, Zeit- schrift fiir Entomologie 11: 377—427. 1992. Individuell unterschiedlicher Leben- slauf bei der Sandbiene Andrena nycthemera Im- hoff. Nachr Blatt Bayerischen Entomologie 39: 116-121. Smith, K. G. V. 1966. The larva of Thecophora occi- densis, with comments on the biology of Conopi- dae (Diptera). Journal of Zoology, London 149: 263-276. Steinmann, E. 1990. Zur Nahorientierung der solitaren Sandbiene Andrena vaga, Panzer 1799 (Hyme- noptera, Apoidea) am Nesteingang. Bulletin de la Société Entomologique Suisse 63: 77-80. Stone, G. N. and P. G. Willmer. 1989. Warm-up rates and body temperatures in bees: the importance of body size, thermal regime and phylogeny. Journal of Experimental Biology 147: 303-328. Wales, B. A. 1972. Vegetation analysis of north and south edges of a mature oak-hickory forest. Eco- logical Monographs 42: 451-471. Weislo, W. T. 1986. Host nest discrimination by a clep- toparasitic fly, Metopia campestris (Fallén) (Dip- tera: Sarcophagidae: Miltogramminae). Journal of the Kansas Entomological Society 59: 82-88. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 123-137 REVISION, CLADISTICS AND BIOGEOGRAPHY OF THE NEOTROPICAL GENUS SOUZALOPESMYIA ALBUQUERQUE (DIPTERA: MUSCIDAE) CLAUDIO JOSE BARROS DE CARVALHO Department of Zoology, Universidade Federal do Parana, C.P. 19020, 81531-990, Cu- ritiba, Parana, Brazil (e-mail: cjbcarva@bio.ufpr.br) Abstract.—Souzalopesmyia Albuquerque, a monophyletic Neotropical muscid genus of five species, is reviewed to include two new species, Souzalopesmyia paraensis Carvalho, new species (Brazil: Para), and Souzalopesmyia sulina Carvalho, new spe- cies (Paraguay: Canindeyu). Ground plan characters of the Phaoniinae as outgroup were used in a cladistic analysis of the genus. The phylogenetic relationships found are (S. amazonica (S. paraensis (S. singularis, S. sulina)) (S. carioca)) and these seem to sup- port at least two biogeographical hypotheses: 1, The basal clade, S. amazonica, suggests a date of origin for the genus as far back as the Late Cretaceous; 2, The occurrence of S. paraensis in Belém, along south side of the Amazon River, suggests a single dispersal event to colonize that region, in a more recent time, which belongs to the northwestern track. Resumo.—Souzalopesmyia Albuquerque, um género monofilético de Muscidae Neo- tropical com trés espécies é revisto para incluir duas novas espécies, Souzalopesmyia paraensis Carvalho (Brasil: Para) e Souzalopesmyia sulina Carvalho, (Paraguay: Can- indeyu). Para a andalise cladistica do género, foram utilizados, como grupo de fora, os caracteres do plano basico de Phaoniinae. A partir do relacionamento filogenético encon- trado (S. amazonica (S. paraensis (S. singularis, S. sulina)) (S. carioca)) podem ser re- tiradas, no minimo, duas hipoteses biogeograficas: 1. O clado basal, S. amazonica, sugere a data de origem do género para o Cretaceo Superior; 2. A ocorréncia de S. paraensis em Belém, ao sul do Rio Amazonas, sugere que um unico evento de dispersao ocorreu para colonizar esta regiao, em uma época mais recente. Key Words: Cladistics, biogeography, phylogenetic analysis, Souzalopesmyia, taxonomy Souzalopesmyia Albuquerque 1951 is an unusual genus of rare Neotropical Muscidae because all of its species are yellow. In gen- eral, yellow species are inhabitants of very dark shaded habitats. The genus was pro- posed by Albuquerque (1951: 53) to ac- commodate two new species, S. carioca and S. amazonica. Subsequently, Pont (1972) included Mydaea singularis Stein 1911, in the genus. The relationship of Souzalopesmyia with other genera of Muscidae is confusing. Malloch (1929) described Peruvia (a pre- occupied name, now a synonym of Souza- lopesmyia) to include only Mydaea singu- laris Stein, and considered it to be close to Charadrella Wulp 1896 (Cyrtoneurininae). Séguy (1937) synonymized Peruvia with Mydaea Robineau-Desvoidy 1830 (Myda- einae), and Albuquerque (1951) considered Souzalopesmyia to be close to Oramydaea Snyder, the latter being an Afrotropical ge- 124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nus of Mydaeinae now synonymized with Myospila Rondani 1856. Hennig (1965), af- ter examination of the ovipositor of the type species of Peruvia put it close to Helina Robineau-Desvoidy (Phaoniinae). Carvalho (1989d) included Souzalopesmyia in his cladistic analysis of Muscidae and consid- ered it as genus of Phaoniinae (Pont 1972, Carvalho et al. 1993). The present paper adds two new species to Souzalopesmyia, presents a cladogram of the genus, and reflects upon the biogeo- graphic relationships among the included species. MATERIAL AND METHODS Specimens from the following institu- tions were studied for comparative purpos- es: Department of Zoology, Universidade Federal do Parana (DZUP), Curitiba, Bra- zil; Museu Paraense Emilio Goeldi (MPEG), Belém, Brazil; Museu Nacional do Rio de Janeiro (MNRJ), Rio de Janeiro, Brazil; Staatliches Museum fiir Tierkunde (SMT), Dresden, Germany; Museo Nacion- al de Historia Natural (MNPA), Asuncion, Paraguay. The terminology and abbreviations used here are those in McAlpine (1981) and Car- valho (1989a), and the descriptions of the type-specimen labels follows O’Hara (1982). The sister group of Souzalopesmyia 1s unknown although the genus is considered as one of the most basal members of the tribe Phaoniini (Pont and Carvalho 1997). The sister group may be found in Afrotrop- ical Phaoniini (Pont 1980), which is com- posed of three genera. Two of these genera, Phaonia Robineau-Desvoidy, 1830 and Helina, are true but paraphyletic genera of Phaoniini (Hennig 1965). On the other hand, the sister group may be found in a more basal group of Neotropical Muscidae. Character polarities for the genus were based on the ground plan of Phaoniinae, and the assignment by Pont (1986) since no phylogenetic analysis for these genera is Table 1. Character state distribution among species of Souzalopesmyia. 0 = plesiomorphic character states; | = apomorphic character states; ? = missing data. Taxonomically useful characters for the species of Souzalopesmyia. Characters with an * were used in phylogenetic analysis. outgroup 000 000 000 000 O00 S. amazonica O]OOE ACO) OMRON AO MS teat S. carioca iO) (Oat shail“ aLfo)at shal S. paraensis OO Os aso} OO) iil S. singularis OOO Oia AO@, aaa wil S. sulina BAO) alae OO) ab alal aval *1. Number of frontal setae in female: 0) three; 1) two. . Frontal setae in male: 2) two cruciate; 1) two, the lower cruciate, the upper reclinate. . Vti: O) parallel; 1) divergent. 4. Postocular setae row in male: 0) complete and distinct, whole row of setulae reaching epistome; 1) incomplete, row of setulae reaching only to basal half of eyes. *5_ Postocular setae row in male: 0) whole row of setulae black; 1) composed of black and yellow setulae; latter beginning after basal half of eye and reaching epistome, but 1—2 with black setu- lae. *6. Number of De prst: 0) two; 1) one. *7. Acr female prst: 0) distinct from the ground se- tulae; 1) not distinct from the ground setulae. *8. Proepisternal seta: 0) strong, similar in length to the upper anepisternal setae; 1) weak, less than the length of upper anepisternal. 9. Crossvein dm-cu: 0) almost straight (Albuquer- que 1951: Fig. 11); 1) weakly curved (Albu- querque 1951: Fig. 13). Fifth sternite shape: 0) without sharp depression on posterior side (Fig. 1); 1) with sharp depres- sion on posterior side (Figs. 2, 3, and Albuquer- que 1951: Figs. 10 and 12). *11. Cercal plate: 0) round outline (Fig. 4, and Al- buquerque 1951: Fig. 4); 1) squared outline (Figs. 5, 6). 12. Head appearance: 0) not elongate; 1) elongate (Albuquerque 1951: Fig. 1). 13. Number of ocellar setae; 0) two; 1) none. 14. General ground color of the flies: 0) not yellow; 1) yellow. tN ies) =i: available (Huckett and Vockeroth 1987, Carvalho 1989d). Table | includes all useful characters and character states distribution of Souzalopes- myia species and their polarities used in the present paper. The program Hennig8s6, ver- sion |. 5 (Farris 1988) was used for the phy- logenetic analysis, applying the implicit enumeration (ie*) option. Consistency (CI) VOLUME 101, NUMBER 1 and retention (RI) indices were calculated excluding uninformative characters (auta- pomorphies and synapomorphies of the ge- nus). TAXONOMY Souzalopesmyia Albuquerque 1951 Peruvia Malloch 1929:104 (preocc. Scud- der 1890). Type-species, Mydaea singu- laris Stein, 1911 (orig. desig.). Souzalopesmyia Albuquerque 1951:53. Type-species, Souzalopesmyia carioca Albuquerque, 1951 (orig. desig.). Diagnosis.—Souzalopesmyia may be rec- ognized by its typical head and antenna shape, by the setulose parafacials for half their length, and by the absence of ocellar setae. Also, they are wholly yellow flies ex- cept for the presence of stripes on the ab- domen (Albuquerque 1951: Figs. 3, 16). Description.—Male head dichoptic (AI- buquerque 1951: Fig. 1), narrower than in female. Frons with orbital setae reclinate, and without crossed setae on frontal vitae. Ocellar setae absent. Antenna long, reach- ing epistome. Arista plumose, longest hairs equal to greatest antennal diameter. Para- facials setulose on upper half. Female pro- boscis as in Fig 14. De 1—2:3. Acr not dis- tinct from ground setulae, except in female of S$. amazonica. 2 postpronotals. Ia: 1:2. Sa: 1:2, second weak, about half length of first. 2 pa. 2 subequal np] setae. Pra absent. Disc of notopleuron bare. Anepisternum with a short seta in upper anterior corner. Anepimeron, greater ampulla, vallar ridge, and meron bare. Ktps 1:2 (not 1:1:1 as stat- ed by Albuquerque 1951). Metathoracic spiracle small, triangular, with yellow se- tulae on margin. Prosternum bare. Fore tib- ia with 1 PD submedian setae. Fore tar- somere | with | V setae. Mid femur with 1 AD, 1 D, 1 PD and 1 P preapical setae. Mid tibia with 2—4 P median setae: | strong V apical setae. Hind tibia without calcar and with | AD median setae, | D, 1 AD, 1 AV apical setae. Veins bare, ex- cept for costa. Vein M ,,, curved slightly 125 toward vein R ,,;. Lower calypter of Phaonia-type. Sternite 1 bare. Male aedea- gus as in Figs. 10—12. Ovipositor long, ter- gites, sternites and membranes covered with microtrichia (Figs. 15, 16). Three elongate spermathecae. Egg: Phaonia- type. Monophyly.—Souzalopesmyia Albu- querque is a monophyletic genus based on the following synapomorphies: |, Head lat- eral appearance elongate; 2, Ocellar setae absent; 3, Ground colour yellow. Remarks.—The species of Souzalopes- myia are rare and have similar facies. Based on current collection records, they are found in rainforest habitats. They may be nocturnal, as they are rarely collected by day. KEY TO SPECIES OF SOUZALOPESMYIA 1 Vti divergent. Crossvein dm-cu_ oblique, weakly curved (Albuquerque 1951: Fig. IB) female: 2°trontals? 452 0iseste ota Mel Shade sautn ane © ees nS S. carioca Albuquerque = Vti parallel. Crossvein dm-cu oblique, al- most straight (Albuquerque 1951: Fig. 11): females Gatton talSe-w- cates eme pee Cree 2 De 2: 3; male: postocular row of setulae in- complete, not reaching epistome; setulae black; female: some acr prst stronger than the ground setulae N p= ~ ae one S. amazonica Albuquerque = De, 1:3: complete; setulae black and yellow, the lat- ter beginning after basal half of eye: female: acr prst undifferentiated from the ground se- tULAS RAR eae goton Peeks ee ea scaie Ge eteast ieee 3 3 1 proepisternal seta weak, about %4 length of the upper anepisternal setae; male: fifth ster- nite without sharp depression on posterior side (Fig. 1); cercal plate heart-shaped (Fig. S. paraensis Carvalho, new species = 1 proepisternal seta strong, similar to the upper anepisternal setae; male: fifth sternite with sharp depression on posterior side (Figs. 2, 3); cercal plate rounded (Figs. 5, (3) ere ae erates 2 uc, Shor Mace cS eee ea 4 4 (3) Species ranging from 8.0 to 9.0 mm in length; posterior ktps strong, about 2 times the length of the anterior one; male: frontal male: postocular row of setulae setae both cruciate; fifth sternite as in Fig. 2-acercalaplate as) neg DS) een. A SOE vo eh eee nea S. singularis (Stein) - Species ranging from 6.5 to 8.0 mm in length; posterior ktps very strong, about 3 126 times the length of the anterior one; male: lower frontal setae cruciate, upper reclinate; fifth sternite as in Fig. 3; cercal plate as in BigeiGy cr aetate S. sulina Carvalho, new species Souzalopesmyia amazonica Albuquerque 1951 Souzalopesmyia amazonica Albuquerque 1951:56; Pont 1972:23 (Neotropical cat- alog); Carvalho et al. 1993: 84 (Neotrop- ical catalog). Diagnosis.—This species is very similar to S. singularis but, it can be easily distin- guished from the other Souzalopesmyia species by dc 2:3 setae. Description.—Male: Head: Frons broad, at narrowest point 0.21 of head width. Eye with only normal pubescence. Fronto-orbit- al plate, parafacial, face and gena silvery white. Fronto-orbital plate broad, broaden- ing gradually from vertex to lunula; at ver- tex plate almost equal to diameter of ante- rior ocellus, at lunula equal to three times anterior ocellus. Frontal vitta broad, parallel to vertex. 2 pairs of strong frontal setae on lower % of frons, former cruciate and latter reclinate; 1 strong orbital, reclinate and di- vergent. Ocellar triangle black, reaching to insertion of orbital setae, with some setulae behind posterior ocellar setae. Vti strong and parallel. Postocular row incomplete, reaching as single row just below mid level of eye and composed of black setulae. Gena below lowest eye margin equal to twice di- ameter of the anterior ocellus. Palpi slender, yellow. Thorax: Ground color yellow, scutum dusted with whitish-grey. Dc 2:3; 5—6 rows of prst acr setulae; 11—12 rows of post acr setulae. 1 proepisternal weak, about % length of upper anepisternal setae. Posterior ktps strong, about 2.5 times length of an- terior one. Scutellum with 1 strong pair of apical and 1 subbasal setae; | preapical weak but stronger than ground setulae; disc with setulae descending below strong setae; bare ventrally. Legs: Yellow. Fore femur with complete rows of AV and D setae; AD row weak and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON just to apical %. Mid tibia with 3 P setae. Hind tibia with 5 AV setae on apical half (Albuquerque 1951: 57). Wings: Clear, veins yellow. Membrane entirely covered with microtrichia. Cross- vein r-m placed before point where vein R1 enters costa. Crossvein dm-cu oblique, al- most straight. Calypters and haltere yellow. Abdomen: Ground-color yellow; in pos- terior view; with a narrow black stripe on tergite 1 which is enlarged on tergite 3, 4 and 5 (Albuquerque 1951: Fig. 16). Tergite 4 and 5 with 2 strong apical setae. Terminalia: See Albuquerque 1951: Fig. 12% Measurements: Length of body, 7 mm (n = 1). Length of wings, 6 mm (n = 1). Female: Differs from male as follows: Head: Frons at narrowest point 0.27 of head width. Frontal vitta broad, divergent to vertex. 3 pairs of strong frontals, former 2 cruciate. Ocellar triangle black, not reach- ing to insertion of 3rd pair of frontals. Some acr prst stronger than ground setulae. Cross- vein dm-cu oblique, more than in male. Terminalia: See Albuquerque 1951: Fig. LPNS: Measurements: Length of body, 8 mm (n=1). Length of wings, 8 mm (n = 1). Remarks.—Adults have been collected in the afternoon on flowers and at night. Type material examined.—Holotype ¢ in MNRJ labelled as follow: ““35/Manaus—no centro [in the center of the city]/ a noite [at night]/15.vi.933 [15 June 1933]/Ant. [An- tonio] Paes Filho”’ [hand label]; Souzalo- pesmyia/amazonica sp.n,/30.8.50 [30 Au- gust 1950 [examined Albuquerque’ date] / D. Albuquerque det.’ [Albuquerque hand label]; “‘Holotipo [holotype]’’ [red label]. Specimen in good condition (Lopes et al. 1997). Right fore tarsi, median and hind legs missing. Segments of the abdomen mounted on slide in Canada balsam. Other examined material.—Total: 1. BRAZIL. Amazonas: Manaus, Ant. Paes Filho, 26.V.1933 (1 2 allotype MNR3J). VOLUME 101, NUMBER 1 Souzalopesmyia carioca Albuquerque 1951 Souzalopesmyia carioca Albuquerque 1951:53; Pont 1972: 23 (Neotropical cat- alog); Carvalho et al. 1993: 85 (Neotrop- ical catalog). Diagnosis.—S. carioca is one of the larg- est Souzalopesmyia species and can be dis- tinguished from the other species by the di- vergent vti. The female has only 2 frontal setae. Description.—Male: Head: Frons broad, at narrowest point 0.24 of head width. Eye with only normal pubescence. Fronto-orbit- al plate, parafacial, face and gena silvery white. Fronto-orbital plate broad, broaden- ing gradually from vertex to lunula, at ver- tex plate almost equal to diameter of ante- rior ocellus, at lunula 3 times diameter of anterior ocellus. Frontal vitta broad, parallel to vertex. 2 pairs of strong cruciate frontals on lower % of frons; 1 strong orbital recli- nate and divergent. Ocellar triangle reach- ing to insertion of orbital setae. Vti strong, divergent. Postoculars in a complete single row, reaching to epistome and composed of black setulae. Gena below lowest eye mar- gin equal to twice diameter of anterior ocel- lus. Palpi slender, yellow. Thorax: Ground-color yellow with scu- tum dusted with whitish grey. Dc 1:3; 6—7 rows of prst acr setulae; 11—12 rows of post acr setulae. | proepisternal seta very weak, about half length of upper anepisternal se- tae. Posterior ktps strong, about twice length of anterior one. Scutellum with 1 strong pair of apical and | subbasal setae; 1 preapical weak but stronger than ground setulae; disc with setulae descending below strong setae; bare ventrally. Legs: Yellow. Fore femur with a com- plete row of AV, D and AD setae, latter weakest. Mid femur with 2—3 PV setae in basal half, shorter than femoral depth. Mid tibia with 2—3 P setae. Hind femur with 1 strong preapical AV seta, longer than fem- oral depth; AD row complete; 1 PD, 1 D, 127 1 AD preapical setae weaker than AV seta. Hind tibia with 4 AV setae on apical half. Wings: Clear, veins yellow. Membrane entirely covered with microtrichia. Cross- vein r-m placed before point where vein R1 enters costa. Crossvein dm-cu oblique, weakly curved in median part. Calypters and haltere yellow. Abdomen: Ground-color yellow; tergite 3 and 4 each with blackish markings increas- ing in size to posterior margin in posterior view; tergite 5 blackish, except hind margin (Albuquerque 1951: Fig. 3). A median black stripe on tergites 3, 4 and 5. Tergite 4 and 5 with 2 strong apical setae. Terminalia: See Albuquerque 1951: Fig. 4-10. Measurements: Length of body, 8 mm (n = 1). Length of wings, 8 mm (n = 1). Female: Differs from male as follows: Head: Frons at narrowest point 0.27 times maximum head width. Frontal vitta broad, divergent to vertex. 2 pairs of strong frontals, cruciate. Ocellar triangle black, not reaching insertion of 2nd pair of frontals. Terminalia: See Albuquerque 1951: Fig. 14515: Measurements: Length of body, 8.5 mm (n=1). Length of wings, 8.5 (n = 1). Type material examined.—Holotype ¢ in MNR3J labelled as follow: “‘Grajahu/Rio de Janeiro/Lopes [Hugo de Souza Lopes]- 6.1.40 [6 January 1940]’’; Souzalopesmyia/ carioca sp.n./30.8.50 [30 August 1950] [ex- amined Albuquerque date] /D. Albuquerque det.”” [Albuquerque hand label]; ““Holotipo [holotype]’’ [red label]. Specimen in good condition (Lopes et al. 1997). Right wing on a slide mounted attached of pin holo- type. Segments of abdomen on a slide mounted. Other examined material.—Total: 1. BRAZIL. Rio de Janeiro: Rio de Janeiro, H.S. Lopes, 6.VI.1940 (1 @ allotype MNRJ). Souzalopesmyia paraensis Carvalho, new species Figs: 1,4, 70, 133.14,,15) Diagnosis.—Souzalopesmyia paraensis can be distinguished from the other Souz- 128 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. 4—6, Cercal plate, dorsal view. 4, S. paraensis. 5, S. singularis. 6, S. sulina. alopesmyia species by male cercal plate rounded in outline and fifth sternite without posterior depression. Description.—Male: Head: Frons broad, at narrowest point 0.21 of head width. Eye with only normal pubescence. Fronto-orbit- al plate, parafacial, face and gena silvery white. Fronto-orbital plate broad, broaden- ing gradually from vertex to lunula, at ver- tex plate less than diameter of anterior ocel- lus, at lunula 3 times diameter of anterior ocellus. Frontal vitta broad, parallel to ver- tex. 2 pairs of strong cruciate frontal setae on lower *% of frons; 1 strong orbital recli- nate. Ocellar triangle reaching insertion of orbital setae. Vti strong, parallel. Postocu- lars in a complete single row, reaching ep- istome and composed of black and yellow setulae; latter beginning after basal half of 1—3, Male fifth sternite, dorsal view. 1, Souzalopesmyia paraensis. 2, S. singularis. 3, S. sulina. eye and reaching epistome, but 1—2 with black setulae. Gena below lowest eye mar- gin twice diameter of anterior ocellus. Palpi slender, yellow. Thorax: Ground-color yellow, dusted with whitish grey. Dc 1:3; 7—8 rows of prst acr setulae; 10—11 rows of post acr setulae. 1 proepisternal seta weak, about *%4 length of the upper anepisternal setae. Posterior ktps strong, twice length of anterior one. Scutellum with | strong pair of apical and 1 subbasal setae, | preapical weak but stronger than ground setulae; disc with se- tulae descending below strong setae; bare ventrally. Legs: Yellow. Fore femur with a com- plete row of AV and D setae; AD row weak. Mid femur with 2 PV setae in basal half, not equal to femoral depth. Mid tibia VOLUME 101, NUMBER 1 129 Figs. 7-12. singularis. 9, S. sulina. 10-12, Phallus and associated structures, lateral view. 10, S. paraensis. 11, S. singularis. 12, S. sulina. with 3 P setae. Hind femur with 1 strong preapical AV seta, longer than femoral depth; AD row complete; 1 PD, 1 D, 1 AD preapical seta weaker than AV seta. Hind tibia with 4 AV setae on apical half. Wings: Clear, veins brownish. Membrane entirely covered with microtrichia. Cross- vein r-m placed before point where vein R1 enters costa. Crossvein dm-cu oblique, al- most straight. Calypters and haltere yellow. Abdomen: Ground-color yellow; in pos- terior view with a narrow, slight, black stripe on tergite 1 which is blackish and en- larged on tergites 3, 4 and 5. Tergite 4 and 5 with 2 strong apical setae. Terminalia: See Figs. 1, 4, 7, 10, 13. 7-9, Epandrium, cercal plate and surstylus, lateral view. 7, Souzalopesmyia paraensis. 8, S. Measurements: Length of body, 8.5 mm (n = 1). Length of wings, 6.6 (n = 1). Female: Differs from male as follows: Head: Frons at narrowest point 0.31 of head width. Frontal vitta broad, divergent to vertex. 3 pairs of strong frontals, first one cruciate, second one reclinate and conver- gent and third one reclinate and divergent. Ocellar triangle black, not reaching inser- tion of 3rd pair of frontals. Postoculars in a complete single row, with black setulae. Measurements: Length of body, 8.5—9.0 mm (n = 2). Length of wings, 7.0—7.7 mm (n = 2). Type material examined.—Holotype ¢ in MPEG, labelled as follow: ““‘Belém Mo- 130 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cambo/ 01-IV-1977”; ‘Brasil Para/A.Y. Harada.” Other examined material.—Total: 2 par- atypes: BRASIL. Para: Belém. A. Y. Ha- rada, 1.IV.1977 (1 2 DZUP); ibidem, same collector G:1V:1977 (1 2 MPEG). Speci- men in reasonable condition. Left fore and right mid legs missing; left mid tarsi and hind right leg glued on to a card attached to the pin: Souzalopesmyia singularis (Stein 1911) Gigs. 255, 8: 11-716) Mydaea _ singularis Stein 1911:91; Stein 1919: 124 (world catalog); Séguy 1937: 282 (world catalog). Peruvia singularis; Malloch 1929: 105 (type species of Peruvia); Hennig 1965: Fig. 31 (tip of female ovipositor). Souzalopesmyia singularis; Pont 1972: 24 (Neotropical catalog); Carvalho et al. 1993: 85 (Neotropical catalog). Diagnosis.—Souzalopesmyia singularis can be distinguished from the other Sou- zalopesmyia species by posterior depression on male fifth sternite and 2 cruciate frontal setae. Description.—Male: Head: Frons broad, at narrowest point 0.23 of head width. Eye with only normal pubescence. Fronto-orbit- al plate, parafacial, face and gena silvery white. Fronto-orbital plate broad, broaden- ing gradually from vertex to lunula, at ver- tex plate less than diameter of anterior ocel- lus, at lunula 2.5 times diameter of anterior ocellus. Frontal vitta broad, parallel to ver- tex. 2 pairs of strong cruciate frontals on lower % of frons; 1 strong orbital reclinate. Ocellar triangle reaching insertion of orbital setae. Vti strong parallel. Postoculars in a complete single row, reaching epistome and composed of black and yellow setulae; lat- ter beginning after basal half of eye and reaching epistome, but 1—2 with black se- tulae. Gena below lowest eve margin equal to 2.5 times diameter of the anterior ocellus. Palpi slender, yellow. Thorax: Ground-color yellow with scu- tum dusted with whitish-grey. De 1:3; 5—6 rows of prst acr setulae; 9-10 rows of post acr setulae. 1 proepisternal seta similar to upper anepisternal setae. Posterior ktps strong, twice length of anterior ones. Scu- tellum with | strong pair of apical and 1 subbasal setae; 1 preapical weak, but stron- ger than ground setulae; disc with setulae descending below strong setae; bare ven- trally. Legs: Yellow. Fore femur with complete row of AV and D setae; AD row weak. Mid femur with 2 PV setae in basal half, less than femoral depth. Mid tibia with 3 P se- tae. Hind femur with | strong preapical AV seta, longer than femoral depth; AD row complete; | PD, 1 D, 1 AD preapical setae weaker than AV seta. Hind tibia with 5 AV setae on apical half. Wings: Clear, veins brownish. Membrane entirely covered with microtrichia. Cross- vein r-m placed just before point where vein R1 enters costa. Crossvein dm-cu oblique, almost straight. Calypters and haltere yel- low. Abdomen: Ground-color yellow; in pos- terior view with slight narrow black stripe on tergite 1 which is enlarged on tergite 3, 4 and 5. Tergite 4 and 5 with 2 strong apical setae. Terminalia: See Figs) 245, 8711- Measurements: Length of body, 8 mm (n = 1). Length of wings, 7.5 mm (n = 1). Female: Differs from the male as fol- lows: Head: Frons at narrowest point 0.32 times maximum head width. Frontal vitta broad, divergent to vertex. 3 pairs of strong frontals, first one cruciate, second one rec- linate and convergent and third one recli- nate and divergent. Ocellar triangle not reaching insertion of 3rd pair of frontals. Postoculars in a complete single row of black setulae. Measurements: Length of body, 8.5—9.0 (n = 3). Length of wings, 8.0—8. 5 mm (n = 3). Type material examined. Holotype male in SMT labelled as follow: *‘Bolivia- Mapiri/ 14.11.03 [hand written]/Sarampioni 700 m [green label]: “‘Coll W. Schnuse/ VOLUME 101, NUMBER 1 131 Figs. 13-16. 13-15, Souzalopesmyia paraensis. 13, Hypandrium, dorsal view. 14, Proboscis, lateral view. 15, Ovipositor, ventral view. 16, Ovipositor of S. singularis, ventral view. 1911-3”; ‘“‘Mydaea/singularis/ sp.n. [light Schnuse, III-1903 (2 2 SMT); S. Carlos, W. green hand written label]: ‘“‘Typus” [red la- Schnuse, I-1901 (1 2 SMT). bel]; ‘“‘Staatl. Museum fur/ Tierlunde Dres- den.”” Specimen in good condition lacking A the left hind leg and hind right tarsi. Ab- Carvalho, new species domen in microvial with glycerine. (Figs. 3, 6, 9, 12) Other examined material.—Total: 3. BO- Diagnosis.—Souzalopesmyia sulina is LIV1A: Mapiri, Sarapioni, 700-800 m, W. one of the smallest Souzalopesmyia species. Souzalopesmyia sulina 132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON It can be distinguished from the other Souz- alopesmyia species by deeply posterior de- pression on male fifth sternite and 2 frontal setae, latter reclinate. Description.—Male: Head: Frons broad. at narrowest point 0.25 of head width. Eye with only normal pubescence. Fronto-orbit- al plate, parafacial, face and gena silvery white. Fronto-orbital plate broad, broaden- ing gradually from vertex to lunula, at ver- tex plate about equal to diameter of anterior ocellus, at lunula 2.5 times diameter of an- terior ocellus. Frontal vitta broad, parallel to vertex. 2 pairs of strong frontal setae on lower % of frons; former cruciate and latter reclinate; 1 strong orbital reclinate and di- vergent. Ocellar triangle black, reaching in- sertion of orbital setae. Vti strong and par- allel. Postoculars in a complete single row, reaching epistome and composed of black and yellow setulae; latter beginning after basal half of eye and reaching epistome, but 1—2 with black setulae. Gena below lowest eye margin twice diameter of anterior ocel- lus. Palpi slender, yellow. Thorax: Ground-color yellow with scu- tum dusted with whitish-grey, more evident in pre sutural area. Dc: 1:3; 5-6 rows of prst acr setulae: 11—12 rows of post acr se- tulae. | proepisternal seta strong, similar to upper anepisternal setae. Posterior ktps very strong, about 3 times length of anterior one. Scutellum with | strong pair of apical and 1 subbasal setae; | preapical weak but stronger than ground setulae; disc with se- tulae descending below strong setae; bare ventrally. Legs: Yellow. Fore femur with complete rows of AV and D setae; AD row weak. Mid femur with 2—3 PV setae in basal half, less than femoral depth. Mid tibia with 3— 4 P setae. Hind femur with 1 strong preap- ical AV seta, longer than femoral depth; AD row weak and complete; 1 PD, 1 D, 1 AD preapical seta weaker than AV seta. Hind tibia with 4—5 AV setae on apical half. Wings: Clear, veins brownish. Membrane entirely covered with microtrichia, cross- vein r-m placed just before point where vein R1 enters costa. Crossvein dm-cu oblique, almost straight. Calypters and haltere yel- low. Abdomen: Ground-color yellow; in pos- terior view with a narrow black stripe on tergite 1 which is enlarged on tergite 3, 4 and 5. Tergite 4 and 5 with 2 strong apical setae. ferminalia: See Figss376;, 9, 12: Measurements: Length of body, 6.5—8.0 mm (n = 5). Length of wings, 6.5—7.7 mm (n = 5). Female: Unknown. Remarks.—Adults have been collected with Malaise traps. Type material examined.—Holotype ¢ in MHPA labelled as follow; “‘Depto Caninde- yu/Reserva Natural del Bosque/Mbaracayu: Jejui-mi/Malaise 3, bosque bajo inundado/ Colr. A.C.E Costa/ 10-18.VIV1996 [18 July 1996]”; ““Holotipo [holotype]” [red label]. Other material examined.—Total: four paratypes. Same label of the holotype: 2 d, 10-18 July 1996 [DZUP, MHPA], 2 6, 18- 28 July 1996 [DZUP, MHPA]. DISCUSSION Phylogenetic Analysis Phylogenetic studies on Muscidae are still scarce (Carvalho 1989d). The family contains about 200 genera with well over 4,000 species worldwide (Pont 1989, Car- valho et al. 1993). The family is undoubt- edly monophyletic (Hennig 1965, Mc- Alpine 1989, Michelsen 1991), but histori- cally the Muscidae has included groups that are doubtfully monophyletic (Carvalho 1989d). Several genera of Phaoniinae in the Neo- tropics (Pont 1972), a paraphyletic subfam- ily pointed by Hennig (1965) and recently by Carvalho (1989b), were transferred to other subfamilies (Carvalho 1985, 1989a, 1989b, 1989c, Couri and Lopes 1986, Car- valho & Pont 1998). Currently the subfam- ily in the region has only four genera (Car- valho et al. 1993): Dolichophaonia Carva- VOLUME 101, NUMBER 1 S. amazonica S. paraensis, sp. n. 10 Fig: 133 S. singularis S. sulina, sp. n. S. carioca 11 5 l 6,7 17 Cladogram of species of Souzalopesmyia. Synapomorphies = solid black rectangles; reversal = dotted rectangles. Character numbers correspond to those stated in Table 1. lho, 1993; Helina, Phaonia and Souzalo- pesmyia. Souzalopesmyia is a small and isolated clade, apparently representing an ancient lineage of Phaoniinae. The species of genus have head elongate which is commonly cor- related with broad male frons (Vockeroth 1972). But these lengthening is considered independent from that in the some genera in the subfamilies Atherigoninae, Cyrtoneu- rininae, Azeliinae-Reinwardtiini or Myda- einae. The Hennig86 phylogenetic analysis of Souzalopesmyia was based on seven char- acters (with an asterisk in Table 1) and re- sulted in a single tree shown in Fig. 17 (length = 9 steps, consistency index = 0.77, retention index = 0.75). Biogeography Hennig (1965) argued that the first in- vasion (Edentata level) of the Neotropical Muscidae fauna may have come from the Northern Hemisphere during the Upper Cretaceous or Early Tertiary period. This time frame was also suggested by Michel- sen (1991) for the invasion of the basal clade of the Anthomyiidae, Coenosopsia + Phaonantho, into South America. There are few papers on biogeography of Muscidae, none of them of Neotropical Re- gion, except Hennig (1965). In Holarctic Region, all taxa of the Eudasyphora s. str. Townsend (Muscinae) are faunal elements of known dispersal centers (Cuny 1980). The speciation of these flies, were correlat- 134 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON c) 9 Caribbean Plate a Vicariance of Northwestern Amazonia Separation of the Northwestern and the Southeastern Components Serra da Mantiqueira Fig. 18. A summary of the main vicariance barriers in the history of the Neotropical region. 1 = Carribbean Plate detachment from the mainland. 2 = Separation between the Northwestern and the Southeastern main components, a line along the Rivers Amazonas/Madeira/Mamoré in the Amazonian Basin. 3 = Epicontinental sea formation in the Maracaibo area. 4 = A large division in northwestern Amazonia (not related to date to any geological event). 5 = Middle to Late Cretaceous water connection between the Parnaiba Basin and the Parana Basin (redrawn from Amorim and Pires 1996). ed with the history of the forest vegetation during the Pleistocene (Cuny 1980). Amorim and Pires (1996), corroborated independently by Grazia (1997), indicated that the first division in the continental re- gion of the Neotropics was in the Late Cre- taceous, showing a northwestern track against a southeastern track (Fig. 18). Alike pattern is showed by Camargo (1996) for some bees (Meliponini, Apinae, Apidae) in Neotropical Region. However, this latter biogeographical reconstruction was postu- lated by modification occurred by the changing forests in the Pleistocene. Souzalopesmyia, based on the position of S. amazonica as the basal clade of genus, may have had its ancestor back in the Late Cretaceous (Fig. 19). This age for the genus is not unrealistic although no fossil record is known for family older than the Eocene (Evenhuis 1994). Pont and Carvalho (1997) studied three fossil species of Muscidae VOLUME 101, NUMBER 1 Fig. 19. Geographical ynica. b, S. paraensis. C, distribution of s c, S. singularis. d. pecie SoSH NS) l of Souzalopesmyia in a. e, S. carioca 136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from Dominican amber dated from Mio- cene, 15-20 mya ago, two of which are Phaonia species. A cladogram by S. M. P. Coelho (unpublished Ph.D. thesis) suggests that the Phaonia fossil species have a more recent origin than the ancestor of Souzalo- pesmyia. The origin of the genus is there- fore probably older than 15—20 mya, sug- gesting that Souzalopesmyia is one of the most basal genera of Phaoniinae in the Neo- tropics. The origin of the Neotropical Phaoniinae fauna cannot be completely understood on the basis of the present paper. The ancestor of Souzalopesmyia may have reached South America by dispersal from North America (Hennig 1965), Africa or have evolved in the Neotropics. The discovery of the sister group of the genus is required. The five species of Souzalopesmyia are morphologically similar and exhibit allo- patric distribution suggesting that the ter- minal branches of this clade could be re- sulted from relatively recent speciation. Nevertheless, the allopatric pattern of speciation of the genus cannot be fully ex- plained with the available geological and biogeographical information. Most of the species are known from only a few speci- mens, suggesting that intensive effort may be necessary before reliable statements can be made about the distribution patterns of the species. Species of Souzalopesmyia are not expect to occur in western side of An- des as it requires tropical rainforest (Figs. 18, 19). The occurrence of S. paraensis in Belém, along south side of the Amazon River, part of the northwestern track, could be the re- sult of a single dispersal event (Figs. 18, 19) to colonize that region, in a more recent time, which belongs to the northwestern track. ACKNOWLEDGMENTS I express my sincere gratitude to the fol- lowing persons who supplied specimens for this study: Uwe Kallweit (SMT), D.M. Pamplona (MNRJ), T. Pimentel (MPEG) and John Kochalka (MNPA). I also thank A. C. FE da Costa, The Fundacion Moises Bertoni and the Darwin Initiative for logis- tic support for the project Biodiversity In- ventory of the Mbaracayu Forest Nature Reserve, Paraguay. I thank the following for critically reviewing the manuscript: L. Marinoni, Universidade Federal do Parana, Curitiba, Brazil; D. S. Amorim, Universi- dade Sao Paulo, Ribeirao Preto, Brazil; M. S. Couri and D. M. Pamplona. Museu Na- cional do Rio de Janeiro, Rio de Janeiro, Brazil; G. S. Carvalho, Pontificia Univer- sidade Catolica, Porto Alegre, Brazil; A. Y. Harada, Museu Paraense Emilio Goeldi, Belém, Brazil; A. C. Pont, University Mu- seum of Natural History, Oxford, U. K; R. Holzenthal, University of Minnesota, U.S.A. I am deeply grateful to R. Cuny, Lakeland College, Alberta, Canada, for comments and additions, which have mark- edly improved the paper. This study was partially supported from CNPq, Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (300.043/86-4). This is paper number 1035 of the Departamento de Zoo- logia, UFPR. LITERATURE CITED Albuquerque, D. de O. 1951. S6bre um género e duas espécies novas de “‘Phaoniinae” neotropical (Di- ptera, Muscidae). 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Revision of the genus Eudasyphora Townsend (Diptera: Muscidae), and reflections on its evolution. The Canadian Entomologist 112: 345-373. Evenhuis, N. L. 1994. Catalogue of the fossil flies of the world (Insecta: Diptera). Backhuys, 600 pp., Leiden. Farris, J. S. 1988. Hennig86 reference. Version 1.5. Published by the author, New York. Grazia, J. 1997. Cladistic analysis of the Evoplitus ge- nus group of Pentatomini (Heteroptera: Pentatom- idae). Journal of Comparative Biology 2: 43—47. Hennig, W. 1965. Vorarbeiten zu einem phylogene- tischen System der Muscidae (Diptera: Cyclor- rhapha). Stuttgarter Beitraege zur Naturkunde 141: 100 pp. Huckett, H. C. and J. R. Vockeroth. 1987. Muscidae, pp. 1115-1131. Jn McAlpine, J. E, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth and D. M.Wood, eds., Manual of Nearctic Diptera, 2. Research Branch Agriculture. Canada Monograph 28: 675-1332. Lopes, S. M., M. Couri, D. Pamplona, and C. J. B. de Carvalho. 1997. Notes on Neotropical types of Diptera described by Albuquerque (Anthomyidae [sic], Fanniidae, Muscidae, Piophilidae, Psilidae, Sapromyzidae, Scatophagidae e Stratiomyidae). 137 Publicagoes Avulsas do Museu Nacional, Rio de Janeiro, n. 69: 1-33. Malloch, J. R. 1929. Exotic Muscaridae (Diptera).— XXVI. Annals and Magazine of Natural History (10) 4: 97-120. McAlpine, J.E 1981. Morphology and terminology, pp. 9-63. In McAlpine, J. E, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth and D. M.Wood, eds., Manual of Nearctic Diptera, 1. Re- search Branch Agriculture. Canada Monograph 27: 1-674. Michelsen, V. 1991. Revision of the aberrant New World genus Coenosopsia (Diptera: Anthomyi- idae), with a discussion of anthomyiid relation- ships. Systematic Entomology 16: 85-104. O’Hara, J. E. 1982. Classification, phylogeny and zoo- geography of the north American species of Si- phona Meigen (Diptera: Tachinidae). Quaestiones Entomology 18: 261—380. Pont, A. C. 1972. Family Muscidae. Jn A Catalogue of the Diptera of the Americas South of the United States, 97. Museu de Zoologia, Universidade de Sao Paulo. 111 pp. 1980. Family Muscidae, pp. 721-761. In Crosskey, R. W., ed., Catalogue of the Diptera of the Afrotropical Region. British Museum (Natural History). London. 1437 pp. . 1986. Studies on Australian Muscidae (Dip- tera) VII. The genus Atherigona Rondani. Austra- lian Journal of Zoology, Supp. Ser., 120: 1-90. . 1989. Family Muscidae, pp. 675-699. In Ev- enhuis, N., ed., Catalog of the Diptera of the Aus- tralasian and Oceanian Regions. Bishop Museum Press, Honolulu & E.J. Brill, Leiden. 1155 pp. Pont, A. C. and C. J. B de Carvalho. 1997. Three spe- cies of Muscidae (Diptera) from Dominican am- ber. Studia Dipterologica 4: 173-181. Séguy, E. 1937. Diptera Family Muscidae. /n Wyts- man, P., Genera Insectorum. Fasc. 205: 604 pp., Bruxelles. Stein, P. 1911. Die von Schnuse in Siidamerika gefan- genen Anthomyiden. Archiv fiir Naturgeschichte 77 (1): 61-189. . 1919. Die Anthomyidengattungen der Welt, analytisch bearbeitet, nebst einem kritisch-syste- matischen Verzeichnis aller aussereuropaische Ar- ten. Archiv fiir Naturgeschichte 83 A 1 [1917]: 85-178. Vockeroth, J. R. 1972. A Review of the World Genera of Mydaeinae, with a Revision of the Species of New Guinea and Oceania (Diptera: Muscidae). Pacific Insects Monograph 29: 134 pp. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 138-142 ASHLOCKOBIUS, A NEW GENUS OF MYODOCHINI FROM VENEZUELA (HEMIPTERA: LYGAEOIDEA: RHYPAROCHROMIDAE: MYODOCHINI) JAMES A. SLATER AND ALEX SLATER (JAS) Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT 06269, U.S.A.; (AS) 901 Alabama Street, Lawrence, KS 66044, U.S.A. Abstract.—Ashlockobius cursorius, new genus and species of myodochine lygaeoid is described from Venezuela. The genus is related cladistically to Orthaea Dallas, Catenes Distant, Heraeus Stal and Myodocha Latreille. It is believed to be an ant mimic. Figures are given of the details of the inflated phallus and of the claspers. A color figure of the habitus is included. Key Words: lockobius cursorius, mimicry The Myodochini constitute one of the dominant elements in the lygaeoid fauna of the Neotropics. It is abundant and diverse not only in the number and variety of taxa present but in the abundance of many in- dividuals of some species. Ant mimicry oc- curs frequently. The new genus described below is apparently an ant mimic although its biology is unknown. Henry (1997) has elevated the former subfamily Rhyparochrominae of the family Lygaeidae to family status with two sub- families, the Plinthisinae and the Rhyparo- chrominae with the former tribes of the lat- ter retained as tribes. We have adopted Hen- ry’s conclusions here, but suggest that the tribes within his new definition of the sub- family may ultimately prove to merit family status cladistically. Henry’s statement that the subfamily Rhyparochrominae as rede- fined consists of taxa with an incomplete suture between abdominal sterna four and five and with carinate pronotal margins is an oversimplification. Genera that occur in at least five tribes within the Rhyparo- chrominae have the abdominal sternal su- ture complete. Rounded, ecarinate pronota Hemiptera, Lygaeoidea, Rhyparochromidae, Myodochini. Venezuela, Ash- occur in several tribes and is one of the di- agnostic features of the Myodochini. All measurements are in millimeters. Ashlockobius Slater and Slater, new genus Description.—Body elongate, slender, nearly parallel sided. Dorsal surface of body subshining. Legs and antennae both extremely elongate. Head moderately decli- vent anteriorly; eyes set a short distance away from anterior pronotal angles. Anten- niferous tubercules divergent. Head below broadly transversely striate from level of distal ends of antenniferous tubercles to level of posterior margins of eyes. Bucculae V-shaped. Vertex convex between eyes with patches of pruinosity present. Anterior pronotal lobe extremely elon- gate and elliptically convex, with a pruinose median stripe on anterior pronotal lobe and large irregular pruinose patches laterally on posterior half of anterior lobe. Posterior pronotal lobe dull with a patch of white pruinosity near transverse impression on ei- ther side of midline. Pronotum with only scattered, relatively inconspicuous, punc- VOLUME 101, NUMBER 1 tures present; anterior collar complete, coarsely punctate, delimited posteriorly by a sharp deeply impressed line, not produced posteriorly at meson; transverse pronotal impression deep and complete; posterior pronotal lobe with prominent punctures. Scutellum with a conspicuous Y-shaped, elevated carina; pruinose, bicolored with a large triangular basal grayish-white patch and a spot on each side near divergence of elevated Y-carina, remainder of scutellum reddish brown. Clavus, corium and mem- brane contrastingly strongly shining former with 3 complete rows of punctures and a partial fourth row on distal half between in- ner and median rows. Corium moderately expanded posteriorly; outer margin irregu- lar and beaded, lacking a stridulitrum. Api- cal corial margin adjacent to membrane lacking a series of punctures. Mesepimeron not emergent. Metathoracic scent gland au- ricle short, straight, tapering distally; evap- orative area large, covering inner two thirds to three fourths of metapleuron, and ex- tending narrowly along posterior margin of mesopleuron. Fore coxa with a prominent spine and a smaller secondary spine present. Fore fe- mur elongate and slender, almost entire ventral surface of each femur heavily spi- nous, with rows of large spines along inner and outer edges and numerous small spines between. Each fore tibia with a widely spaced series of 4 short sharp spines on in- ner face. Shaft of tibia not strongly curved. Lateral and ventral surfaces completely dull and chiefly pruinose except for a large quadrate shining patch mesally on meso- sternum, this latter with a narrow median groove. Antenna very elongate, slender, te- rete with fourth segment fusiform. Posterior margin of abdominal sternum 2 not finely scalloped. Posterior margin of pygophore broadly rounded, without a median impression. Clasper (Fig. 3) with a distinct thumb-like exterior projection, interior margin with a pronounced flange narrowing distally but extending almost to apex. Phallus (Figs. 2, 139 4) without sclerotized conjunctival or vesi- cal spines; conjunctiva short, bearing low lateral subapical lobes; vesica short; heli- coid process present; a large lobe bearing many projections on each side of ejacula- tory reservoir. Ejaculatory reservoir with well developed wings (Fig. 2), holding sclerites short, narrowing distally. Type species.—Ashlockobius cursorius, new species. Monotypic. Discussion.—Ashlockobius keys without difficulty to couplet 51 in Harrington’s (1980) key to myodochine genera. It differs from Jogo Bergroth (from Japan) in having a relatively much longer anterior pronotal lobe, more than one and one-half times the length of the posterior lobe in Ashlocko- bius; in Togo the anterior lobe is at most only slightly longer than the posterior lobe. Ashlockobius also does not have strongly curved anterior tibiae. Most specimens of Togo are brachypterous, the male fore tibiae are prominently curved, and the members of the genus are relatively stout and robust. Ashlockobius differs from those genera reached through couplet 52 of Harrington by having the combination of armed male fore tibiae, as described above, and an an- terior pronotal lobe at least 1.5 times as long as the posterior lobe. The absence of vesical or conjunctival spines and the robustly winged ejaculatory reservoir with short holding sclerites places Ashlockobius in the group of genera with Harrington’s Type IV phallus. This place- ment is supported by the broadly rounded posterior margin of the pygophore (Fig. 5). Within this group the presence of pruinose areas on the dorsum and the somewhat elongate head and rounded vertex place the genus with a group on Harrington’s clado- gram consisting of Orthaea Dallas, Catenes Distant, Heraeus Stal, and Myodocha La- treille. Analysis of other phallic characters must await study of the inflated phallus of more species. The illustration of an inflated Myodocha phallus provided by Ashlock (1957), which exhibits an elongate conjunc- tiva with several lobes and an apparently 140 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Ashlockobius cursorius, Dorsal view. VOLUME 101, NUMBER 1 141 Figs. 2-5. Ashlockobius cursorius. 2, Ejaculatory reservoir, dorsal view. 3, Claspers, outer and inner views. 4, Aedeagus, lateral view. 5, Genital capsule, dorsal view. asymmetric set of vesical lobes, indicates Ashlockobius cursorius Slater and that such studies will prove to be fruitful. Slater, new species Etymology.—It gives us great pleasure to (Figs. 1—5) be able to dedicate this striking new genus Description.—Male: Body very elon- to the memory of Dr. Peter D. Ashlock for gate, slender, attenuated, with extremely his many contributions to the systematics of elongate appendages. Color bright reddish the Lygaeoidea. brown almost throughout, including ap- 142 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pendages. Antenna with distal ends of seg- ments II and III infuscated with chocolate brown. Corium marked with chocolate brown as follows: a small spot along lateral corial margin at level of distal third of api- cal corial margin, a small apical spot, an elongate dash along radial vein running from level of distal fifth of claval commis- sure nearly to anterior end of apical corial margin, a stripe at inner angle of corium that extends from inner margin at apical third of claval commissure to adjacent row of punctures then narrowly along apical corial margin to a level slightly posterior to lateral corial spot. Anterior collar and pos- terior pronotal lobe yellow, contrasting with red-brown anterior lobe. Corium light yel- low brown except as noted above. Eyes large, protrudant but not stalked, head strongly narrowing behind eyes but without a distinct stalked neck. Length head 1.34, width 1.20, interocular space 0.62. Length pronotum 2.60, length anterior pronotal lobe (less anterior collar) 1.60, width 1.84. Length scutellum 1.24, width 0.96. Length claval commissure 0.84. Mid- line distance apex clavus-apex corium 1.76. Midline distance apex corium-apex abdo- men 1.72. Length labial segments I 0.76, I 0.84, Il 0.52, IV 0.44. Labium reaching posterior third of prosternum but remote from fore coxae. Length antennal segments I 1.88, If 3.48, If 2.80, [IV 1.72. Total body length 9.80. Holotype.—d. VENEZUELA: Aragua, 16 km S. Telerias, May 10, 1978 (C.W. & L.B. O’Brien and Marshall). In American Museum of Natural History, New York. Etymology.—The name is from the Latin word for a runner. Discussion.—The male specimen upon which the preceding description is based was collected in Venezuela twenty years ago. It is a striking, elongate, long-legged myodochine, which we have held for many years in the hope that additional specimens would become available. Unfortunately this has not happened. ACKNOWLEDGMENTS We thank Drs. Charles and Lois O’Brien (Florida State University, Tallahasee) for the gift of the holotype and Mrs. Mary Jane Spring (University of Connecticut) for ex- ecution of the dorsal view illustration. LITERATURE CITED Ashlock, P. D. 1957. An Investigation of the Taxonom- ic value of the Phallus in the Lygaeidae (Hemip- tera-Heteroptera). Annals of the Entomological Society of America 50: 407—426. Harrington, B. J. 1980. A generic level revision and cladistic analysis of the Myodochini of the World (Hemiptera, Lygaeidae, Rhyparochrominae). Bul- letin of the American Museum of Natural History 167: 49-116. Henry, T. J. 1997. Phylogenetic Analysis of Family Groups within the Infraorder Pentatomomorpha (Hemiptera: Heteroptera), with Emphasis on the Lygaeoidea. Annals of the Entomological Society of America 90: 275-301. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 143-148 BIOSYSTEMATIC STUDIES OF CEYLONESE WASPS, XXII: BETHSMYRMILLA, A NEW GENUS OF MUTILLID WASPS (HYMENOPTERA: MUTILLIDAE: MYRMILLINAE) KARL V. KROMBEIN AND ARKADY S. LELEJ (KVK) Department of Entomology, MRC 165, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0165, U.S.A.; (ASL) Institute of Biology and Pedology, Vladivostok-22, 690022, Russia (e-mail: entomol @online.marine.su) Abstract.—Bethsmyrmilla alticola, new genus and new species is described from Sri Lanka. A key is given to the Oriental genera of Myrmillinae. The halictine bee Lasio- glossum (Sudila) alphenum (Cameron) is the probable host of B. alticola. Key Words: Mutillid wasps, Bethsmyrmilla alticola, new genus and new species, Sri Lanka, Lasioglossum (Sudila) alphenum (Cameron) Borge Petersen estimated that the mutil- lid fauna of Sri Lanka was 74 species based on the literature (in litt. to KVK, 1976). He mentioned that he found 45-50 species among the specimens that he had borrowed. The latter figure, however, does not incor- porate a study of all of the rich mutillid fauna collected during the later years, 1975-1981, of the Smithsonian’s “‘“Ceylon Insect Project’? and two visits in 1993 and 1997 by K. V. Krombein and B. B. Norden. We anticipate that a study of these collec- tions will result in a much more accurate inventory of the mutillid fauna. During the latest trip to Sri Lanka, Beth Norden found two females of an unusual small mutillid wasp in the ground nests of the halictine bee Lasioglossum (Sudila) al- phenum (Cameron). These wasps belong to a new genus and species that we place in the Myrmillinae because the side of the tho- rax is evenly concave, the mesopleuron has a strong supracoxal carina and the pronotal- mesopleural suture is lacking except a small section above. The subfamily presently comprises three genera in the Oriental re- gion: Spilomutilla Ashmead, Squamulotilla sensu Mickel and Bethsmyrmilla, new ge- nus. The genus Squamulotilla Bischoff (type species Squamulotilla denticollis Bischoff, male, North Cameroon, Nigeria) includes seven Afrotropical species which are known from males only. For many years Guido Nonveiller has collected and studied the mutillid fauna of Cameroon. We support his opinion (1995) that true females of Squamulotilla are probably described in the Afrotropical genera Clinotilla Arnold (males still unknown). Oriental species placed in Squamulotilla by Mickel (1933, 1935) comprise several undescribed genera of Myrmillinae which will be treated sep- arately. Depositories for specimens listed are as follows: IBPV _ Institute of Biology and Pedology, Vladivostok, Russia. USNM National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. U.S.A. 144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON KEY TO THE ORIENTAL GENERA OF SUBFAMILY MYRMILLINAE Ia yRemaley ecg sce tes yo see Ie ees Oy DD Male (unknown for Bethsmyrmilla) . Mandible widened apically, without subbasal tooth on inner margin (Fig. 8); gena dentate beneath; gastral tergum 2 posteriorly with three spots (Fig. 3) or a wide band of pale pubes- cence shallowly concave medially; gastral ster- num 2 with median longitudinal carina ending usually invanvacutestubercle, say «ea 2 ee Sri MeRa th cat) Geo eC Tn ee te Spilomutilla Ashmead — Mandible not widened apically, with subbasal tooth on inner margin (Figs. 6, 7, 9); gena not dentate beneath; gastral tergum 2 posteriorly with a band of pale pubescence that widens medially to an obtuse angle (Figs. 1, 2, 5) or with a large median spot (Fig. 4); gastral ster- num 2 if carinate not ending in a tubercle... 3 3. Mid trochanter with narrow apical process (Fig. 12); posterolateral angles of head tuber- culate behind eyes (Figs. 14, 15); thorax gently sloping posteriorly, upper margin not dentate nor serrate (Figs. 15, 16); mandible apically with a single tooth (Fig. 6) .. Bethsmyrmilla Krombein and Lele}, new genus — Mid trochanter without process; posterolateral angle of head not tuberculate but gena tuber- culate below eye in Squamulotilla lamellata Mickel and S. arundinacea Pagden; thorax abruptly sloping posterorly, upper margin usu- ally serrate or dentate, rarely with only an acute median denticle, or if not dentate at all (S. puerilis Cameron and S. afghanica Lele}) inner mandibular margin with two small preapical denticles (Fig. 9) and gastral tergum 2 with large spot of pale pubescence on posterior mar- gin (Fig. 4) Oye dart Papers. SeaNNuateld aeeee Squamulotilla sensu Mickel 4. Apterous; mandible extremely widened apical- ly, with deep preapical emargination; hind coxa ventrally with small, sharp posterolateral den- ticle; gastral sternum 8 membranous laterally, narrow median part with parallel lateral ridges se ES ys TEL TO RR pte Rp pe Spilomutilla Ashmead — Alate; mandible not widened apically, inner border with two small preapical denticles and larger basal one (cf Fig. 9); hind coxa ventrally without denticles; gastral sternum 8 not mem- branous laterally, with median carina or tuber- cle; gastral terga 2—5 very often with median longitudinal’ carina? As): 443 - serene ec aac Nucl yoni cast wane i Squamulotilla sensu Mickel we Bethsmyrmilla Krombein and Lelej, new genus (Figs. 1, 6, 10-18) Type species.—Bethsmyrmilla alticola Krombein and Lelej, new species. The ge- nus is monotypic. Female.—Head large, distinctly wider than thorax, viewed from above the sides gradually widened behind eyes (Fig. 14), widened area forming a sharp tubercle as viewed from behind (Fig. 15); mandible not widened apically, inner margin with sub- basal tooth (Fig. 6); medial clypeal lobe with prominent lateral tooth (Fig. 10); fla- gellomere [I 1.8—1.9X its maximal width and 1.8—2.0 as long as flagellomere II (Fig. 11), the latter wider than its length. Mesosoma dorsally more or less quad- rangular, laterally crenulated (Fig. 15); me- sopleuron with trituberculate supracoxal ca- rina (Fig. 16, sc); propodeal dorsum pos- teriorly without denticles, posterior face of propodeum sloping gently downward; fore tarsi with weak comb (Fig. 18); mid coxa ventrally with blunt tubercle and mid tro- chanter with narrow apical process (Fig. I) Dorsum of gaster with spots and bands of golden pubescence (Fig. 1); basal half of gastral sternum 2 with weak median carina that does not terminate in a tubercle. Male.—Unknown. Discussion.—Differences of this new ge- nus from other Oriental Myrmillinae are noted in the key. The female of Bethsmyr- milla is similar to those of Squamulotilla lamellata Mickel and S. arundinacea Pag- den in having a lateral tubercle on the head, similarly shaped mandibles (cf Figs. 6, 7) and similar spots and bands of golden pu- bescence (cf Figs. 1, 2) on the gaster. It is easily separated from the latter species in lacking a tubercle on the fore coxa, in hav- ing a narrow apical process on the mid tro- chanter and the position of the lateral tu- bercle on the head, behind the eye in Beths- myrmilla and beneath the eye in the latter two species. Also, the propodeal dorsum in the former is not denticulate posteriorly whereas the latter species have at least an acute median denticle. The female of Bethsmyrmilla is superfi- cially similar to another larger group of Squamulotilla species in having spots and bands of golden pubescence on some of the VOLUME 101, NUMBER | 145 \ 1.0 mm Figs. 1-12. Females of Oriental Myrmillinae. 1-5, Patterns of pale pubescence on abdominal terga. 1, Bethsmyrmilla alticola holotype, Sri Lanka, terga 1-3 only. 2, Sqguamulotilla lamellata, Vietnam. 3, Spilomutilla consolidata (Cameron), Sri Lanka. 4, Sq. puerilis, South India. 5, Sg. strangulata (Smith), South China. 6-9, Mandible. 6, B. alticola. 7, Sq. lamellata. 8, S. consolidata. 9, Sq. puerilis. \O—12, B. alticola. 10, Clypeus, frontal view. 11, Antennal segments 1-5. 12, Coxa, trochanter, base of femur, mid leg above, hind leg below. 146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON + o Dg 231m ne Sa) tn’ 7 ¥ roy Bere oe. Figs. 13-18. Bethsmyrmilla alticola, female paratype uncoated. 13, 14, Head. 13, Frontal view (note genal tubercle extending laterad of each eye on lower margin). 14, Dorsal view. 15, 16, Thorax. 15, Dorsal view. 16, Oblique view, bracket and sc indicate supracoxal carina. 17, Head, posterolateral view, note sparse, long setae that margin eye. 18, Foretarsus, note that basal segment is angled, foreshortened. VOLUME 101, NUMBER 1 gastral terga but differs markedly in having the lateral tubercle on the head, in lacking denticles along the posterior margin of the thoracic dorsum and in the gently rather than abruptly sloping posterior surface of the thorax. Etymology.—We take great pleasure in naming this remarkable mutillid for Beth B. Norden, collector of the type series of B. alticola and discoverer of its probable host species. Bethsmyrmilla alticola Krombein and Lelej, new species (Figs. 1, 6, 10—18) Female.—Length (through gastral seg- ment 3) 5.0 mm. Red; gaster black except sternum | totally and tergum | basally red; mandible red, brownish apically; palpi brownish; scape red, flagellomeres red, darkened above; legs red, mid and hind tib- iae with darker spines; mandible and clyp- eus with long pale erect setae, scape with shorter ones; frons and vertex with sparse recumbent, short reddish setae, genae with whitish ones; vertex and frons along eye or- bit with sparse, long, erect black setae (Fig. 17); thoracic dorsum with sparse, short, subappressed black setae mixed with long, erect black ones; legs with subappressed and erect yellowish setae; pattern of golden pubescence on gaster (Fig. 1); gastral sterna 2 and 3 posteriorly with whitish fascia; pro- podeal hindface, gastral tergum 1 anterior- ly, gastral sternum 2 and gastral terga 2 and 3 laterally with sparse, erect, whitish setae. Head parallel behind eyes, gena postero- laterally with strong acute tubercle (Figs. 13-17); antennal scrobe well developed be- tween antennal tubercle and lower part of eye (Fig. 13); clypeus delimited above by a weak convex carina with four small tuber- cles and ending laterally in a strong tooth (Figs. 10, 13); antenna with short flagello- meres (Fig. 11), flagellomere I 1.8—1.9X its apical width and 1.8—2.0 flagellomere II, the latter 0.75 its width; hypostomal ca- rina without projection; gena below not car- inate; head above with dense large punc- 147 tures (Figs. 13, 14, 17). Thorax viewed from above (Fig. 15) more or less quadran- gular with slightly widened pronotal and propodeal areas; humeral angles developed, lateral margin of pronotum with vertical ridge (Fig. 16); thoracic dorsum noticeably convex, weakly serrate laterally, with one lateral tubercle on the middle of mesono- tum, and with dense reticulate punctures; propodeal dorsum posteriorly not dentate nor serrate; mesopleuron concave with su- pracoxal carina well developed, the latter with three blunt tubercles (Fig. 16, sc); mid coxa ventrally with blunt tubercle and mid trochanter with narrow apical process (Fig. 12); hind coxa carinate along posterior mar- gin of ventral surface, carina ending in a blunt tubercle. Gastral sternum 1 with well developed median carina; gastral tergum 2 with rather short lateral felt line and dense small punc- tures; gastral sternum 2 with more or less flattened central disc and dense punctures which are much sparser and larger on disc; gastral segments 4—6 lacking in holotype and 3—6 in paratype (these parts acciden- tally amputated and lost during nest dig- ging). Male.—Unknown. Range.—The species is known only from a trail at about 1,950 m altitude along the upper border of the Hakgala Botanic Gar- den about 10 km S of Nuwara Eliya. We believe that it may be more widely distrib- uted. Its probable host has been collected only at high altitudes, 1,700—1,950 m, at various localities in the districts of Nuwara Eliya (Nuwara Eliya, Hakgala, Horton Plains) and Kandy (Adam’s Peak Trail). Type material.—Holotype 2, Sri Lanka, Nuwara Eliya District, Hakgala Botanical Garden, 6°55’N, 80°49’E, 21-22 April 1997, B.B. Norden [USNM]. Paratype °&, same data as holotype but 24—26 February 1997 [IBPV]. Etymology.—The specific name is from the Latin altus, high, and -cola, dweller. Natural history.—Beth Norden excavated two nests of Lasioglossum (Sudila) alphe- 148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON num one each on 26 February and 22 April 1997. Each nest contained a single female mutillid. Both wasps were found at the ba- ses of the nest tunnels and are presumed to have traveled to the farthest reaches of the vertical shafts in an effort to avoid capture. The February nest shaft reached a depth of 7 cm and also contained a female L. al- phenum and a female of the parasitic bee Nomada priscilla Nurse that were captured a few mm above the mutillid. The April nest shaft reached a depth of 13 cm. In this nest, two female L. alphenum were collect- ed at depths of 12 and 12.5 cm respectively. The mutillids moved more vigorously than the bees and were more difficult to col- lect. Pollen and immature bees (larvae and pupae in various stages of development) were found in cells located above the base of the nest where the wasps were retrieved. Discussion.—Norden et al. (1994) found two females of the mutillid wasp Pseudo- methoca bethae Krombein within a com- munal nest of an Arizona bee Exomalopsis (Phanamalopsis) solani Cockerell. One mutillid was within a provisioned cell pre- sumably to feed on the pollen-nectar mass. Although neither specimen of B. alticola was in a cell with provisions, we suspect that they would use this convenient source of food while they remained within the nest. And when bee brood reached the ap- propriate developmental stage it would be available for parasitism. Such cryptic be- havior by the mutillid would greatly lessen exposure to predators and may explain the rarity of this species in collections. ACKNOWLEDGMENTS Within the Smithsonian Institution we thank Beth B. Norden for her notes on the natural history of B. alticola and preparing specimens for SEM study, Susann G. Bra- den for skillful preparation of the scanning electron micrographs and George L. Vena- ble for preparation of the plates. We are grateful to P- Klimov (IBPV) for preparing Figures 1-12. ASL thanks the late Borge Petersen for generously providing valuable exchange material of Oriental mutillid wasps and V. Kuznetsov (IBPV) for the gift of material. KVK thanks Bryan N. Dan- forth, Cornell University, Ithaca, NY, for identification of Lasioglossum (Sudila) al- phenum and Maximilian Schwarz, Ansfel- den, Austria, for identification of Nomada priscilla. We are grateful to G. Nonveiller, Zemun, Yugoslavia, for critical reading of the manuscript and valuable comments, and also an anonymous reviewer. LITERATURE CITED Mickel, C. E. 1933. The Mutillidae of Eastern Asia. Lingnan Science Journal 12(3): 289-325. . 1935. The Mutillid Wasps of the Islands of the Pacific Ocean (Hymenoptera: Mutillidae). Transactions of the Royal Entomological Society of London 83(2): 177-312. Nonveiller, G. 1995. Recherches sur les Mutillides de Afrique XVII. Note pour servir a la connais- sance du Pristomutilla Ashmead, 1903 (male, femelle) avec description du male encore genre inconnu du genre, d’espece nouvelles du genre et des nouveaux sous-genres Diacanthotilla (femel- les) et Acanthomutilla (femelles) (Hymenoptera, Mutillidae). Entomofauna 16(5): 29-119. Norden, B. B., K. V. Krombein, and S. W. T. Batra. 1994. Nests and Enemies of Exomalopsis (Phan- amalopsis) solani Cockerell (Hymenoptera: Apo- idea, Mutillidae; Diptera: Asilidae). Proceedings of the Entomological Society of Washington 96(2): 350-356. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 149-163 TWO NEW WESTERN NEARCTIC CULICOIDES LATREILLE (DIPTERA: CERATOPOGONIDAE) DESCRIBED FROM ALL STAGES MaArK S. BREIDENBAUGH AND BRADLEY A. MULLENS Department of Entomology, University of California, Riverside, California 92521, U.S.A. (BAM, corresponding author for reprints) Abstract.—The egg, larva, pupa, and adult of two new species of Culicoides La- treille from desert mountains in southern California and Baja, Mexico are described: C. kettlei and C. vetustus. Descriptions of immatures were made from laboratory- reared material. Rearing techniques and notes on behavior observed in the laboratory are presented. Key Words: Bluetongue and epizootic hemorrhagic disease viruses have been suspected of con- tributing to lamb mortality in desert bighorn sheep populations in southern California (DeForge et al. 1982, Wehausen et al. 1987, Elliott et al. 1994). A comprehensive sur- vey of potential vectors (Culicoides Latre- ille) occurring in bighorn habitats within the desert mountains of southern California (Mullens and Dada 1992a) revealed three undescribed species of Culicoides. The adults of one species, Culicoides boydi Wirth and Mullens have been previously described (Wirth and Mullens 1992). The present study describes all life-stages of these remaining two species and provides data on their biology and geographic distri- bution. MATERIALS AND METHODS Host-seeking females were collected us- ing suction traps baited with | kg of dry ice. Some females collected at the Philip L. Boyd Deep Canyon Desert Research Center near Palm Desert, Riverside Co., CA, or near the town of Morongo Valley, San Ber- nardino Co., were given the opportunity to feed on heated, defibrinated sheep blood Culicoides, Ceratopogonidae, immatures, morphology through a parafilm or chick-skin membrane (Hunt 1994). Engorged females were sep- arated from other midges in the collections while they were immobilized on a chill ta- ble. Female midges were then held at 21°C for 7 days. Some individual females would oviposit onto damp filter paper, when held overnight in a petri dish. Gravid females which did not oviposit were decapitated to induce oviposition. Filter paper with eggs from an individual female was placed in a petri dish containing nutrient-enriched 1.5% noble agar. Eggs were held in a humid chamber and checked daily for hatch. First-instar larvae were of- fered several food sources, including a nu- trient rich liquid diet consisting of bacteria, algae, and yeast used for rearing colonized Culicoides variipennis sonorensis Wirth and Jones (Jones et al. 1969). In addition, the bacterial feeding nematodes, Pelodera sp. and Panagrellus redivivus (L.), were supplied as potential prey on a biweekly ba- sis (Mullens and Velten 1994). Larval growth and feeding behavior were observed daily and the duration of egg, larval, and pupal development recorded. Samples of eggs from associated females 150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON were placed into 70% EtOH. Specimens were fixed (Day et al. 1997), then critical- point dried, transferred onto stubs backed with sticky tape, sputter-coated with gold- palladium, and viewed on either a JOEL JSM-35C or Phillips XL30 scanning elec- tron microscope. The parental adult female was preserved in 70% EtOH and slide-mounted in balsam after Wirth and Marston (1968). Fourth-in- star larvae, pupae, and adults from reared offspring of the parental female were also preserved. Offspring were mounted in Can- ada balsam or Hoyer’s medium. Descrip- tions of immature stages and males of both species were made from laboratory-reared material. The terminology of Downes and Wirth (1981) is used for adults, of Lamberson et al. (1992) and Nevill and Dyce (1994) for pupae, of Murphree and Mullen (1991) for larvae, and of Becker (1961) and Campbell and Kettle (1975) for eggs. Nomenclature agrees with Borkent and Wirth (1997) and Spinelli and Ronderos (1997). The following measurements were made from fourth-instar larvae: total length (TL), head length (HL), head width (HW), sub- genal width (SGW), mandible length (ML), width across the lateral arms of the epi- pharynx (LAW), total width across the paired dorsal comb sclerites of the epiphar- ynx (DCW), caudal-segment length (CSL), caudal-segment width (CSW), length of se- tae ‘o’ (OL), and the distance between their bases. From values listed above, the following ratios were calculated: head ratio (HR = HL/HW), subgenal ratio (SGR = HW/ SGW), and caudal-segment ratio (CSR = CSL/CSW). Illustrations were made of the morphology and chaetotaxy of the head capsule and caudal segment. The hyposto- ma, epipharynx, hypopharynx, and mandi- ble were illustrated, and thoracic pigmen- tation and anal papillae were drawn when appropriate. For pupae the following structures were described and illustrated: respiratory horn, operculum, caudal segment, and the ad, dl, dasm, dpm, lasm, lpm, and vpm tubercles. The holotype, allotype, and some para- types are deposited in the National Museum of Natural History (USNM), Smithsonian Institution, Washington, DC; paratypes, as available, will be deposited in the collec- tions of the University of California, Riv- erside and the California Academy of Sci- ences, San Francisco. Culicoides (Haematomyidium) kettlei Breidenbaugh and Mullens, new species (Figs. 1—4) Egg.—Banana-shaped. Surface with two types of ansulae arranged in longitudinal rows (Fig. 1A). Moderately stalked ansulae present on concave surface, ridges com- posed of flattened papillate ansulae on con- vex surface (Fig. 1B). Average length = 356) 2) 17 we widthy— 535-222 (=) 11): Larva.—Total length = 2.79 (2.16—3.51, n = 15) mm. Head capsule (Figs. 2A—C): Light brown. Small larvae, HL = 128 (122- 134, n = 29) p, HW = 89 (82-102, n = 28) ww; shape somewhat long and narrow, SGW = 58 (54-67, n = 30) pw; HR = 1.4 (1.2-1.6, n = 27); SGR = 1.5 (1.4-1.8, n = 27). Mandible (Fig. 2F) medium-small, ML = 35 (22-38, n = 29) wp; with pointed apex and subapical rounded prominence; base with distinct point of articulation. Hy- postoma (Fig. 2G) rounded, entire. Labium small, crescent-shaped and sclerotized. Epi- pharynx (Fig. 2H): Dorsal-comb sclerites with 5—7 unequal angular teeth; moderately- wide, DCW = 11 (10-13, n = 17) p. Comb 2 narrow with short, rounded teeth; comb 4 wider, with rounded teeth. LAW = 49 (43-— 52, n = 30) p; lateral curtains composed of short, dense, filamentous processes. Hypo- pharynx apparently poorly sclerotized, not seen. Thoracic pigmentation (Fig. 21): dis- tinct lateral pattern on thoracic segments. Prothorax with arched pattern; mesothorax and metathorax with crescent and sagittate patterns, respectively. Caudal segment (Figs. 2D, E): Short, CSL = 231 (189-285, n = 32) w and moderately-wide, CSW = 126 VOLUME 101, NUMBER 1 Fig. 1. Culicoides kettlei, egg. A, Lateral view. B, Detail of surface. O0=V79 ym —3)1 Oval CSRe= 129 5 — 2.2, n = 31). Setae ‘“‘o”’ short relative tothe 45, n = CSL, OL = 46 (38-51, n = 19) pw; bases of setae moderately separated, OD = 37 (26— 19) w. Anal papillae not observed. Material studied.—Laboratory-reared from 152 Setae “QO” al Figs 2: Setae “O” 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Culicoides kettlei, larva. A-C, Head capsule. A, Dorsal view. B, Ventral view. C, Lateral view. D, Caudal segment, dorsal view. E, Caudal segment, ventral view. K Mandible. G, Hypostoma. H, Epipharynx. I, Thoracic pigmentation, dorsal view. eggs deposited by individual females collect- ed in San Bernardino County, CA, Morongo Valley, 15-VI-95 (n = 3); 22-V-96 (n = 5); 14-VI-96 (n = 24), paratypes: slides 29,202 Pupa.—Light yellow brown. Respiratory horn (Fig. 3A): yellow brown; basal half crenulated; spines, sparsely distributed. Four lateral spiracules present and 8-11 apical spiracles. Tracheal annulations asso- ciated with tracheae visible in basal portion and a reticulated pattern is visible in distal portion. Operculum (Fig. 3B): Yellow brown, with two types of spines, neither ex- tending beyond the am tubercles. Sagittate spines occurring laterally; smaller, rounded spines occurring medially. Anterior margin moderately notched. Caudal segment (Fig. 3C): Spines present at base of posterolat- eral processes, absent apically; band of spines along anterior margin and a patch of spines located basimedially on dorsum. Chaetotaxy: Dorsal tubercles (Fig. 3D): 1 and 2 round with stout spine; 3 round with a short seta; 4 a slender seta; 5 a small, round pore. ad tubercle (Fig. 3E): spinate with two subequal spines. d/ tubercle (Fig. 3F): with 3 unequal, slender spines. Abdo- men (Fig. 3G): dasm tubercles: 1 with a spine and 2 a bristle. dpm tubercles: round- ed; | and 2 with short spine; 3 and 4 lack setae; 5 with a short hair. /asm tubercle: spi- VOLUME 101, NUMBER 1 153 F prue~ WoW Fig. 3. E, ad tubercle. E dl tubercle. G, Abdominal chaetotaxy. nate with short spine. /pm tubercles: 1—3 spinate; | and 3 with short spine; 2 with a bristle. vpm tubercles: rounded; 1 with a spine; 2 and 3 with long and medium length setae, respectively. Material studied.—Laboratory-reared from individual females collected in San Bernardino County, CA, Morongo Valley, 14-V-95 (n = 6), paratypes: slides 10,11. Adults.—Female: Wing length 1.06 (0.92—1.19, n = 28) mm. Head: Eyes (Fig. 4A) bare; separated by a single facet width; without interfacetal hairs. Antenna (Fig. Culicoides kettlei, pupa. A, Respiratory horn. B, Operculum. C, Caudal segment. D, Dorsal tubercles. 4B) with lengths of flagellomeres of holo- type 43-27-28-33-29-31-30-33-36-37-43- 47-63 (in w); antennal ratio 0.91 (0.74— 1.04, n = 28); well-developed sensilla coe- loconica present on flagellomeres 1, 4—8; 26% (7/27) of observed specimens lack sensilla on flagellomere 4. Palpus (Fig. 4C) with lengths of segments 24-46-61-20-26; palpal ratio 2.07 (1.75—2.36, n = 29); P/H ratio 0.79 (0.70—0.86, n = 18); third seg- ment swollen with broad shallow sensory pit. Thorax: Brown, scutum without con- spicuous pattern. Legs brown (Fig. 4D), 154 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Tip : : oe 1 aa : KAN SKA Fue Wp ene wis 2 Zee Re Wg ory aS AIL ARENA ee RK ES eee TPE te ON eee ee eins 5 ; alos ANSON Sie 5 x Tee get eH mite Wises - = ~ oy Sy, a ae By ya = mA wolst oy : oo BG SE eto coal. oo a aN . Ee, ees eal E Weck ve et \ i nore wy POTTY prspep yey opiate | Culicoides kettlei, adult. A-E Female, G-H, Male. A, Eye separation. B, Antenna. C, Palpus. D, Legs, left to right, front, mid, and hind. E, Wing. F. Spermathecae. G, Genitalia, parameres omitted. H, Parameres. femorotibial joints blackish; fore and mid femora with narrow subapical pale rings, all tibiae with narrow subbasal pale rings and hind tibiae with distal fourth pale; hind tib- ial comb with five spines, second from the spur longest. Wing (Fig. 4E) with a slightly angular poststigmatic pale spot; distinct transverse pale spot present in r5; cell ml with narrow pale spot; cell m2 with round- ed pale spot located distally and with a lon- gitudinal pale spot lying posterior to medial fork. Macrotrichia in moderate numbers VOLUME 101, NUMBER 1 evenly distributed over wing. Halter pale. Abdomen: Brown. Two slightly unequal spermathecae, plus rudimentary third and sclerotized ring (Fig. 4F). Functional sper- mathecae with long slender necks; length, including neck, 32 (29-37, n = 25) w for the larger, 29 (26-35, n = 26) wp for the smaller. Male: Genitalia (Figs. 4G—H): Ninth sternum with scarcely perceptible caudo- median excavation, ventral membrane spic- ulate; apicolateral processes moderately long, pointed, caudal margin between them slightly cleft medially. Gonocoxite moder- ately stout; ventral root with anterior point longer, more slender than dorsal root; gon- ostylus moderately curved and slender with moderately bent, pointed tip. Aedeagus with basal arch extending more than two- thirds of total length; distal portion short, tapering to a simple tip. Paramere (Fig. 4H) with strong basal knob; midsection arched; distally, greatly narrowed and reflexed upon midsection, constricting to a noticeably fringed filamentous tip. Distribution.—Southern California and northern Baja California, Mexico. Material studied—HOLOTYPE: 2°, Riv- erside County, CA, Deep Canyon, Bighorn Overlook, nr. Palm Desert, 22-IX-88 (B. A. Mullens), CO,-baited trap. ALLOTYPE: <6, San Bernardino County, CA, Big Morongo Canyon, Morongo Valley, 15-VI-95 (Brei- denbaugh), laboratory-reared from wild- collected female. PARATYPES: CALI- FORNIA. 6 2, same data as holotype, ex- cept 1 2 24 May 1988; 6 2, Riverside Co., Bighorn Drive, Palm Desert, 8 October 1988, CO,-baited trap, 1 2, same except 22 September 1988; 9 2, 4 6, same data as allotype except from June 1995 to July 1996 (some K. Luhring); MEXICO. 7 2°, Mexico, Baja California, Cadavina, 19 March 1994 (C. Szijj), CO,-baited trap. Behavior and rearing.—In the laboratory, the average female fecundity was 74 + 24 eggs (n = 10). From a single reared cohort observed daily, pupation first occurred 26 days following egg hatch. In culture, larvae 155 fed on Pelodera sp. nematodes. Adults have been collected from soil-emergence traps along a seasonal creek in the Deep Canyon watershed (Breidenbaugh and Mullens, in preparation), indicating that creek margins are used by this species as a larval devel- opment site. Feeding behavior is poorly known with a single report of an unfed fe- male recovered from a bighorn sheep (Mul- lens and Dada 1992b). Discussion.—Culicoides kettlei, was originally recognized by the late W. W. Wirth who referred to this species as #120 (W. W. Wirth, personal communication). Following Vargas (1960) the male genitalia fit nicely into the subgenus Haematomyi- dium. The female wing pattern is inconclu- sive, since the r-m crossvein is not dark. However, we feel that placement in the sub- genus Haematomyidium is reasonable. Adult morphology is similar to that of Cu- licoides (Haematomyidium) debilipalpis Lutz, a common biting midge in the eastern U.S.. From 1985-1997 C. debilipalpis was synomized with C. lahillei Iches (Spinelli and Wirth 1985). Recently, however, the validity of C. debilipalpis was confirmed (Spinelli and Ronderos 1997). The range of C. debilipalpis includes the southeastern U.S. as far west as Louisiana, with a dis- junct distribution that includes Costa Rica south to Argentina. In contrast, C. lahillei is strictly South American. The females of C. kettlei can be easily separated from C. debilipalpis and all other Nearctic members of Haematomyidium by the antennal sen- sorial pattern (1,4(5)-8) (Wirth et al. 1985). In addition, no Neotropical species of Hae- matomyidium are known to occur north of southern Mexico (Wirth et al. 1988), and their range thus does not overlap with that of C. kettlei. Apart from the similarity in size, the lar- vae of this species are noticeably different from those of the only other North Ameri- can species in this subgenus with described larvae, Culicoides paraensis (Goel- di)(Murphree and Mullen 1991). The hy- postoma is round and smooth in this species 156 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON while lateral teeth are present in C. paraen- sis. Furthermore, the hypostoma lacks the distinct subapical notch seen in C. paraen- sis. The pupa of this species is similar to C. paraensis in the type and location of spines on the operculum. Etymology.—This species is named in honor of Dr. D. S. Kettle, Emeritus Profes- sor, Department of Entomology, University of Queensland, Australia, for his pioneering work on the biology of Culicoides, specif- ically in the discipline of the morphology of immatures. Culicoides vetustus Breidenbaugh and Mullens, new species (Figs. 5-8) Egg.—Cigar-shaped. Surface with flat- tened longitudinal rows, many not contig- uous from end to end (Fig. 5A). Ansulae flattened, not distinct, merge to form lon- gitudinal ridges (Fig. 5B), present on all lat- eral surfaces, and not varying with curva- ture. Average length = 267 + 28 p and width = 49 + 3 p (n = 8). Larva.—Total length = 3.25 (2.81—4.0, n = 15) mm. Head capsule (Figs. 6A—C): Yellow; medium sized, HL = 180 (166— 192, n = 28) p, HW = 123 (109-138, n = 27), 2, SGW. =, 90 (80-102, n= 28)-p; overall shape long and narrow, HR = 1.5 (1.2—1.7, n = 27), very oblong, SGR = 1.4 (1.2—1.5, n = 27). Mandible (Fig. 6D) me- dium length, ML = 51 (48—54, n = 24) p; base wide; pointed marginal prominence basimedially, followed by a subapical notch, sharply angled to pointed apex. Hy- postoma (Fig. 6E) difficult to see but round- ed medially and smooth. Epipharynx (Fig. 6F): Dorsal-comb sclerites moderately wide, DCW = 14 (13-15, n = 24) yp, with 5 subequal pointed teeth/sclerite; comb 4 with many unequal pointed, rounded teeth; lateral curtains wide; teeth thin, hair-like; LAW wide relative to DCW, LAW = 60 (52—67, n = 21) wp, indented near lateral apex. Hypopharynx (Fig. 6G) with hypo- pharyngeal fringe separated into two dis- tinct prominences by a medial notch. Tho- racic pigmentation (Fig. 6H): Absent. Cau- dal segment: Short with length varying considerably among individuals, CSL = 286 (131-326, n = 25) wp; narrow, CSW = 139 (86-157, n = 26) pw, CSR = 2.1 (1.5— 2.4, n = 25); setae “‘o”’ of medium length, OL = 82 (61-99, n = 18) wp, and well sep- arated, OD = 45 (35-53, n = 21) w. Anal papillae (Fig. 61) four deeply bifurcate pairs. Material studied.—Laboratory-reared from eggs deposited by individual females collected in San Bernardino County, CA, Morongo Valley, 15-VI-95 (n = 6); 14-V- 96 (n = 15); 14-V-96 A42 (n = 15), para- types: slides 2-7. Pupa.—Light to dark brown. Respiratory horn (Fig. 7A): Proximal portion light brown; roughly divided into 3 subequal portions. First section with annular trache- ae; second portion annulations obscured by reticulation pattern; 3 lateral spiracular pro- tuberances present; terminal section, dark brown with 5—9 spiracular openings apical- ly; few scales on horn. Operculum (Fig. 7B): Mostly smooth, 3—8 large spines on lateral margins and patch of setae on central portion; no large spines medial to the am tubercles; large spines triangular. Caudal segment (Figs. 7C—D): Narrow V-shaped cluster of small spines on dorsal surface of both sexes. Anterior band of spines com- plete on both sexes; large spines on proxi- mal % of posterolateral processes; distal third, smooth dark brown. Chaetotaxy: Dorsal tubercles (Fig. 7E): setae | and 2 of medium length, stout; seta 3 short, stout; seta 4 a long, slender bristle; 5 a circular pore. ad tubercle (Fig. 7F) with one long, one shorter setae. d/ tubercle (Fig. 7G) with a long slender bristle and second shorter thicker bristle. Abdomen (Fig. 7H): dasm tubercles: setae | and 2 of medium thickness, | longer than 2. dpm tubercles: setae 1 and 2 equal; 3 and 4 lack setae; 5 short. /asm tubercle: rounded with short spine. /pm tubercles: spinate with shorter, stouter, equal bristles on | and 3, bristle 2 longer and slender. vpm tubercles: 2 with a VOLUME 101, NUMBER I bys “yA / 4. Ba aly +4. gf” LF BEKY K2606 HH13 16.6U Culicoides vetustus, egg. A, Lateral view of entire egg. B, Detail of egg surface. fe 158 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Mandible. E, Hypostoma. EK Epipharynx. G, Hypopharynx. H, Head and thorax, dorsal view. I, Anal papillae, caudal segment. slender bristle, longer and thinner than 1 and 3. Material examined.—Laboratory-reared from eggs deposited by individual females collected in San Bernardino County, CA, Morongo Valley, 14-V-96 (n = 8); 15-VI- 95 A5 (n = 17), paratypes: slides 17-18; 14-V-96 A42 (n = 3 ), paratypes: slide 1. Adults.—Female: Wing length 1.24 (1.1-— 1.4, n = 20) mm. Head: Eyes (Fig. 8A) separated by single facet width; without in- terfacetal hairs. Antenna (Fig. 8B) lengths Culicoides vetustus, larva. A-C, Head capsule. A, Dorsal view. B, Ventral view. C, Lateral view. D, of flagellomeres of holotype 46-28-28-30- 30-30-33-33-43-46-46-50-65; antennal ratio 0.97 (0.92-1.0, n = 18); well-developed sensilla coeloconica present on flagello- meres 1, 11-13. Palpus (Fig. 8C) with lengths of segments 24-50-78-30-33; third segment swollen, with round moderately deep sensory pit near apex. Palpal ratio 2.06 (1.8—2.3, n = 21); proboscis long, P/ H ratio 0.95 (0.81-1.3, n = 19). Thorax: Brown, lateral aspects of scutum darker. Legs brown, lacking pattern; hind tibial VOLUME 101, NUMBER 1 Ipm Fig. 7. comb (Fig. 8D) with 5 spines, second from spur longest. Wing (Fig. 8E) uniformly grayish with pale spot on anterior margin just past second radial cell. Halter pale. Ab- domen: Brown. Two slightly unequal sper- mathecae (Fig. 8F), plus rudimentary third; functional spermathecae with short slender necks; length 44 (35—51, n = 18) p for larg- er, 42 (34-50, n = 17) p for smaller. Male: Genitalia (Figs. 8G—H): Sternite 9 with broad moderately deep caudomedian excavation, ventral membrane spiculate; tergite 9 with prominent triangular apico- lateral processes, moderately broad, slightly divergent; the caudal margin between them transverse without medial cleft. Gonocoxite moderately stout; ventral root more slender 159 Wr Culicoides vetustus, pupa. A, Respiratory horn. B, Operculum. C, Caudal segment, male. D, Caudal segment, female, E, Dorsal tubercles. F ad tubercle. G, dl tubercle. H, Abdominal chaetotaxy. than dorsal root; gonostylus curved distally, moderately slender, with moderately broad blunt tip. Aedeagus Y-shaped; basal arch extending half of total length; basal arms short, moderately slender, recurved at ex- treme apex, distal portion moderately broad. Paramere (Fig. 8H) with distinct an- terior process; midpoint straight; apex di- verging with three sharp medial spines, tip sharply pointed, diverging. Distribution.—Mojave and Colorado de- serts of southern California. Material examined.—HOLOTYPE: 2°, Riverside County, CA, Deep Canyon, nr. Palm Desert, 29-VI-89 (B. A. Mullens), CO,-baited trap. Allotype: d, San Bernar- dino County, CA, Big Morongo Canyon, 160 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 8. Hind leg. E, Wing. K Spermathecae. G, Genitalia, parameres omitted. H, Parameres. Morongo Valley, 15-VI-95 (Breidenbaugh), laboratory-reared from wild-collected fe- male. PARATYPES: | 2, same data as al- lotype; 16 2, 11 ¢ same data as allotype, except collected between 14-V-96 and 2- VII-96, some reared from wild-collected °; 9 2, San Diego County, CA, Anza Borrego State Park, Yaqui Well, 26-IV-97 (Breiden- baugh), CO,-baited trap. Biology and rearing.—Adult females collected in CO,-baited traps readily fed as Abid bidldbefe ih eg id Cldlatyy, oO LLL ALO EPR aa ~ Fi x 7 os . ‘ “ 2 mes : aS " P: . : 2 : Itaeaeee ‘ o% "vp spangp yon i anny E H Culicoides vetustus, adult. A-F Female. G-H, Male. A, Eye separation. B, Antenna. C, Palpus. D, through an artificial membrane and depos- ited eggs in the laboratory. Larvae exhib- ited the slowest development of any species reared on the agar system. Average clutch size was 86 + 35 eggs (n = 52 females); these hatched in 7 + 1 days with 67 + 25% fertility (n = 25 females). Average devel- opment period to pupation was 117 + 28 days (n = 46). Adults emerged 4 + 1| days later (n = 39). It is not known if the rearing medium VOLUME 101, NUMBER 1 was responsible for the slow development or whether this is the normal condition for larvae of this species. The percentage of larvae reaching adulthood (approx. 31%) was similar to the other species reared (Breidenbaugh and Mullens, in prepara- tion). Despite extensive observations, lar- vae were never seen feeding on nematodes; thus, this species was probably utilizing other microorganisms present in the cul- tures. Discussion.—As discussed by Blanton and Wirth (1979), adults of a number of Nearctic Culicoides do not key readily to the subgenera described by Vargas (1960). This is true for C. vetustus. From adult and larval characteristics presented here, we are tentatively placing C. vetustus in the bigut- tatus species group. For example, the man- dibles of the larvae resemble those illus- trated by Murphree and Mullen (1991) of Culicoides biguttatus (Coquillett) and Cu- licoides spinosus Root and Hoffman except for the depth of the notch on the subapical margin of the mandible of C. vetustus. The number of dorsal-comb teeth (5) on the epi- pharynx is the same for C. spinosus. How- ever, the lateral arms of the epipharynx are notched in C. vetustus, but not in C. spi- nosus or C. biguttatus. The hypostoma of C. biguttatus and C. vetustus are very sim- ilar. The pupal operculum of C. vetustus re- sembles that of C. biguttatus and C. spi- nosus. However, the operculum of C. ve- tustus has fewer spines, and these are lim- ited to the lateral margins. The respiratory horn of C. vetustus is more similar to that of C. spinosus, darkened at the tip and with 3 spiracular openings visible. The range of C. vetustus overlaps only with C. sublettei Atchley and C. usingeri Wirth of the biguttatus group. The male genitalia of these species are similar in sev- eral respects, including the general shape of the parameres. The apices of the parameres of C. vetus- tus are divided into 3—4 short spines and the distomedian process of the aedeagus is 161 bluntly rounded. The aedeagus of C. vetus- tus is more slender than that of C. sublettei or C. usingeri, and there are 5 tibial spines in C. vetustus and C. sublettei, but 4 in C. usingeri. The females of C. vetustus lack a very distinctive wing pattern. However, they can be distinguished clearly from the other species in the biguttatus group and from Culicoides piliferus Root and Hoff- man, which it superficially resembles, by the sensorial pattern (1, 11—13). Etymology.—Latin vetustus for long- lived, referring to the lengthy development period of the larvae. DISCUSSION Most descriptions of Culicoides species have included only the adult stage or in some cases, only the female. This is a result of the relative difficulty in locating devel- opmental sites or collecting and associating males. The collection of host-seeking fe- males, use of an artificial host, and a labo- ratory rearing system proved a very suc- cessful way of associating immature stages with adults. This approach should be useful for many species whose adults are known. The rearing technique is advantageous in that the entire cohort from an isofemale is unquestionably conspecific. In contrast, field-collected larvae, if they are located, generally must be killed and slide-mounted for identification. Consequently, an incor- rect association with other larvae in the col- lection which have been reared to adults is possible. The rearing method used here probably results in a narrower range of measurements for morphometric analysis, than if field-col- lected material were examined. Environ- mental homogeneity and the genetic simi- larity of laboratory-reared sibling larvae can result in morphological measurements that are unnaturally similar. The descrip- tions herein, however, will allow field-col- lected immatures to be identified and mea- sured to better characterize the range of nat- ural variability in characters such as size. 162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ACKNOWLEDGMENTS We thank K. Luhring (University of Cal- ifornia, Riverside) for help with collections and rearing; W. W. Wirth (deceased) for as- sistance with initial discovery of these new species; W. L. Grogan (Salisbury State Uni- versity, MD); and the California Academy of Sciences which supplied adults of C. de- bilipalpis. We appreciate the comments of W. L. Grogan and C. S. Murphree (Belmont Uni- versity, TN) on the manuscript as well as A. Borkent (Enderby, British Columbia), W. L. Kramer (Nebraska Dept. of Health and Human Services, NE), and G. R. Spi- nelli (Museo La Plata, Argentina) for ad- vice. This study was supported in part by the Anderson Endowed Fellowship award- ed to MSB and by USDA-NRICGP #94- 37312-06232 to BAM. LITERATURE CITED Becker, P. 1961. Observations on the life cycle and immature stages of Culicoides circumscriptus Kieff. (Diptera: Ceratopogonidae). Proceedings of the Royal Society Edinburgh 67: 363-388. Blanton, F S. and W. W. Wirth. 1979. The sand flies (Culicoides) of Florida (Diptera: Ceratopogoni- dae). Arthropods of Florida and Neighboring Land Areas, Vol. 10, 204 pp. Borkent, A. and W. W. Wirth. 1997. The World species of Ceratopogonidae (Diptera). Bulletin of the American Museum of Natural History 233, 257 Pp. Campbell, M. M. and D. S. Kettle. 1975. Oogenesis in Culicoides brevitarsis Kieffer (Diptera: Cerato- pogonidae) and the development of a plastron-like layer on the egg. Australian Journal of Zoology 23: 203-218. Day, J., D. Duzak, Y. Braverman, and A. Chizov-Ginz- burg. 1997. Ultrastructure of the eggs of Culicoi- des circumscriptus, Culicoides gejgelensis, and Culicoides imicola (Diptera: Ceratopogonidae). Journal of the American Mosquito Control Asso- ciation 13: 76-83. Deforge, J. R. and J. E. Scott. 1982. Ecological inves- tigations into high lamb mortality. Transactions of the Desert Bighorn Council 26: 65-76. Downes, J. A. and W. W. Wirth. 1981. Chapter 28, Ceratopogonidae, pp. 393—421. In McAlpine, J. FE et al., eds. Manual of Nearctic Diptera, Vol. 1. Research Branch Agriculture Canada Monograph DT Elliott, L. F, W. M. Boyce, R. K. Clark, and D. A. Jessup. 1994. Geographic analysis of pathogen ex- posure in bighorn sheep (Ovis canadensis). Jour- nal of Wildlife Diseases 30: 315-318. Hunt, G. J. 1994. A procedural manual for the large- scale rearing of the biting midge, Culicoides var- lipennis (Diptera: Ceratopogonidae). United States Department of Agriculture Research Service, ARS-121. Jones, R. H., H. W. Potter, and S. K. Baker. 1969. An improved larval medium for colonized Culicoides varlipennis (Diptera: Ceratopogonidae). Journal of Economic Entomology 62: 1483-1486. Lamberson, C., C. D. Pappas, and L. G. Pappas. 1992. Pupal taxonomy of the tree-hole Culicoides (Cer- atopogonidae: Diptera) in eastern North America. Annals of the Entomological Society of America 85: 111-120. Mullens, B. A. and C. E. Dada. 1992a. Spatial and seasonal distribution of potential vectors of hem- orrhagic disease viruses to peninsular bighorn sheep in the Santa Rosa Mountains of southern California. Journal of Wildlife Diseases 28: 192— 205. 1992b. Insects feeding on desert bighorn sheep, domestic rabbits, and Japanese quail in the Santa Rosa Mountains of southern California. Journal of Wildlife Diseases 28: 476—480. Mullens, B. A. and R. K. Velten. 1994. Rearing Cu- licoides variipennis sonorensis (Diptera: Cerato- pogonidae) on agar and nematodes. Journal of Medical Entomology 31: 175-177. Murphree, C. S. and G. R. Mullen. 1991. Comparative larval morphology of the genus Culicoides La- treille (Diptera: Ceratopogonidae) in North Amer- ica with a key to species. Bulletin of the Society of Vector Ecology 16: 269-399. Nevill, H. and A. L. Dyce. 1994. Afrotropical Culi- coides: Description and comparison of the pupae of seven species of the similis supergroup (Cera- topogonidae: Diptera). Onderstepoort Journal of Veterinary Research. 61: 85-106. Spinelli, G. R. and W. W. Wirth. 1985. Clave para la identificaci6n de las especies del género Culicoi- des Latreille presentes al Sur de la Cuenca Ama- zonica. Nuevas citas y notas sinonimicas (Diptera: Ceratopogonidae). Review of the Society of En- tomology, Argentina 44: 49-73. Spinelli, G. R. and M. M. Ronderos. 1997. Description of the male of Culicoides lahillei and revalidation of Culicoides debilipalpis (Diptera: Ceratopogon- idae). Transactions of the American Entomologi- cal Society 123: 303-309. Vargas, L. 1960. The subgenera of Culicoides of the Americas (Diptera, Ceratopogonidae). Revista de Biologia Tropical 8: 35—47. Wehausen, J. D., V. C. Bleich, B. Blong, and T. L. Russi. 1987. Recruitment dynamics in a southern VOLUME 101, NUMBER 1 California sheep population. Journal of Wildlife Management 51:86—98. Wirth, W. W. and N. Marston. 1968. A method for mounting small insects on microscope slides in Canada balsam. Annals of the Entomological So- ciety of America 61: 783-784. Wirth, W. W., A. L. Dyce, and B. V. Peterson. 1985. An atlas of wing photographs, with a summary of the numerical characters of the Nearctic species of Culicoides (Diptera: Ceratopogonidae). Contribu- tions of the American Entomological Institute 22: 1—46. 163 Wirth, W. W., A. L. Dyce, and G. R. Spinelli. 1988. An atlas of wing photographs, with a summary of the numerical characters of the Neotropical spe- cies of Culicoides (Diptera: Ceratopogonidae). Contributions of the American Entomological In- stitute 25: 1-72. Wirth, W. W. and B. A. Mullens. 1992. Culicoides boy- di (Ceratopogonidae: Diptera): A potential vector of hemorrhagic disease viruses to desert bighorn sheep in southern California. Journal of Medical Entomology 29: 1006-1010. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 164-174 BLASTOBASIS GRAMINEA, NEW SPECIES (LEPIDOPTERA: GELECHIOIDEA: COLEOPHORIDAE: BLASTOBASINAE), A STEM BORER OF SUGAR CANE IN COLOMBIA AND VENEZUELA DAVID ADAMSKI Research Associate, Department of Entomology, MRC-127, Smithsonian Institution, Washington, DC 2056-0127, U.S.A. Abstract.—Blastobasis graminea, new species, a stem borer of sugar cane in Colombia and Venezuela, is described and illustrated. For the first time, a larva of Blastibasini is described in detail. Scanning electron micrographs of the larva, illustrations of the larval mandible, setal maps, and photographs of larval damage and pupation sites are provided. Auximobasis obstricta Meyrick 1918, is transferred to Blastobasis Zeller 1855, n. comb.., and Blastobasis subolivacea Walsingham 1897, is transferred to Holcocera Clemens 1863, n. comb. Key Words: For decades, entomologists have known that larvae of at least one species of micro- lepidoptera other than Diatraea saccharalis (Fabricius) (Crambidae) feed on sugar cane and related grasses in Latin America. Al- though adult specimens of one species of Coleophoridae (Blastobasinae) have been collected since the late 1940’s and 1950’s by H. E. Box in Venezuela and during the early 1970’s and 1980’s by L. Cardenas and others in Colombia, this moth remained nameless. Because many Blastobasinae are similar in wing pattern, they are frequently mis- identified. For example, the species de- scribed herein, Blastobasis graminea, had been previously misidentified as Auximo- basis obstricta Meyrick 1918, (Box 1953, Guagliumi 1962) and Blastobasis suboli- vacea Walsingham 1897, (Martorell 1976). In addition, type specimens of Neotropical Blastobasinae have not been studied sys- tematically until recently. Since Meyrick (1894) the Blastobasinae have long been considered to be monophy- Coleophoridae, Blastobasinae, Blastobasis, sugar cane, Colombia, Venezuela letic; recent studies (Adamski and Brown 1989, Hodges, in press) have corroborated this notion and postulated phylogenetic re- lationships of the Blastobasinae within Ge- lechioidea. In this study, the Blastobasidae (sensu Adamski and Brown 1989) are treat- ed as a subfamily within the Coleophoridae, following Hodges (in press). The purpose of this paper is to describe and illustrate Blastobasis graminea, new species, and to make available to entomol- ogists and sugar cane growers a means by which to identify it. Adult and larvae were examined using an incandescent light source (reflected light). Kornerup and Wanscher (1978) was used as a color standard for the description of the adult. Genitalia were dissected as described by Clarke (1941), except Mercurochrome and chlorazol black were used as stains. Slide preparations were examined with dis- secting and compound microscopes. Mea- surements were made with a calibrated oc- ular micrometer. All specimens examined are deposited in The National Museum of VOLUME 101, NUMBER 1 165 I Holotype of Blastobasis graminea. Fig. 1. Natural History, Smithsonian Institution, Washington, D.C., (USNM), except where indicated otherwise. Label data taken ver- batim are expressed with quotations, while bracketed data are used to complete label data written in abbreviated form, or to help with the recognition of certain labels by de- scription of condition, e.g., [round label]. The ultrastructure of the larva was stud- ied with an Hitachi HH-S-2R scanning electron microscope at an accelerating volt- age of 20 kV. For SEM examination, larvae were fixed in 3% glutaraldehyde in 0.1 M potassium phosphate buffer (pH 7.3), rinsed in phosphate (pH 7.3), and postfixed in 2% osmium tetroxide in 0.1 M potassium phos- phate (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 in a Polaron E5100 sputter coater. Blastobasis graminea Adamski, new species (Figs. 1-26) Diagnosis.—Blastobasis graminea can be distinguished from other Blastobasis by the orange gray ground color, wide base of the uncus, rounded outer margin of the proximal flange, and wide ostial opening. Adult.—Head: Cephalic vestiture pale orange gray, except inner surface of labial palpus pale orange gray intermixed with brown scales tipped with white and few dark-brown scales, outer surface mostly brown intermixed with brown scales tipped with white, pale orange-gray scales, and dark-brown scales; segments paler near api- cal region. Thorax: Tegula and mesoscutum pale or- ange gray; legs with outer surface pale or- ange gray intermixed with orange-gray scales tipped with white, most specimens with foreleg and midleg with outer surface mostly grayish brown intermixed with grayish-brown scales tipped with white, or- ange-scales, and orange-gray scales tipped with white, leg segments and tarsomeres paler near apical region; forewing (Fig. 1), length 7.1—-10.0 mm [n = 37], orange gray intermixed with orange gray scales tipped with white, brown scales tipped with white, and brown scales; several unrubbed speci- mens with discal cell region paler than out- 166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2, 3. Blastobasis graminea. 2, Wings, scale er region of wing; holotype with a brown streak on basal part of posterior margin (Fig. 1); some specimens with veins de- marcated with white scales; a dark brown 1.00 mm. 3, Male genitalia, scale = 0.5 mm. midcell spot and two distal spots usually present; fringe scales mostly orange gray tipped with white intermixed with orange gray scales; undersurface grayish brown; VOLUME 101, NUMBER 1 cubitus four-branched, divergent from ra- dials and M, (Fig. 2); hindwing with both surfaces pale grayish brown; cubitus four- branched in a series typical of all New World Blastobasis (Fig. 2). Abdomen: Orange gray. Male Genitalia (Fig. 3): Uncus wide at base, posteriorly curved and narrowed api- cally, apical setae shorter than basal setae; gnathos bidentate; dorsal strut narrow; ter- gal setae numerous; diaphragma with mi- crotrichia throughout, extending to proxi- mal flange; proximal flange with stout mar- ginal setae, margin rounded; lower part of valva with marginal setae, numerous along apical third; juxta bandlike; aedoeagus api- cally rounded, with several stout anellar se- tae. Female Genitalia (Fig. 4): Ovipositor telescopic, in four membranous divisions; ostium within membranous area slightly posterior to seventh sternum; ostial opening wide; antrum membranous, narrowed abruptly anteriorly forming a common in- ception for ductus seminalis and ductus bursae; ductus bursae long, with two rows of platelike sclerotizations within anterior part; corpus bursae with posterior lobe near inception of ductus bursae; signum horn- like. Larva.—Length 6.5—14.9 mm [n = 207]. Body white, smooth, with head capsule, prothoracic shield, anal shield, pinacula and crochets yellowish orange. Head (Figs. 5— 12, 17, 18): Hypognathous; epicranium smooth; adfrontal sclerites narrow, delim- iting frons dorsolaterally; frons closed; C1, C2, and C3 about equal in length, about three times length of Fl and F2; C3 closer to Fl than to C2; C2 slightly closer to mid- line than Cl or C3; C1 setae broadly curved, convergent; Fl subapical (Fig. 5); P1 long, closer to P2 than A2; A2 closer to Al than A3; A3 nearly equidistant to A2 and L1 (Figs. 5—7); S1 between stemmata 2 and 3, and closer to S2 than to S3; SS1 near mandibular articulation, and closer to SS2 than to SS3; SS2 between stemmata 5 and 6 (Figs. 6, 7); labrum bilobed, each 167 Fig. 4. Female genitalia of Blastobasis graminea. Scale = 1.00 mm. lobe with four subequal marginal setae and two subequal medial setae (Figs. 5, 6); mandibles slightly asymmetrical, with two distinct dentitions and two subequal setae on outer margin (Figs. 5, 6, 17); labium smooth with microtrichia along lateral mar- gin of proximal half; distal part of labium with median submental pit (Figs. 6, 8); la- bial palpus two-segmented, with dorsally directed subapical seta on basal segment. Sensilla types and arrangement on median 168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5-10. > - SEM of larva of Blastobasis graminea. 5, Frontal view of head capsule, 70X. 6, Ventral view of head capsule, 70. 7, Lateral view of genal region of head capsule, 250. 8, Ventral view of labium, LBP = labial palpus, SPIN = spinneret, 250. 9, 10, Sensilla on apex of maxillary palpus, A2 = sensillum stylo- conicum; Al, M1, M2, L1, L2, L3, = sensilla basiconica; SD = sensilla digitiform, 2,500. lobe and apex of palpus similar to that of Glyphidocera juniperella Adamski and Brown 1987, except for elongate depression near digitiform sensillum on part near L3 sensillum (Figs. 8-10). Sensilla types on antenna (Fig. 11) similar to other Lepidop- tera (Schoonhoven and Dethier, 1966). Pro- thorax (Figs. 12, 18): Prothoracic shield with SD1 and D2 about equal in length, twice length of XD1 and XD2; SD1 and D2 VOLUME 101, NUMBER 1 Figs. 11-16. SEM of larva of Blastobasis graminea. 11, Sensilla on apical portion of antenna. 1 = sensilla basiconica; 2 = sensillum chaetica; 3 = sensullum styloconicum; 4 = sensillum trichodeum, 950. 12, Lateral view of head capsule and thorax, 45x. 13, Ventral view of right proleg on A4, 200. 14, A8-10, 60. 15, Lateral view of A10, 80X. 16, Posetrior view of A10, 80%. about four times length of SD2 and D1; D1 usually slightly longer than SD2; SD2 clos- er to SD1 than to XD2; SD2 and D2 anter- iorad to D1; L1 about twice length of L2 and L3; SV1 about twice length of SV2; V1 short (not illustrated). Mesothorax and metathorax (Figs. 12, 18): D1 anterodorsal to D2, on same pinaculum; D2 about three times length of D1; SD1 anterioventral to SD2, on same pinaculum; SD1 about three 170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 17-20. 19, Al-3. 20, A7-10. times length of SD2; L2 anterioventral to L1, on same pinaculum; LI] slightly longer than L2 and about three times length of L3; SV1 about equal in length to L1 and slight- ly caudal to L3; V1 short (not illustrated). Abdomen (Figs. 13—16, 19, 20): Al and A2 with D2 three times length of D1; SD1 about same length as D2; SD2 very short (Figured larger than normal), on same pin- aculum as SD1 above spiracle; L1 antero- dorsal to L2, L2 about twice length of L1; L3 caudal to D2, about equal in length to L1; SV3 2-3 times length of SV2 and SV1; D2 Larva of Blastobasis graminea. 17, Left mandible. 18, Lateral view of head capsule and thorax. SV2 and SV1 in nearly straight line per- pendicular to longitudinal body axis, SV3 slightly anterior to SV1; V1 short (not il- lustrated); A3—-A6 with SV1 posterior to SV2; prolegs with crochets uniserial and triordinal, crochets smaller along outer mar- gin of planta; A7 with SV3 absent; A8 with SD1 hairlike; one SV seta present; SV1 nearly in verticle line with L3 and V1; SV1 and L3 about equal in length, V1 short (not illustrated); spiracle slightly larger than pro- thoracic and other abdominal spiracles; A9 with SD2 absent; L3 ventral to L2; SV1 VOLUME 101, NUMBER 1 caudal to D2; V1 short (not illustrated); A10 (Figs. 14-16, 20): D2, SD1, and SD2 about four times length of D1; crochets uni- serial and triordinal. Holotype.— °, ““Colombia: Instituto Col- ombiano Agropecuario, Experiment Station ‘‘Palmira,’’ Cauca Valley, 1 March—15 March 1991, Ex. Sugar cane, Coll. Lucero Cardenas Duque, Emerged 21 April—1 May 1991.”’ The holotype is not dissected and is deposited in USNM. Paratypes.—3 ¢, Same data as holotype. Paratypes are not dissected and are depos- ited in USNM. Other specimens examined.—COLOM- BIA: 2 6,7 2, “Miranda (Val.), en. cafia a [zucar], Jul[y] 1984, L[ucero] Cardenas”’; 1 36,1 2 from Vitor Becker Collection [yel- low label]; 1 3, 5 2, ‘“‘Miranda, VI-28-— [19]84, L[ucero] Cardenas,” ‘“‘Tallos cafia de azucar,’ “? Wing Slide by DA 3349, USNM 81585,” [green label],’’ ““2 Geni- talia Slide by D. Adamski 2885, USNM 81422,” [green label]; 3 2, “‘Riopaila, Par- asita Diatraea,”’ ‘“I-6—[19]65, 723-4,” “‘II- 1-[19]65, 9651-28,” “‘II-15[19]65, 446— lerling. lao. “Ex. sugar cane; Ag: Exp. Sta. Palmira, Valle, Let: Oct. 3, 1941, B. Losada S,”’ “2 Genitalia Slide by R. B. Selander, USNM 11, 157,” [green label]; 2 2, “Valle Ingenio del Cauca, H: cafia de azticar, Barrenador, Dic/[19]82, D-83,” ““@ Genitalia Slide by D. Adamski 2849, USNM 81393,” [green label], ‘‘ 2 Genitalia Slide by D. Adamski 2850, USNM 81394,” [green label]; 2 6, 1 ¢, ‘‘Valle Ing[enio] del Cauca, H: cana de azucar Col: L[ucero] Cardenas y Y.P. Chacon, II-[19]83, D-83,” “d Genitalia Slide by D. Adamski 2847, USNM 81391,” [green label], “‘d Genitalia Slide by D. Adamski 2846, USNM 81390,” [green label], ““2 Genitalia Slide by D. Adamski 2848, USNM 81392,” [green la- bell? VENEZUELA: 2°, °“Tachira, El Cobre, 12,00 m[e]t[e]rs, May 1947,” and melachirass Wa) Grita;, 1,450: mileltlelrs, 14.V.1949,”” ‘“‘Reared from larva in Sugar cane,” “‘2 Genitalia Slide by R. B. Selan- der, USNM 11160,” [green label], ‘‘ 2 Gen- 171 italia Slide by R. B. Selander, USNM 11161" [green label]; 2 ¢d, 1 2, “Maracay, 450 mfejt[e]rs, 28.11.1949, H.E: Box’’, “February 1951,” “‘June 1948,” ‘‘Reared from larva in Sugar cane,” ‘‘d Genitalia Slide by D. Adamski 3038, USNM 81488” [green label], ““d Wing Slide by J. G. Clarke, USNM 11209” [green label], “dé Genitalia Slide by J. G. Clarke, USNM 11209”’ [green label], ““2 Genitalia Slide by R. B. Selander, USNM 11164” [green label]; 1 2, “‘Carabobo, Cent. Tacarigua, 450 m[e]t[e]rs, September 1947, H.E. Box,”’ “‘Reared from larva in Sugar cane,” ““® Genitalia Slide by R. B. Selander, USNM 11159 [green label], 1 6; 3° 2, ‘“Yaracuy, Chivacoa, 230 m[e]t[e]rs, Feb- ruanye, 19505 es 42S 195052 oSan Pa[illegible], 400 m[e]t[e]rs, 22.11.1949, H.E. Box,” “‘Reared from larva in Sugar cane,” ‘“‘d Genitalia Slide by D. Adamski 3037, USNM 81487,” [green label], “‘? Genitalia Slide by R. B. Selander, USNM 11163” [green label], ‘“‘Reared from larva in Sugar cane,”’ “‘? Genitalia Slide by J. G. Clarke, USNM 11210” [green label], ‘“‘Reared from larva in Sugar cane,” “@ Genitalia Slide by R. B. Selander, USNM 11162,” [green label], ““Reared from larva in Sugar cane’’; 1 d, “‘Merida, nr. Egido, 1,500 mf[e]t[e]rs, 8.VI.1949, H. E. Box,” ‘*$ Genitalia Slide by D. Adamski 3039, USNM 81489” [green label]; 1 2, ‘‘Miran- da, Sta. Lucia, 180 mfe]t[e]rs, 5.111.1948, HE: Box,” “2 “Genitalia Shide by. J. 1G: Clarke, USNM 11207’ [green label], ‘‘Reared from larva in Sugar cane’; 1 2, ‘‘Aragua, El Conseja, 550 m[e]t[e]rs, March 1951, H.E. Box,” “‘Reared from lar- va in Sugar cane,” ““? Genitalia Slide by R. B. Selander, USNM 11158” [green la- bel]; 1 do, ‘“‘El Limon, nr, Maracay, 460 m[e]t[e]rs, 31.i1i.1950, H.E. Box,” ““Reared from larva in Ciox lachryma-jobi [L]”; 1 ?, “Zulia, Perijo, Mts. close to Colombia, Dec. 1950, E Fernandez Yepoz,”’ “‘Reared from larva in Setaria paniculifera |Four- nier],’’ ““2 Genitalia Slide by J. EK Clarke, USNM 11208” [green label]; 1 2, ‘““Vene- 172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON zuela, Turbio Valley, nr. Barquisimeto, 1956, P. Guagliumi, Larva boring sugar- cane; COM. INSEE ENTE COLE NO: 15211, Press[ented] by Com. Inst. Ent., BM 1957-256.” Fourteen additional adult spec- imens were examined at The Natural His- tory Museum, London, with above label data. Larvae studied were collected and preserved in alcohol with the following data, ‘““Colombia: Instituto Colombiano Agropecuario, Experiment Station ‘*Palmi- ra,” Cauca Valley, 15 December—20 Janu- ary 1990, Ex. Sugar cane, Coll. Lucero Car- denas Duque.”’ All larval specimens are de- posited in the USNM alcohol collection. Types examined.—Lectotype designated by Clarke, 6, Blastobasis obstricta Mey- rick, ‘““Lectotype’’ [round label], “‘Bartica, Brit[ish] Guiana, Parish 1.13,” “‘Lectotype, Auximobasis obstricta Meyrick, J.EC.C. 1948,” ‘““d Genitalia on Slide 5-X-1948, J.EG.C. 8078,” Auximobasis obstricta Meyr., E. Meyrick det., in Meyrick Coll. 211.) “Sobstricta Meyr;’, “Meyrick: Goll, BM 1938-290,”’ [Natural History Museum, London, England]. Lectotype, 6, Blasto- basis subolivacea Walsingham, ‘‘S[aint] Thomas, 9.IV.[18]94” [hand-written pink label], ““Blastobasis subolivacea 125.2089 WLSM, <6, TYPE” [hand-written label], ‘““$ Genitalia Slide by D. Adamski, 3470” [green label], “Holotype, Blastobasis su- bolivacea Wlism, <6,” [“‘Grigore Antipa”’ National Museum of Natural History, Bu- charest, Romania]. Etomology.—Blastobasis graminea is named after the plant family Gramineae be- cause larvae feed on several grass hosts. DISCUSSION Auximobasis obstricta Meyrick 1918, is transferred to Blastobasis Zeller, 1855, and Blastobasis subolivacea Walsingham, 1897, is transferred to Holcocera Clemens 1863 (new combinations). Blastobasis graminea is probably more closely related to Blastobasis obstricta Meyrick, 1918, n. comb. than to any other described Blastobasis. Both species differ markedly in wing pattern and in several male and female genitalic features. How- ever, males of both species share an uncus with a widened base, a bidentate gnathos, and stout marginal setae along the outer margin of the proximal flange. Females share a widened ostium. Martorell (1976) reported two species of Blastobasidae feeding within sorghum heads in the Vieques Islands east of Puerto Rico, but I have not seen any specimens to substantiate this. BIOLOGY Cardenas and Hernandez (1985) de- scribed the biology of Blastobasis grami- nea on sugar cane in Colombia; these find- ings are summarized below. The most se- vere damage by B. graminea occurs within the terminal third of the sugar cane plant, however, damage can occur in lower re- gions as well. When the damage is extreme the apical portion of the plant dies. Early instars of B. graminea feed on the surface tissue layers. When the larva is able, it bores into the stem. Galleries are usually irregularly shaped (Figs. 22—24); the larvae never excavate more than two internodes of the plant. Mature larvae usu- ally pupate between the stem and the sheath (Fig. 21), but sometimes within the stem (Fig. 22). In addition to sugar cane, larvae feed on corn (Figs. 25, 26), sorghum, Coix lacryma-jobi L., and Setaria paniculifera Fournier. There appears to be a strong correlation (Ratio of 8:1) between the presence of Dia- traea saccharalis and Blastobasis grami- nea, however, it is not known which moth species attacks the plant first. ACKNOWLEDGMENTS I thank Ingeborg Zenner-Polania, former director, Programa de Entomologia, Insti- tuto Colombiano Agropécuario, Bogota, Colombia, for the coordination of activities related to the acquisition of specimens of Blastobasis graminea; Lucero Cardenas Duque, of the above institution, for live and VOLUME 101, NUMBER | 173 Figs. 21-26. within sheath of sugar cane (see arrow). 23, Larvae and damage within sugar cane stem (see arrow). 24, Larval pho) Larval damage of Blastobasis graminea. 21, Pupal site on sugar cane (see arrow). 22, Pupa gallery within sugar cane stem (see arrow). 25, Larval damage in corn stem (see arrow). 26, Larva within corn & 7 o S stem (see arrow). preserved larvae, and photographic prints of larval damage; Klaus Sattler, Michael Shaffer and Kevin Tuck, of the Natural History Museum, London, England, for their help with examination and photog- raphy of type specimens; Dorel Rusti, “Grigore Antipa’’ National Museum of Natural History, Bucharest, Romania, for the loan of the lectotype of Blastobasis su- bolivacea Walsingham; Greta Tyson and Michael Sullivan, of the Electron Micro- scope Center, Mississippi State University, for their help with the preparation of the specimens and photographic plates; Carl Hansen of the Office of Imaging, Printing and Photographic Services for the photo- graph of the holotype; the late John EF Gates Clarke, Smithsonian Institution, for referring this research problem to me. This research was supported in part by NSF Grant BSR85-01212 and a grant from Sig- ma Xi. 174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Adamski, D. and R. L. Brown. 1987. A New Nearctic Glyphidocera with descriptions of all stages (Lep- idoptera: Blastobasidae: Symmocinae). Proceed- ings of the Entomological Society of Washington 89(2): 329-343. . 1989. Morphology and Systematics of North American Blastobasidae (Lepidoptera: Gelechioi- dea). Mississippi Agricultural Forest Experiment Station Technical Bulletin 165. Mississippi Ento- mological Museum Publication No. 1, 70 pp. Box, H. E. 1953. List of Sugar-Cane Insects: A Syn- onymic Catalogue of the Sugar-cane Insects and Mites of the World, and of their Insect Parasites and Predators, Arranged Systematically. Com- monwealth Institute of Entomology, London, 100 Pp- Cardenas Duque, L. and Miria del Pilar Hernandez. 1985. Barrenador de la cana de azucar en Colom- bia. Miscelanea, Sociedad Colombiana de Ento- mologia 1: 12-17. Clarke, J. EF G. 1941. The preparation of slides of the genitalia of Lepidoptera. Bulletin of the Brooklyn Entomological Society 36: 149-161. Clemens, B. 1863. American micro-lepidoptera. Pro- ceedings of the Entomological Society of Phila- delphia 2: 119-129. Guagliumi, P. 1962. Las plagas de cana de aztcar en Venezuela. Ministerio de Agricultura y Cria. Ma- racay, Venezuela. Monografia no. 2. 2 partes. 789 PP: Hodges, R. W. In press. Gelechioidea. /n Kristensen, N. P., ed., Handbuch der Zoologie. Kornerup, A. and J. H. Wanschner. 1978. Methuen Handbook of Colour. Third Ed. Methuen and Co., Ltd., London. 252 pp. Martorell, L. EF 1976. Annotated Food Plant Catalogue of the Insects of Puerto Rico. Agricultural Exper- iment Station, University of Puerto Rico, Depart- ment of Entomology. 303 pp. Meyrick, E. 1894. On a collection of Lepidoptera from upper Burma. Transactions of the Entomological Society of London 1894: 1-29. . 1918. Exotic Microlepidoptera. 2. pp. 385-— 480 (1916-23) Taylor and Francis, London. Re- printed by C. W. Classey, Ltd., 1969. 640 pp. Schoonhoven L. M. and V. G. Dethier. 1966. Sensory aspects of host-plant discrimination of lepidopter- ous larvae. Archives Neerlandaises de Zoologie 16(4): 497-530. Walsingham, Lord (Thomas de Grey). 1897. Revision of the West Indian micro-Lepidoptera with de- scriptions of new species. Proceedings of the Zoo- logical Society of London 1897: 54-183. Zeller, P. C. 1855. Die Arten der Gattung Butalis be- schreiben. Linnaea Entomologica 10: [169]—269. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 175-196 A REVISION OF THE FLAVIDUS GROUP OF THE GENUS CHRYSOPS MEIGEN (DIPTERA: TABANIDAE) TeEvIsS M. BAIER Department of Zoology, University of New Hampshire, Durham, NH 03824, U.S.A. (e-mail: tbaier @jlc.net) Abstract.—A taxonomic revision of the Chrysops flavidus species group is provided with a key to species, descriptions, distribution maps, and illustrations of wings and an- tennae for each of the nine species currently recognized in this group. The relationships and identification of these nine species are based on analysis of morphological similarity involving 13 critical characters. A total of 4,843 specimens from 14 institutions were examined during this study. Two new species in the Chrysops flavidus group are described, Chrysops sandyi and Chrysops tumidicornis, both previously misidentified as Chrysops atlanticus. The male of Chrysops dixianus is described for the first time. Key Words: The genus Chrysops Meigen is currently represented by 87 described species in the Nearctic Region north of Mexico (Burger 1995), and 291 world wide. This genus was last revised in North America by Philip (1955). Philip’s revision included a key to 95 species and 14 subspecies, with corre- sponding notes on these taxa, and descrip- tions of 4 new species and 5 new subspe- cies (but it also treated species from Central America and the Caribbean). Although no one has divided all of the Nearctic Chrysops into definitive species groups, some species groups have been rec- ognized by tabanid taxonomists. The three commonly used are the Chrysops callidus group, the C. carbonarius group, and the C. flavidus group (Pechuman 1949). The C. flavidus group is particularly difficult be- cause of the large amount of intraspecific variation within the included taxa. Al- though three species have been described in this group during the past 45 years, no one has studied the group as a whole. The Chrysops flavidus group currently Diptera, Tabanidae, Chrysops flavidus group, key to species includes 7 previoously described species and two new species: Chrysops atlanticus Pechuman, C. brunneus Hine, C. celatus Pechuman, C. dixianus Pechuman, C. flav- idus Wiedemann, C. pudicus Osten Sacken, and C. reicherti Fairchild, C. sandyi, n.sp., and C. tumidicornis, n. sp. I will redescribe, clarify relationships and summarize the bi- ology and geographic distribution of this group. Immature stages and habitat are dis- cussed under the description of each spe- cies. MATERIALS AND METHODS I examined 4,843 specimens, (4,684 fe- males and 159 males), from the collections listed below. The acronyms are those of Ar- nett et al. (1993), except for the University of New Hampshire Collection (UNHC). AMNH: Department of Entomology, American Museum of Natural History, New York, NY; David A. Grimaldi. BMNH: Department of Entomology, The 176 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Natural History Museum, Lon- don, U. K.; John E. Chainey. Department of Entomology, Cali- fornia Academy of Sciences, San Francisco; Paul H. Arnaud, Jr. Canadian National Collection, Centre for Land and Biological Resources Research, Agriculture Canada, Ottawa, Ontario; J. M. Cumming. Department of Entomology, Clemson University, Clemson, SC; Michael A. Floyd. Department of Entomology, Cor- nell University, Ithaca, NY; E. Ri- chard Hoebeke. Division of Insects, Field Museum of Natural History, Chicago, IL; Alfred E Newton, Jr. Florida State Collection of Ar- thropods, Division of Plant Indus- try, Gainesville; G. B. Fairchild. Illinois Natural History Survey Insect Collection, Champaign; Kathleen R. Methven. Ohio State University, Collection of Insects and Spiders, Columbus; Andrey Sharkov. Department of Entomology Insect Collection, Texas A. & M. Uni- versity, College Station; Edward G. Riley. Entomology Collection, Universi- ty of Georgia, Athens; Cecil L. Smith. Entomological Museum, Depart- ment of Zoology, University of New Hampshire, Durham; Donald S. Chandler. National Museum of Natural His- tory, Smithsonian Institution, Washington, DC; R. V. Peterson. CASC: CNEI: CUCE: CUIC: FMNH: FSCA: INHS: OSUC: TAMU: UGCA: UNHC: USNM: The following insect collections are also cited in this paper: MNHN: National Collection of Insects, Muséum National d’Histoire Na- turelle, 45, Rue Buffon, Paris 75005 France. MRSN: Museo Regionale Scienze Natur- ali, Via Gioletti 36, Torino 10128, Italy. MZLU: Museum of Zoology, Lund Uni- versity, Helgonavagen 3, S-223 62 Lund, Sweden. ZMHB: Museum ftir Naturkunde der Humboldt Universitat zu Berlin, Bereich Zoologischer Museum, InvalidenstraBe 43, 1040 Berlin, Germany. The Chrysops flavidus group has few dis- tinctive structural characters, and the spe- cies can be difficult to identify. Color pat- terns are important in identification but must be used with caution because of var- iation. The characters described below are the most useful in separating the species. The morphological terminology used in this paper follows that used by Teskey (1990). Principal characteristics used are: body length, in millimeters, from the base of the antennae to the apex of the abdomen; the size and shape of the scape and pedicel as well as the length ratio and color of the scape, pedicel, and flagellum; shape of the frontal callus; color of the mesoscutum; wing pattern, particularly the width of the apical spot, the location of the outer margin of the crossband, and the infuscation of the 5th posterior cell; color of the hind femur; and abdominal color pattern. For illustrations, I chose a specimen that most closely resembled the description of the holotype. One wing from each specimen was removed, placed between two 2 X 2 glass slides and scanned into a Dell 486p/ 25 computer using MICROTEK Scan- Maker 35t. The scanned picture was printed and used as a template for the final drawing. The antennal drawings were produced with a camera lucida. The locality data from all specimens ex- amined were entered into a database using Wordtech Systems, dBXL®, (a dBASE III Plus® compatible format). The information collected was placed into 9 “‘fields’’: spe- cies, country, state, county, town, collector, VOLUME 101, NUMBER 1 aS aa eater OES GAN flagellum pedicel scape basal callus 2 antennal socket Figs. 1-3. Chrysops species. museum, count (number of specimens with identical data), and sex (male or female). Locality data for specimens I did not ex- amine were taken from the distribution re- cords of L. L. Pechuman, now stored in the Zoology Department at the University of New Hampshire. Distribution maps were then produced for each species from the above database and from the records of L. L. Pechuman. DIAGNOSIS OF THE CHRYSOPS FLAVIDUS GROUP Frontoclypeus glossy yellow with no mid-facial stripe; frontal callus yellow to light brown, occasionally with brown upper margin; frons yellow pollinose; mesoscu- tum with 3 longitudinal stripes, darker than ground color; proepimeron and proepister- nal callosity yellow tomentose with long yellow hair; wing pattern with broad apical spot entering 2nd submarginal cell, cross- band reaching hind margin of wing, hyaline Wa hyaline triangle crossband 1, Wing of Chrysops species. 2, Antenna of Chrysops species. 3, Anterior view of head of triangle not crossing vein R,,;, cell br at least % infuscated basally; abdominal ter- gites 2—4 patterned with median inverted ““V°’ set over pale median triangle. KEY TO SPECIES OF THE CHRYSOPS FLAVIDUS GROUP 1. Scape and pedicel distinctly swollen, together longer than flagellum (Fig. 14). Frontal callus twice as wide as high, with no black markings. Abdomen with inconspicuous median trian- gles. Wing pattern with extensive apical spot, reaching crossband posteriorly and enclosing hyaline crescent (Fig. 23) brunneus Hine — Scape and pedicel less swollen, combined length equal to or shorter than flagellum. With- out remaining combination of characters .... 2 NO . Usual hyaline areas of wing slightly smoky to very smoky, outer margin of crossband angled from vein R, to middle of vein M, (Fig. 22). Scape and pedicel moderately swollen, com- bined length equal to or rarely longer than fla- gellum (Fig. 13). Mesoscutum green-gray iri- descent with black longitudinal stripes oe RE aa ott LS od BU S38 atlanticus Pechuman — Without above combination of characters ... 3 178 . Mesoscutum greenish gray or steel gray in ground color, longitudinal stripes black. Outer margin of crossband sinuous ............ 4 Mesoscutum yellow in ground color, longitu- dinal stripes brown. Outer margin of cross- band, straight, concave, or sinuous ......... 6 . Antenna not swollen, flagellum at least 4% lon- ger than scape and pedicel combined (Fig. 15). Frontal callus spade shaped, width and height subequal. Without dark spot under scutellum BRD 5 At Rane Giese cee celatus Pechuman Scape and pedicel slightly swollen to swollen, combined length subequal to flagellum. Frontal callus, width and height not equal. Abdomen with dark spot under scutellum, often extend- ing to lateral edges of tergite 1 . Hyaline triangle not reaching vein R,,,, cells br and bm % and ¥% infuscated basally (Fig. 30). Scape and pedicel swollen, pedicel barrel shaped (Fig. 21). Frontal callus light brown. Mesoscutum gray green with black stripes MALS era Ce RARCe toe: Ae tumidicornis, Nn. sp. Hyaline triangle reaches vein R,,3;, apical spot narrowly enters 2nd submarginal cell, cells br and bm '% and infuscated basally (Fig. 27). Scape and pedicel slightly swollen (Fig.18). Frontal callus reddish brown with black upper margin. Mesoscutum steel gray with black SUHPeS erie eee oe pudicus Osten Sacken . Abdominal tergites 1—2 light yellow to straw color, dark inverted “‘V” on tergite 2 com- pressed, not reaching beyond the middle of ter- gite 2. Flagellum 1.3 to 1.5 times longer than scape and pedicel combined Abdominal tergites 1—2 yellow, orange or light brown, inverted “‘V” pattern not compressed, extending beyond the middle of tergite 2 to- ward anterior margin. Flagellum equal to or up to 1.2 times longer than scape and pedicel Combined ware a cer heey Reese: see eye 8 . Fifth posterior cell infuscated, outer margin of crossband sinuous, cells br and bm % and % infuscated basally (Fig. 28). Hind femur red- dish-brown. Tergites 1—2 straw-colored, medi- an anterior portion of tergite 2 with a greenish cast. Length 8-9.5 mm ..... Fifth posterior cell hyaline, vein Cu, infuscated at border, outer margin of crossband straight or convex, cells br and bm % and ¥, infuscated basally (Fig.25). Hind femur dark brown. Ter- gites 1-2 reicherti Fairchild yellow without a greenish cast. Length 6-8.5 mm dixianus Pechuman . Hind femur yellow. Mesoscutum yellow with brown stripes. Outer margin of crossband straight or concave, cells br and bm % and 4% infuscated basally (Fig. 26). Scape and pedicel slightly swollen (Fig. 17). Length 8-10 mm Blog a Sodas teams hen Sect ra eee ee: flavidus Wiedemann PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON — Hind femur dark brown basally. Mesoscutum yellow green with brown stripes. Outer margin of crossband straight, hyaline triangle slightly tinted, cells br and bm %—% and Y, infuscated basally (Fig. 29). Scape and pedicel not swol- len (Fig. 20). Length 5.7—7.5 mm Ss SN EW sc end ebor i ar ee taet te wR een sandyi, n. sp. Chrysops atlanticus Pechuman (Fig. 4, 13, 22) Chrysops canifrons Walker 1848: 197-198. Holotype female: Florida (BMNH). Se- nior synonym of atlanticus, suppressed by the International Commission on Zoo- logical Nomenclature, Opinion No. 1711, 1993. Chrysops atlanticus Pechuman 1949: 79— 82. Holotype female: Rehoboth Beach, Delaware (CUIC); Jamback and Wall 1959: 23-24, description of egg, larva and pupa; Teskey 1969: 29-30, descrip- tion of larva; Goodwin 1972: 104, de- scription of pupa. Diagnosis.—Length 6.5—9.2 mm. Scape and pedicel moderately swollen, flagellum equal to length of scape and pedicel com- bined. Mesoscutum grayish green in ground color. Wing pattern with smoky tinge, hy- aline areas not clearly defined, hyaline tri- angle not extending beyond lower half of Ist submarginal cell. Female.—Light to dark brown, length 8.5—9.2 mm. Scape and pedicel moderately swollen, light brown, basal flagellomere light brown, apical flagellomeres black, length of flagellum subequal to combined length of scape and pedicel, antennal ratio 15:10:25. Frontoclypeus glossy yellow, eye margins bearing yellow tomentum that ex- tends medially beneath tentorial pits to frontoclypeal suture. Maxillary palp brown with sparse black hairs. Frons yellow and gray tomentose with yellow hairs at vertex, width 1.13 times height. Frontal callus ob- long, bulbous, light brown, width 1.54 times height. Vertex with glossy black in- tegument surrounding shining brown ocelli. Mesoscutum grey green iridescent with three black longitudinal stripes, sublaterals VOLUME 101, NUMBER 1 179 Figs. 4-9. Distributions. 4, Chrysops atlanticus. 5, C. brunneus. 6, C. celatus. 7, C. dixianus. 8, C. flavidus. 9, C. pudicus. 180 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON wider than median one, scutellum predom- inantly brown, lighter medianly. Legs pre- dominantly yellow, hind coxa and fore ta- rusi brown, fore tibia with black hair, mid and hind tibia with mixed yellow and black hair. Wing pattern with usual hyaline areas smoky, apical spot broad, fading into 2nd submarginal cell, crossband brown with ir- regular outer margin, crossing Ist posterior cell at its center, a perpendicular line drawn from vein R, to hind margin of wing not forming line parallel to outer margin of crossband, hyaline triangle not extending beyond posterior % of Ist submarginal cell, discal cell, 4th and 5th posterior cells in- fuscated, 5th posterior cell paler in center, cells br and bm % and % basally infuscated. Halter brown stalk yellow-brown knob. First abdominal tergite dark yellow, black haired medianly, tergites 2—5 with subme- dian oblique dark spots, gray posterior mar- gin extending medianly into pale orange, equilateral triangle, dark spots on tergite 2 outline triangle and join anteriorly forming inverted “‘V”’ pattern, tergites 3—5 with sub- median dark spots slightly separated ante- riorly. Sternites 1-2 yellow, sternite 3 mot- tled yellow and brown, sternites 4—7 dark brown. Male.—Similar to female except for usu- al sexual differences and following char- acteristics. Length 6.5—8 mm. Scape and pedicel slightly more enlarged with black hair that is longer and finer than on female. Mesoscutum sparsely covered with long yellow hair. Cell br entirely infuscated ex- cept for a subapical hyaline spot. Abdomen with yellow and black hair scattered over pattern. Material examined.—934 2 and 18 ¢ examined from the following collections: GASE,..,CNCI_ CUCE. CUICCEMNET, FSCA, INHS, OSUC, TAMU, UGCA, UNHC, USNM. Distribution.—Atlantic coast of North America from Maine to Florida. I have ex- amined specimens from AL, DE, FL, GA, LA, MA, ME, MD; MS, NC, NH, NJ, NY, SC, TN, and VA. There are also published records for CT and RI. Biology.—The larvae have been found in salt marsh and brackish pools. Based on their narrow distribution along the coast, they are probably restricted to this saline environment. Flight times for C. atlanticus are between April and September in Loui- siana (Tidwell 1973), June to September in Virginia (Pechuman 1973), and late May to mid October in New York (Pechuman 1981). The adults are abundant during June and July throughout their range and are eco- nomically important due to their aggressive biting behavior. Thirty C. atlanticus bites were counted on 1 human in 90 seconds in a cultivated field near Cedarville, New Jer- sey, (Hansens, 1980). Anderson (1971) de- scribed C. atlanticus as being autogenous, depositing the first egg mass before seeking a blood meal. Subsequent work by Mag- narelli and Anderson (1976) concluded that populations of C. atlanticus can be main- tained by the first oviposition, making them difficult to control. Chrysops atlanticus is not known to transmit any disease agent in nature; how- ever, in the laboratory, tests have demon- strated it to be an effective experimental vector of the African filarial worm Loa loa (Orihel and Lowrie 1975). Discussion.—Chrysops atlanticus 1s most similar to C. sandyi, C. tumidicornis, and to C. brunneus, based on the smoky wing pattern, association with a saline en- vironment, and swollen antennae. Chrysops atlanticus differs from C. brunneus in its narrower scape and pedicel (Fig. 22) that when combined are equal in length to the flagellum, moderately inflated frontal cal- losity, grayish-green mesoscutum, and wing pattern without clearly defined hyaline ar- eas. Chrysops atlanticus differs from C. sandyi in its larger size, blackish longitu- dinal stripes on the mesoscutum, and cross- band with irregular outer margin. Chrysops atlanticus differs from C. tumidicornis in its larger size, less swollen pedicel, smokier wing pattern, and lighter abdominal pattern VOLUME 101, NUMBER 1 that does not have a dark spot on tergite | under the scutellum. There appears to be no overlap in the geographical distribution of C. atlanticus with either C. sandyi or C. tumidicornis. Color variation in C. atlanticus is com- mon throughout its range, making it partic- ularly difficult to define. The abdominal pattern can vary from tergites 2—6 having a pale inverted ““V”’ formed by hair, to a dark, well defined, integumental inverted ““V”’ pattern that includes *% or more of the tergite. In all these cases, the characteristic wing pattern described above and the in- flated scape and pedicel will define C. at- lanticus. Chrysops brunneus Hine (Figs. 5; 14, 23) Chrysops brunneus Hine 1903: 34. Syn- types female and male: Sandusky, Ohio, 2? 6 (OSUC), 1 2 (BMNH):; Goodwin 1976: 343, description of larva and pupa. Diagnosis.—Robust species, length 8—10 mm. Antenna with scape and pedicel swol- len, flagellum distinctly shorter than com- bined length of scape and pedicel. Frontal callus strongly inflated, width twice height. Wing with apical spot extensive, reaching around margin of wing to crossband and enclosing hyaline crescent. Cells br and bm % and ¥%2 infuscated basally. Abdominal pat- tern indistinct. Female.—Yellowish brown. Scape and pedicel distinctly swollen, scape twice as long as broad, light brown, black setose, basal flagellomere yellow basally, dark brown apically, apical flagellomeres black, flagellum distinctly shorter than scape and pedicel combined, antennal ratio 18:12:25. Frontoclypeus glossy yellow, eye margins bearing yellow tomentum that extends me- dianly beneath tentorial pits to frontocly- peal suture. Maxillary palp light brown with sparse black hairs. Frons yellow to- mentose with scattered yellow hairs, con- vergent above, height 1.3 times width at base. Frontal callus elliptical, distinctly nar- 181 rowed laterally, strongly inflated, light brown, upper margin dark brown, width twice height. Vertex dark brown tomentose, except glossy around each ocellus, ocelli dark brown. Mesoscutum light brown to- mentose and yellow pilose, except lateral margins pale yellow-gray tomentose, bear- ing 3 longitudinal stripes, median stripe very narrow, dark brown, sublateral stripes brown and broad. Scutellum dark brown basally, brown apically, yellow pilose. Pleuron yellow-gray tomentose. Fore coxa light brown tomentose, mid and hind coxa dark brown tomentose, femur, tibia and tar- sus light brown, femur and tibia bearing yellow hairs, except apices of tibia bearing mixed yellow and black hairs, tarsus bear- ing black hairs. Wing pattern distinct, apical spot broad, extending around wing margin to crossband, leaving narrow hyaline band along outer margin of crossband that does not reach vein R,,;, outer margin of cross- band straight or sinuous, 5th posterior cell infuscated, cells br and bm % and ¥% their length respectively. Halter with dark brown stalk and light brown knob. Abdomen uni- formly brown, lacking dark integumental markings, tergite 1 with black hair medi- anly, yellow hair laterally, tergites 2—4 pre- dominantly black-haired, except for yellow hairs on posterior margin that expand me- dianly into indistinct yellow-haired trian- gles, tergites 5—7 with mixed black and yel- low hairs forming no distinct pattern. Ster- nites 1—5 light brown, sternites 3—5 with progressively larger median and lateral brown integumental spots, 6—7 dark brown, all sternites with mixed black and yellow hairs. Male.—Resembles female except for the usual sexual differences and the following characteristics. Length 8.2-10 mm, scape and pedicel slightly more inflated than in female, with longer, finer black hair, fron- toclypeus with yellow tomentose triangle at vertex, cheeks yellow pollinose. Long yel- low hairs scattered over mesoscutum, scu- tellum, and abdomen. Material examined.—S02 @ and 14 6 182 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 10-12. were examined from the following collec- tions: AMNH: .-CASC.> ENEL,,.CUECC, CUIC, \BMNE: FSCA... INHS;,OSUC, TAMU, UGCA, UNHC, USNM. Distribution.—New York to Florida, west to Texas, and north to Michigan and southern Ontario. It is not found in the Ap- palachian Highlands and surrounding ter- rain. I have examined specimens from: AL, AR... DE, FL, GA, IL, INKS; KY,1LA;,MD, MI,. MS, -NJ; NY.. OH,..OK,. PAs SG,.TN, TX, and VA. It is also known from IA, MO, NC, and Ontario, Canada. Biology.—Chrysops brunneus is associ- ated with both fresh and salt water habitats. Goodwin (1976) reared a larva of this spe- cies collected from a salt marsh in Mc- Clellanville, SC. Adults are commonly Distributions. 10, Chrysops reicherti. 11, C. sandyi. 12, C. tumidicornis. found in marshes near lakes such as Lake Michigan, Lake Erie and Lake Ontario. Discussion.—Chrysops brunneus is very distinctive and can be separated from other species of the C. flavidus group by its strongly inflated scape and pedicel that combined are longer than the flagellum, and by its reduced hyaline triangle appearing as a crescent-shaped area between the cross- band and broad apical spot. The smoky wing pattern of C. brunneus and its pres- ence in a saline environment are similar to that of C. atlanticus, but is easily separated by those characters noted above. The only variation observed in this spe- cies was the abdominal pattern that has ter- gites 2—3 with small, median, black dashes on the integument in approximately 5% of VOLUME 101, NUMBER 1 183 Gee oi Ge atlanticus 13 dixianus 16 reicherti 19 Figs. 13-21. brunneus 14 flavidus Ay] sanayi 20 Female antennae. 13, Chrysops atlanticus. 14, C. brunneus. 15, C. celatus. 16, C. dixianus. ce/atus iS) oS Ogee tumidicornis pa 17, C. flavidus. 18, C. pudicus. 19, C. reicherti. 20, C. sandyi. 21, C. tumidicornis. the specimens examined. Ordinarily the ab- dominal pattern is indistinct with only some dark shadowing from dark hairs. Chrysops celatus Pechuman (Figs. 6, 15, 24) Chrysops flavidus celatus Pechuman 1949: 82-83. Holotype female: Medford Lakes, New Jersey (CUIC); Pechuman 1957: 30, description of male. Chrysops celatus: Teskey 1969: 34, de- scription of larva and pupa. Diagnosis.—Black and yellow, length 8— 9.8 mm. Scape and pedicel not swollen, fla- gellum % longer than scape and pedicel combined. Frontal callus spade shaped, width and height subequal. Mesoscutum greenish gray iridescent with black longi- tudinal stripes. Cells br and bm % and /¥, infuscated basally. Female——Scape and pedicel not swollen, light brown, black setose, basal flagellomere light brown basally, black apically, apical fla- gellomeres black, 1.3 times length of basal flagellomere, antennal ratio 15:10:34. Fron- toclypeus glossy yellow, eye margins bearing yellow tomentum that extends medianly be- neath tentorial pits to frontoclypeal suture. Maxillary palp and labrum yellow. Frons pre- dominantly yellow-gray tomentose with scat- tered yellow hair, slightly convergent above, height 1.5 times width at base. Frontal callus spade shaped, bulbous, yellow brown, width and height subequal. Vertex gray tomentose, except glossy brown integument associated with each ocellus which extends narrowly to upper margin of frontal callus. Mesoscutum greenish-gray iridescent, except lateral mar- gins yellow tomentose, bearing 3 black lon- gitudinal stripes, the median one narrow and flanked with gray. Scutellum black ba- sally. Pleuron yellow tomentose. Legs mostly yellow, fore leg with tarsus and api- cal portion of tibia dark brown with black hair, mid and hind coxa brown, tarsus with black hairs. Wing pattern with broad apical spot entering apical half of 2nd submarginal cell, apex of hyaline triangle not reaching vein R,,, outer margin of brown crossband sinuous, 5th posterior cell infuscated basal- ly, hyaline apically, cells br and bm % and Y. infuscated basally. Halter brown. Tergites 1-2 yellow brown, tergite 2 with darker median integumental marking appearing as an inverted “‘V”’’ set over median yellow- 184 brown triangle, tergites 3—4 predominantly brown except for lighter posterior margin that expands medianly into light brown pol- linose triangle, remaining tergites with an- terior % brown, posterior 4% yellow-brown. Sternites yellow-brown pollinose with ster- nites 3—7 bearing narrow yellow pollinose line along the posterior margin, sternites 5— 7 predominantly brown. Male.—Similar to female except for the usual sexual differences and following characters. Length 6.5—7.5 mm. Pedicel only slightly shorter than scape, with finer, longer black hairs, length of basal flagel- lomere and combined apical flagellomeres subequal. Apical spot reaches posterior half of 2nd submarginal cell and then fades along hind margin, cells br and bm %4 and Y infuscated basally. First tergite with dark spot under scutellum or entirely dark. Fifty percent of males examined with abdominal pattern of yellow and black with a black scutellum. Material examined.—574 2, 30 6, ex- amined from the following museums: AMINE: CAS GC.) CNCke CUCE, "CuIC, FSCA, INHS, TAMU, UGCA, UNHC, USNM. Distribution.—Massachusetts south to Florida, west to Texas; disjunct population in Michigan, southern Ontario, and north- ern Ohio, Illinois, and Indiana. I have ex- amined specimens from the following States) AE; -GE DE: BE, GAIN, KYLA, MA, MD, MI, MS, NC, NH, NJ, NY, SC, TN, TX, and VA. L. L. Pechuman has re- corded specimens from WV, and Ontario, Canada. Biology.—Teskey (1969), based his de- scription of the larva on 35 specimens. They were collected from 3 sites with fresh, standing water: wet sand, organic soil, and around roots of aquatic vegetation. In 1972, Teskey also collected larvae from the edge of a slow-moving stream in Virginia (Pe- chuman 1973). The flight period for Chrysops celatus is late May to September in its northern range, and from April to late October in the south. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Peak flight time over its entire range is in June. Although it is a common species, it is not abundant enough to be considered a pest. Discussion.—Pechuman (1949) first de- scribed C. celatus as a subspecies of C. flavidus, and it was so treated until Teskey (1969) found the larvae and pupae of the two subspecies to be specifically distinct. The adult of C. celatus most closely resem- bles C. flavidus and C. reicherti, and can be separated by its greenish-grey mesoscutum and the longer, more slender antennae. The sinuous outer margin of the crossband in C. celatus is similar to that of C. reicherti, but in C. flavidus this margin is either straight or concave. The larva resembles C. flavidus (Goodwin 1976) and C. dixianus (Teskey 1969). The characters used to define C. celatus are consistent, in most cases, but there is some variation in the color pattern of the abdomen and wings, and in the color of the scutellum. The abdominal pattern on ter- gites 2-3 may vary from a black inverted ““V’’ on the integument to lighter markings of dark hairs only. The apical spot of the wing may be extended and fade out along the hind margin of the wing, approaching a pattern much like that of C. reicherti. Spec- imens with this wing pattern tend to be more robust and have a more darkly infus- cated wing. The scutellum, which is usually dark basally and reddish apically, is either wholly dark or reddish. Chrysops dixianus Pechuman (Pigss.7/,9116;/25) Chrysops dixianus Pechuman 1974: 185-— 187. Holotype female: Wedge Plantation, McClellanville, South Carolina (CUIC); Goodwin 1976: 345-347, description of larva and pupa. Diagnosis.—Yellow and brown species, length 6—8.5 mm. Antenna not swollen, fla- gellum % longer than scape and pedicel combined. Frons width at base subequal to width at vertex. Hind coxa and femur dark VOLUME 101, NUMBER 1 185 tumidicornis _ ) » YY 7 . Figs. 22-30. Female wing patterns. 22, Chrysops atlanticus. 23, C. brunneus. 24, C. celatus. 25, C. dixianus. 26, C. flavidus. 27, C. pudicus. 28, C. reicherti. 29, C. sandyi. 30, C. tumidicornis. 186 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON brown. Wing picture with 5th posterior cell hyaline, infuscated area bordering vein Cu,. Female.—Scape and pedicel not notice- ably enlarged, yellow with black setae, bas- al flagellomere yellow basally, remainder brown, apical flagellomeres dark brown to black, flagellum approximately % longer than length of scape and pedicel combined, antennal ratio 13:9:32. Frontoclypeus glossy yellow, eye margins bearing yellow tomentum that extends medianly beneath tentorial pits to frontoclypeal suture. Max- illary palp light brown with sparse black hair basally, more dense apically. Frons grayish-yellow pollinose with scattered yel- low hairs, width at base subequal to width at vertex, height 1.15 times width at base. Frontal callus oval, slightly inflated, light brown, width 1.4 times height. Vertex yel- low gray, glossy surrounding each ocellus, ocelli yellow. Mesoscutum yellow-gray to- mentose in ground color with 3 dark brown longitudinal stripes, median one flanked with lighter shade of brown, the two sub- lateral stripes merge with median one near the scutellum, yellow hair scattered over entire mesoscutum. Scutellum dark brown basally, light brown apically. Pleuron yel- low-gray tomentose. Fore coxa and femur yellow, fore tibia with basal % yellow, api- cal % and fore tarsus dark brown, bearing black hair. Mid coxa dark brown, femur, tibia and metatarsus yellow, remainder of tarsus dark brown. Hind coxa and femur dark brown, tibia light brown basally, dark- ening to apex, metatarsus light brown, re- maining portion of tarsus dark brown, tibia and tarsus bearing stiff black hair. Wing pattern distinct, apical spot broad, filling half of 2nd submarginal cell, hyaline tri- angle not quite reaching vein R,,,;, outer margin of crossband straight or convex, 5th posterior cell hyaline, vein Cu, infuscated along border, cells br and bm ¥% and Y; in- fuscated basally respectively. Halter with stalk brown, knob dark brown. Abdominal pattern distinct, tergite 1 yellow, light brown under scutellum, tergite 2 with an- terior half entirely yellow, posterior half yellow with 2 median oblique dark brown dashes making a flattened inverted *“‘V”’ not reaching posterior margin of tergite, poste- rior margin with median yellow triangle and 2 submedian brown spots, tergite 3 with wide dark brown band that narrows medianly, not reaching anterior or posterior margins, posterior margin yellow, tergites 4—5 with anterior % dark brown, remaining 3 yellow, tergites 6—7 dark brown. Sternites 1—4 yellow, 4th sternite with dark brown median spot, 5th sternite brown with yellow hind margin, remaining sternites dark brown. Male.—Not previously described. Length 7.4 mm. Scape and pedicel not no- ticeably enlarged, yellow with black hair, basal flagellomere yellow at base, remain- der brown, apical flagellomeres black, an- tennal ratio 15:12:34. Maxillary palp yel- low with long yellow hair and a few black hairs, length twice width. Mesoscutum yel- low-gray tomentose, above wing bases yel- low tomentose, with 3 dark brown longi- tudinal stripes that merge at the base near the scutellum. Scutellum dark brown at base, reddish brown apically. Fore coxa and femur yellow, tibia with basal % yellow, apical 4% and tarsus dark brown, bearing black hair. Mid coxa dark brown, femur, tibia and metatarsus yellow, remainder of tarsus dark brown. Hind coxa and femur dark brown, tibia light brown basally, dark- ening to apex, metatarsus light brown, re- maining portion of tarsus dark brown, tibia and tarsus bearing stiff black hair. Wing pattern with cells br and bm ¥% and % in- fuscated basally, hyaline triangle reaching vein R,,;, infuscated area bordering Cu, vein wider than in female. First abdominal tergite yellow, with light brown shading un- der scutellum, 2nd tergite with anterior % yellow, posterior ¥% with 2 median oblique dark brown dashes not attaining posterior margin of tergite, posterior margin with vague, median yellow triangle, tergite 3 with wide dark brown median band not reaching anterior or posterior margins, bor- dered laterally by black setae, hind margin VOLUME 101, NUMBER 1 yellow, tergites 4—5 with anterior *% dark brown, remaining % yellow, tergites 6—7 dark brown. Sternites 1-3 yellow, the 3rd sternite with a dark brown median spot, sternites 5—6 brown with yellow hind mar- gin, remaining sternite dark brown. Based on 1 specimen from Alachua Co. Florida, collected by G. B. Fairchild, V-24/25-1975, (FSCA). Material examined.—468 @, 1 6, ex- amined from the following museums: ANINEL, “CNCI, CUCGE, CUIC, FSCA, INHS, TAMU, UGCA, UNHC, USNM. Distribution.— Virginia to Florida, west to Louisiana and Arkansas. I have exam- ined specimens from: AL, AR, FL, GA, LA, MS, NC, and SC. L. L. Pechuman has locality data from VA. Biology.—Little is known about the bi- ology of Chrysops dixianus. The larva was taken from the edge of a freshwater lake in mud and decomposing leaves (Goodwin 1976). Pechuman (1974) stated that C. dix- ianus could be a common pest, as suggested by 124 specimens collected on July 6, 1971, in Berkeley County, South Carolina, by D. C. Sheppard. Discussion.—This species was once con- fused with C. pudicus but differs in having a longer, thinner flagellum, a brown meso- scutum, and the anterior portion of tergite 2 without markings. The adult of C. dixi- anus appears most similar to C. reicherti. The 2nd abdominal tergite of both has markings only on the posterior 4%, but C. dixianus is smaller, has a defined apical spot, and the Sth posterior cell is hyaline. There is little variation in this species. The mesoscutum, normally with a narrow median brown stripe, may have the stripe as wide as the two sublaterals. The scutel- lum may be all dark, and the abdominal markings on tergites 2—3 can vary from me- dian dashes to a more extensive pattern reaching the lateral margins. The hind fe- mur can be up to % light brown. Chrysops flavidus Wiedemann (Eigs 38;, 17/2426) Chrysops flavidus Wiedemann 1821: 55. Holotype female: Savannah, Georgia 187 (MZLU); Krober 1926: 291—292, rede- scription; Teskey 1969: 39, description of larva and pupa. Chrysops pallida Macquart 1838: 162 (1838: 166). Holotype female: locality unknown (MNHN). Synonomized by Philip 19653325. Chrysops pallidus Bellardi 1859: 73. Ho- lotype female: Mexico (MRSN). Preoc- cupied by Macquart 1838. Chrysops guiterasi Brunetti 1923: 401. Syntypes female and male: Manzanillo, Cuba, 2 (BMNH), ¢ (ZMHB). Synon- omized by Bequaert, 1940: 279. Diagnosis.—Light brown, length 8-10 mm. Scape and pedicel slightly swollen. Mesoscutum yellow with brown stripes. Hind margin of crossband straight or con- cave, cells br and bm ¥% and ¥% infuscated basally. Female.—Scape slightly enlarged, scape and pedicel light brown, black setose, basal flagellomere light brown, apical flagello- meres black, equal to or longer than basal flagellomere, antennal ratio 15:10:30. Fron- toclypeus glossy yellow, eye margins bear- ing yellow tomentum that extends medianly beneath tentorial pits to frontoclypeal su- tures. Maxillary palp and labrum yellow. Frons predominantly yellow tomentose, slightly convergent above, height 1.28 times width at base. Frontal callus oblong, bulbous, light brown, width 1.83 times height. Vertex yellow tomentose, except glossy around each ocellus, ocelli black. Mesoscutum yellow tomentose, yellow pi- lose, bearing three subshiny brown longi- tudinal stripes, median one narrower than the sublaterals. Scutellum yellow basally, orange-yellow apically. Pleuron yellow to- mentose. Legs predominantly yellow, mid and hind coxa brown, hind tibia with scat- tered black hair apically, fore, mid, and hind tarsi with black hair. Wing pattern with clearly defined hyaline areas, apical spot entering posterior half of 2nd submarginal cell, apex of hyaline triangle not quite reaching vein R,,;, outer margin of cross- 188 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON band straight or slightly concave, 5th pos- terior cell infuscated basally, hyaline api- cally, cells br and bm % and % infuscated basally. Halter with brown knob and yellow stalk. Abdominal tergites golden brown, tergites 2—5 with median yellow pollinose triangles, each triangle outlined submedi- anly with indistinct brown spots, first tergite with two submedian brown spots, hind mar- gin of tergites 3—6 narrowly yellow tomen- tose, tergites bearing mixed black and yel- low hairs. Sternites golden yellow tomen- tose, sternites 2—6 bearing a narrow yellow posterior border, sternites 4—5 with median indistinct brown spot. Male.—Similar to female except for the usual sexual differences and the following characters. Length 6.5—7.2 mm. Scape and pedicel with longer, finer, black hair. Wing faintly tinted below the apical spot and along the hind margin, basal % of cell br infuscated. Abdomen without distinct me- dian triangles. Material examined.—1,245 2, 75 ¢ ex- amined from the following museums: AMINH BMNH, CASC,..CNCI,..CUCC, CUIC, FMNH,.:FSCA,INHS, .OSUC, TAMU, UGCA, UNHC, USNM. Distribution.—Missouri to Massachusetts south to Texas, Florida, Mexico, the Baha- mas, Belize, and Cuba. I have examined specimens from AL, AR, DE, FL, GA, IL, IN, KS, LA, MA, MD, NC, NJ, NY, OK, SC, TN, TX, VA, Belize, the Bahamas, and Cuba. L. L. Pechuman recorded specimens from CT, IA, KY, MO, MS, OH, PA, RI, WV, and Mexico. Biology.—The larva described by Teskey (1969) was taken from sand on the edge of a pond. Tidwell (1973) collected most of the Louisiana specimens from the banks of ponds and waterways in bottomland hard- wood forests associated with the Mississip- pi flood plain. Jones and Bradley (1923) reared a larva found at the bottom of a brook. Jones and Anthony (1964) collected larvae from the margins of brackish water and from highly alkaline soil and reared 38 specimens whose pupal period averaged 8 days. Adults in northern regions, are active from June to October and most abundant in July. In the south they fly as early as March and continue until late September. In some areas of Florida there are two distinct pop- ulation peaks of Chrysops flavidus, one oc- curring in April and May, the other in Au- gust and September (Jones and Anthony 1964). This species was described as pestiferous to humans (Hine 1906, Jones and Anthony 1964), generally attacking the head and neck, and considered one of the worst “stock pests”’ in the genus Chrysops (Hine 1906). Discussion.—Chrysops flavidus is most commonly confused with C. atlanticus, C. reicherti and C. celatus, all considered at one time to be either subspecies or varieties of C. flavidus. It is also similar to C. sandyi, described below. Chrysops flavidus differs from C. atlanticus in having thinner anten- nae, a yellow mesoscutum, and a wing pat- tern with well defined hyaline areas. Chrys- ops flavidus differs from C. celatus in its shorter, stouter antennae, yellow mesoscu- tum, straight or slightly concave outer mar- gin of the crossband, and its less distinct abdominal pattern. It differs from C. reich- erti in having the outer margin of the cross- band straight or slightly concave, yellow hind femur, and an inverted *“‘V”’ pattern that reaches the anterior half of tergite 2. Chrysops flavidus differs from C. sandyi in its larger size, yellow mesoscutum with dark brown stripes, and its more divergent frons with the width at the vertex greater than the width of the basal callus. Variation in C. flavidus is found in the tinting of the wing and in the abdominal color pattern. Approximately 10% of the specimens examined had tinting of the wing below the apical spot, approaching the smoky wing pattern of C. atlanticus. How- ever, C. flavidus has a consistently straight or slightly concave outer margin of the crossband and yellow mesoscutum. Speci- VOLUME 101, NUMBER 1 mens with the apical portion of the wing tinted have been collected from the coastal areas of South Carolina, Georgia, and Flor- ida. A collection of 27 specimens from Monroe County, Florida, exhibit a tinted wing, have a stouter scape, a grayer meso- scutum, and a reddish brown abdomen. Other than these differences, these speci- mens do not vary in size and other char- acters. The female syntype of Chrysops guiter- asi, described from Cuba and considered a synonym of C. flavidus, was examined. I have no doubt it is conspecific with C. flav- idus, although the abdomen is reddish- brown. Chrysops pudicus Osten Sacken (Figs. 9; 185.27) Chrysops pudicus Osten Sacken 1875: 381— 382. Lectotype female: Beverley, Mas- sachusetts, (MCZC). Lectotype exam- ined, missing head; Goodwin 1976: 350— 351, description of pupa. Diagnosis.—Brown and black, length 6.5-8.2 mm. Frontal callus reddish-brown with black upper margin. Mesoscutum steel gray with black stripes. Hyaline triangle reaches vein R,,;, apical spot narrowly en- ters 2nd submarginal cell. Cells br and bm Y% and , infuscated basally. Female.—Scape and pedicel slightly swollen, ground color shiny brown, bearing stiff black hairs, basal flagellomere light brown gradually darkening at apex, apical flagellomeres black, flagellum subequal to length of scape and pedicel combined, an- tennal ratio 13:9:23. Frontoclypeus glossy reddish brown, eye margins bearing light brown tomentum that extends medianly be- neath tentorial pits to frontoclypeal suture. Maxillary palp yellow. Frons light brown tomentose, yellow pilose, slightly conver- gent above, length 1.3 times width at base. Frontal callus oblong, bulbous, reddish brown with black upper margin, margin concolorous with smooth integument sur- rounding ocelli. Vertex dark brown, glossy 189 around each ocellus, ocelli black. Scutellum black. Fore coxa light brown, trochanter slightly darker, femur light brown, apical 4 darker brown, tibia with black hair apically, tarsi black, mid coxa dark brown, femur light brown with scattered stiff yellow hair, tibia light brown with short black setae api- cally, tarsus light brown basally, dark brown apically, covered with black setae, hind femur dark brown with yellow hair, tibia light brown with yellow hair basally, remainder with yellow and black hair, tar- sus light brown basally, black apically. Wing pattern with clearly defined hyaline areas, apical spot narrow, entering extreme apical portion of 2nd submarginal cell, hy- aline triangle reaches vein R,,,, outer mar- gin of crossband straight or slightly con- cave from vein R,,, to vein M,, cell br and bm % and Y, infuscated basally. Abdomen with first 2 segments yellow, Ist tergite with large black spot under scutellum not reaching posterior margin, extending later- ally to approximately % width of tergite, 2nd tergite with black integumental invert- ed “‘V”’ not reaching anterior or posterior margin, black hair scattered over black in- tegumental pattern, orange median triangle under black inverted ““V” pattern, tergites 3—4 black anteriorly with median orange triangle posteriorly, black pattern not reach- ing lateral margin of the tergite, tergites 5— 6 black anteriorly, yellow posteriorly. Ab- domen yellow ventrally, sternites 3—4 with median black spot, spot on sternite 3 small- er than on sternite 4, remaining sternites black. Male.—Similar to female except for the usual sexual characteristics and the follow- ing characters. Scape and pedicel covered with long, fine, black hair. Mesoscutum al- most entirely black, the 3 stripes indistinct, with scattered yellow hair. Apical spot slightly more extensive. Cell br %4 infuscat- ed basally. Material examined.—91 @, 1 3d exam- ined from the following collections: CUIC, FSGA, UNHE: Distribution.—Southern Illinois to Mas- 190 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sachusetts, south to eastern Texas and Flor- ida with some disjunct populations in north- ern Indiana, Michigan, New York, southern Ontario, and Nova Scotia. I have examined specimens from AL, CT, FL, GA, LA, LA, NC, NH, NY, RI, SC, and Nova Scotia. L. L. Pechuman has records from NJ, DE, IN, IL, MD, MI, MS, OK, TN, TX, and WI. Biology.—The larva of this species, al- though not formally described, has report- edly been taken from wet soil along a road- side ditch (Jones and Anthony 1964, Pe- chuman 1973). The pupa was described by Goodwin (1976) from Baldwin County, Al- abama, but the habitat is unknown. Jones (1953) indicated that in parts of Florida there are two generations of C. pudicus an- nually, one that emerges in May—June and another in August-September. Large num- bers can occur in wooded areas of Florida during April and May (Jones and Anthony 1964). Discussion.—Pechuman et al. (1983) treated C. pudicus as a member of the Chrysops callidus group, which includes C. dimmocki Hine, a species that can resemble C. pudicus. Members of the C. callidus group all have a black frontal callus, in- verted ““V”’ pattern on 2nd abdominal ter- gite that reaches the anterior margin and, in some species, a hyaline triangle that ex- tends beyond vein R,,;. Chrysops pudicus differs from members of the C. callidus group in having an abdominal pattern on tergite 2 not reaching the anterior margin, a hyaline triangle not crossing vein R,, and a yellow frontal callus with dark top margin. The frontal callus is rarely entirely dark (Brennan 1935, Teskey 1990), and the hyaline triangle rarely reaches beyond vein R,,3;. Therefore, I have chosen to place C. pudicus in the Chrysops flavidus group. Chrysops pudicus most closely resembles C. tumidicornis, described below, in having similar size, dark coloration, and Ist ab- dominal tergite with a dark spot under the scutellum. They differ in that C. pudicus has a narrower scape and pedicel, a less ex- tensive, well defined apical spot, and a hy- aline triangle that reaches R,,. Variation within C. pudicus occurs in the extent of the hyaline triangle (Pechuman 1973) and in the color of the frontal callus. Of the 91 females examined, 10 specimens from Florida had a hyaline triangle that did not reach vein R,,,;. Although variation in color of the frontal callus is mentioned in various publications (Brennan 1935, Teskey 1990), all the specimens I examined had a yellow frontal callus with a dark upper mar- gin. Chrysops reicherti Fairchild (Figs. 10, 19; 28) Chrysops reicherti Fairchild 1937: 60-61. Holotype female: Monticello, Jefferson County, Florida (MCZC). Holotype ex- amined. Goodwin 1972: 105-107, de- scription of larva and pupa. Chrysops flavida reicherti: Philip, 1947: 273, treated as a subspecies. Pechuman 1957: 30-31, description of male; Philip 1965: 325, treated as a variety of C. flav- idus. Diagnosis.—Yellow and brown, length 8-9.5 mm. Antenna not swollen. Meso- scutum yellow in ground color with 3 brown stripes. Tergites 1—2 light yellow, darker pattern on posterior half of tergite 2. Cells br and bm are % and % infuscated basally. Female.—Scape and pedicel not swollen, light brown with black setae, flagellum black, antennal ratio 15:10:35. Frontocly- peus glossy yellow, eye margins bearing grayish yellow tomentum extending medi- anly beneath tentorial pits to frontoclypeal suture. Maxillary palp brown. Frons yellow tomentose, slightly convergent above, height 1.3 times width at base. Frontal cal- lus oblong, bulbous, light brown, width 1.33 times height. Vertex lhght brown around each ocellus, ocelli yellow. Meso- scutum yellow tomentose with three sub- shiny brown longitudinal stripes, median one narrower than sublaterals, flanked with VOLUME 101, NUMBER 1 lighter shade of brown. Scutellum yellow brown. Pleuron yellow tomentose. Legs predominantly light brown, fore and mid tarsi brown, mid and hind tibiae with dark hair apically, hind femur reddish brown, apical tarsomeres brown to black. Wing with extensive apical spot reaching poste- rior % of 2nd submarginal cell fading into 3rd posterior cell, hyaline triangle not reaching vein R,,;, outer margin of cross- band bowed outward at center, 5th posterior cell infuscated basally and along vein mar- gins, hyaline apically, cells br and bm % and %4 infuscated basally. Halter with brown stalk and knob. First abdominal tergite light brown, second tergite with basal half yel- low, green tinted medianly, apical half with oblique brown spots reaching hind margin and enclosing pale triangle. Tergite 3 with brown median band set over median orange triangle, hind margin of tergite gray. Ter- gites 4—6 with basal half brown, apical half yellow. Sternites predominantly golden brown, 3—6 with median brown spot pro- gressively enlarged on posterior segments. Male.—Similar to female except for the usual sexual differences and the following characters: Length 6.5—7.0 mm. Scape swollen, pedicel /; longer than in female, scape and pedicel with longer, finer, black hair. Mesoscutum with long yellow hair. Wing with apical spot more extensive, cells br and bm % and % infuscated basally. Material examined.—720 ¢, 20 ¢ ex- amined from the following collections: CASCS ENC: CUIC;, -FSCA, INHS, TAMU, UGCA, UNHC, USNM. Distribution.—Southern Illinois to Dela- ware, south to eastern Texas and Florida. I have examined specimens from AL, AR, RENGAHIL, KY, LA, MA; MD, MS, NC; SC, TN, TX, and VA. L. L. Pechuman has records from IN and MO. Biology.—The larvae of Chrysops reich- erti have been collected in very wet mud and organic debris from small ponds and lakes (Goodwin 1972, Tidwell 1973). Adults have been collected between April and September in the southern states. The 19] one Massachusetts specimen was collected on May 8. Discussion.—Chrysops reicherti was originally described as a species by Fair- child (1937). Philip (1947) listed it as a sub- species of C. flavida. Goodwin (1972), after describing the larva and pupa of C. reich- erti and comparing them to those of C. flav- idus, considered them separate species and elevated C. reicherti to a full species. Chrysops reicherti is most commonly confused with C. flavidus, but is separated by its pale yellow first two abdominal seg- ments, sinuous outer margin of the cross- band, and the slightly darker hind femora. In the key above, C. reicherti shares with C. dixianus the color pattern on the poste- rior portion of the second tergite. They dif- fer in that C. dixianus has a straighter outer margin of the crossband, an apical spot that reaches slightly beyond the apex of the wing, and Sth posterior cell extensively hy- aline. Variation in C. reicherti includes tinting in the usual hyaline areas of the wing, 2nd abdominal tergite without the greenish cast described by Fairchild and, hind femur light or dark brown. Chrysops sandyi Baier, new species (Rigs. 12, 205.29) Diagnosis.—Yellow and brown, length 6-8 mm. Antenna slightly enlarged. Me- soscutum with brown iridescent stripes. Wing slightly tinted in usual hyaline areas. Cells br and bm are % and % infuscated basally. Holotype female.—Length 6.2 mm. Scape slightly enlarged, antenna light brown except apical 3/4 of Ist flagellomere slightly darker, apical 4 flagellomeres black, antennal ratio 13:9:24, length of flagellum subequal to combined length of scape and pedicel. Frontoclypeus glossy yellow, eye margins bearing yellow tomentum that ex- tends medianly beneath tentorial pits to frontoclypeal suture. Maxillary palp light brown with sparse black hairs, length 4 times width. Frons yellow tomentose with 192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON scattered yellow hairs, convergent above, height slightly more than width at base, width at vertex less than width of frontal callus. Frontal callus oval, yellow brown, width 1.7 times height. Vertex yellow to- mentose, glossy around each ocellus, ocelli light brown. Mesoscutum yellow-green iri- descent tomentose, except lateral margins iridescent yellow gray, bearing 3 brown to dark brown stripes, the median one darker and narrower than the two sublaterals. Scu- tellum dark brown basally, brown apically. Pleuron yellow gray. Legs mostly light brown, fore tarsus black, apical portion of tibia dark brown with black hair, mid coxa dark brown, tarsus with black hair, hind coxa dark brown, femur and basal tibia red- dish brown, apical tibia brown, apical tibia and tarsus with black hair. Wing with broad apical spot reaching posterior half of 2nd submarginal cell, apex of hyaline triangle including posterior half of Ist submarginal cell, not reaching vein R,,;, Outer margin of crossband straight, 5th posterior cell slightly tinted, infuscated along vein Cu,, cells br and bm % and % infuscated basally. Halter with light brown stalk and brown knob. Tergites 1—2 yellow brown, tergite | with a few dark hairs under scutellum, ter- gite 2 with a median brown integumental marking appearing as a flattened inverted “V"’, not reaching anterior margin, fading into pale posterior margin, median orange- brown triangle behind inverted ‘“*V’’ pat- tern, tergite 3 mottled dark brown anteri- orly, orange brown posteriorly, extending medianly to form orange triangle, tergites 4—6 dark brown anteriorly, yellow orange posteriorly, tergites 3-6 with gray hind margin. Sternites 1—3 yellow brown, ster- nite 3 bearing a median brown spot, ster- nites 4—7 dark brown, posterior margin pale yellow tomentose. Male.—Unknown Material examined.—Holotype 2, AL- ABAMA: Mobile County, VH-1952, T. R. Adkins (CUIC). Paratypes: 105 2. ALA- BAMA: Mobile County, VII-1952 (62 @), VII-24-1952 (4 2), T. R. Adkins (CUIC); 1 2 Corden, VII-8-1962, M. Tidwell (FSCA). LOUISIANA: St. Tammany Parish, 4 @ Slidell, 7 mi SE, VI-23-1980, E. A. Lisow- ski (CUIC). MISSISSIPPI: Hancock Coun- ty, Ansley, VI-10-1966 (5 @), VI-20-1966 (6 2), Diamond & Bradford (FSCA); 2 @ V-5-1976, M. Tidwell (FSCA); 1 2 Pear- lington, VI-9-1966, ? collector (FSCA); 1 2 MTO fee area, IV-27-1965, R. Hepburn (FSCA). Jackson County, Fountainbleau, VI-26-1966 (2 2), VII-27-1968 (1 ¢), B. Byrd (FSCA); 4 2 Gautler, 4.5 mi WSW, VI-24-1980, E. A. Lisowski (CUIC); 12 2 Ocean Springs, V-10-1970, G. Ross (FSCA). Etymology.—Named in honor of G. B. **“Sandy”’ Fairchild for his extensive contri- bution to the organization of Chrysops taxa and his desire to see the southeastern spe- cies clarified. His examination of specimens and critical comments on this work has been greatly appreciated. Distribution.—Gulf coast of Alabama, Mississippi, and Louisiana. Biology.—Chrysops sandyi has been col- lected between April 27 and August 8. Based on its distribution along the Gulf coast, it is probably limited to a litoral en- vironment. Discussion.—This species has been con- fused with C. atlanticus because of the tint- ed wing and greenish iridescent color of the mesoscutum. Chrysops sandyi has a nar- rower antennae, the pedicel %4 the length of the scape, longer palpi, length 4 times width of apical palpomere, mesoscutum with brown longitudinal stripes, and lightly tint- ed wing pattern. Chrysops atlanticus inhab- its the Atlantic coast from Maine to Florida while C. sandyi is found along the Gulf coast of Alabama, Mississippi, and Louisi- ana. Variation in C. sandyi occurs in the ground color of the mesoscutum, and in the abdominal pattern. Ground color of the me- soscutum, although usually greenish irides- cent, may have yellow tomentum. Abdom- inal pattern may be indistinct, with a pattern of black hair and no integumental markings. VOLUME 101, NUMBER 1 Specimens with these features appear sim- ilar to C. flavidus, but C. sandyi is smaller, wing tinted in the usual hyaline areas, and flagellum subequal to the combined length of the scape and pedicel. Chrysops tumidicornis Baier, new species (Figs. dil; 2 129) Diagnosis.—Black and brown, length 5.8—8 mm. Antenna swollen, pedicel barrel shaped. Frontal callus width approximately 2 times height. Mesoscutum grey green with black stripes. Wing with distinct hya- line triangle, cells br and bm % and % in- fuscated basally. Holotype female.—Length 7.4 mm. Scape and pedicel swollen, pedicel barrel shaped, brown, covered with black setae, basal flagellomere brown, apical 4 flagel- lomeres black, flagellum subequal to length of scape and pedicel combined, basal fla- gellomere equal to combined length of the apical 4 flagellomeres, antennal ratio 14:11: 26. Frontoclypeus glossy yellow, eye mar- gins bearing yellow tomentum that extends medianly beneath tentorial pits to frontocly- peal suture. Maxillary palp light brown with scattered stiff black hairs, length 4 times width of apical palpomere. Frons yel- low tomentose, slightly convergent above, height 1.2 times width at base. Frontal cal- lus oval, slightly bulbous, yellow, width 1.9 times height. Vertex yellow brown except glossy black around each ocellus, ocelli black. Mesoscutum grayish-green ground color with 3 longitudinal black stripes not merging basally, median one narrower than two sublaterals, yellow tomentose above wing bases. Scutellum black basally, red- dish brown apically. Fore coxa, femur and basal % of tibia yellow, remaining tibia brown, black setose, tarsus entirely dark brown, black setae, mid coxa brown, femur, tibia and basal tarsomeres yellow, tarsi and apical % of tibia with black setae, hind coxa, femur and tibia brown, scattered yel- low hair and black setae, black setae be- coming more dense toward apical portion 193 of tibia and tarsus. Wing pattern distinct, apical spot includes apical %4 of vein R,, reaching posterior half of 2nd submarginal cell, and fading along hind margin of wing to crossband, hyaline triangle slightly tint- ed, not reaching vein R,,,, outer margin of crossband straight, 5th posterior cell infus- cated along margins, center hyaline, cells br and bm ¥% and % infuscated basally respec- tively. Abdominal tergites orange yellow, Ist tergite dark brown under scutellum, ter- gite 2 with median black inverted ‘‘V”’ pat- tern on integument not reaching anterior, posterior or lateral margins, scattered black hair over dark pattern, median orange tri- angle behind inverted ““V”’ pattern, hind margin gray, tergite 3 with gray posterior border widened medianly to form an indis- tinct triangle, anterior half with black pat- tern of 2 half circles that merge above tri- angle and occupy % width of tergite, ter- gites 4—6 black anteriorly, posterior border gray. Abdomen orange yellow ventrally, sternites 3—5 with median dark spot becom- ing wider on each successive sternite, ster- nites 6—7 dark brown. Male.—Unknown. Material examined.—Holotype 92, FLORIDA: Levy County, Cedar Key Shell Mound, VII-7-1976, R. H. Roberts (FSCA). Paratypes: 206 2. FLORIDA: Bay County, 2 2 V-14-1957, E W. Mead (CASC); 1 V-25-1982, J. Hogsette (FSCA); Saint An- drews St. Park, 1 2 V-14-1957, E W. Mead (CASC). Citrus County, 3 2 Ozello, VII- 29-1959, H. V. Weems (CASC). Collier County, | 2 Collier-Seminole State Park, XI-26-1976, John Edward Rawlins (CUIC). Dade County, | 2 Everglades National Park, I-29-1959, H. A. Denmark (CASC); 1 2 Royal Palm Hammock, VI-22-1951, Price, Beamer & Wood (CASC); 1 2 West Lake, XII-4-1970, P H. & M. Arnaud (CASC). Dixie County, 2 2 Jena, VII-27- 1991, Jena, 10 mi S on RT 361, VII-22- 1985 (2 2), VII-13-1991 (8 2), VII-21- 1991 (5 ¢), VII-27-1991 (4 2), VII-31- 1991 (13 @), VII-11-1992 (1 2) L. R. Da- vis, Jr. (FSCA); 3 2 Suwannee, V-19-1964, 194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON C. E Zeiger (CASC); 2 2 Suwannee, V-19- 1964 (FSCA). Franklin County, Carrabelle, IV-25-1980 (2 2), IV-27-1980 (3 @), IV- 28-1980 (1 2), IV-29-1980 (4 2), V-4-1980 (5 ¢), IV-14-1977 (1 2), L. L. Pechuman (CUIC); IV-28-1980 (1 2), V-4-1980 (1 2), L. L. Pechuman (FSCA); Eastpoint, 1 2 V- 19-1935 (FSCA); Lake Morality, IV-27- 1980 (1 ¢), V-10-1980 (1 2), 1 2 Timber Island, V-4-1980, L. L. Pechuman (CUIC); 1 2 Timber Island, V-4-1980, L. L. Pechu- man (CASC); 2 ¢ St. George Island, IV-5- 1976, L. L. Pechuman (FSCA). Gilchrist County, 1 @ VI-2—-1950, (CASC). Gulf County, 4 2 V-11-1973, J. T. Goodwin (FSCA), V-4-1973 (1 @), VII-14-1971 (1 2), H. V. Weems, Jr (FSCA). 2 2 St. Jo- seph State Park, V-1/3-1970, W. W. Wirth (USNM); 1 2 V-5-1987, L. Strange and J. Wiley (FSCA). Hernando County, 1 2? Bay- port, [V-22-1978, L. A. Strange (FSCA). Hillsborough County, 1 @ Tampa, IV-29- 1950, G. B. Worth (CASC). Jefferson County, 1 2 Monticello, IX-18-1935, G. B. Fairchild (FSCA). Lee County, 2 2? Sanibel Island, IV-8-1933, W. J. Clench (USNM); 2 2 Sanibel Island IV-8-1933, W. J. Clench (FSCA). Levy County, 2 2 VII-8-1980, E. Davis (FSCA); 1 2 V-6-1955, H. V. Weems (FSCA); 3 2 Cedar Key VI-20-1974, (FSCA); 5 2 Cedar Key area, VI-4-1991, J. Huether (CUIC); 3 2 Cedar Key, VII-12- 1939; R.-H; Beamer (CASC): 3 2 VI-12- 1939, Oman (USNM); 1 2 VII-29-1977, L. R. Davis, Jr. (FSCA); 1 2 VI-28-1973, Carl Shleck, (FS@A);\ 1.9) VI-15-1979; 14) As Wood (FSCA); Cedar Key, Shell Mound, VI-9-1971 (5 ¢), VI-16-1973 (1 2), VI-29- 1976 (6 2), VII-3-1976 (7 2), G. B. Fair- child (FSCA); 1 @ 3 mile E. Shell Mound, 2 @ VI-9-1971, (FSCA); 8 2 VII-7-1976, R. Roberts (FSCA); 3 2 VI-21-1973, R. Wilkerson (FSCA); Yankeetown, V-8/13- 1980 (1 2), VII-8-1980 (1 2), V-16-1980 (3 2), VI-24-1980 (5 ¢), E. Davis (FSCA); V-21/23-1979 (1 2), V-14/16-1979 (2 2), V-11/14-1979 (1 2), V-23/24-1979 (3 Q), V-25/30-1979 (2 ¢), V-30/31-1979 (1 @), Roberts (FSCA); 3 2 V-30/31-1985, D. L. Kline (FSCA); VIII-10-1966 (1 @), VII-28- 1965 (4 2), C. E Zeiger (FSCA). Manatee County, 1 @ VI-12-1925, T. H. Hubbell (CASC). Monroe County, 1 2 Everglades National Park, IV-8-1970, W. W. Wirth (USNM). Okaloosa County, 4 @ V-12- 1972, J. T. Goodwin (FSCA). Pasco Coun- ty, 5 2 Hudson, VH-13-1939, R. H. Beamer (CASE);. 11 2 -VITI-13-1939;' Oman (USNM); 1 2 VII-13-1939, Oman (FSCA); 2 2 VII-13-1939, D. E. Hardy (CASC): Pi- nellas County, 1 2 IV-19-1930, B. P. Moora (USNM); 1 @ V-2-1955, R. P. Essar (FSCA); 1 2 V-10-1955, E. W. Holder, Jr. (FSCA); 1 2 Dunedin, V-14-1959, O. L. Cartwright (USNM); 2 2 Pass-a-grille, IV- 7-1930, W. G. Fargo (CASC). Santa Rosa County, | 2 Santa Rosa Island, V-24-1971, H. V. Weems Jr. (FSCA); Taylor County, 1 2 Steinhatchee, VII-12-1966, R. P. Esser (CASC); 1 2 Steinhatchee, VII-20-1991, L. R. Davis, Jr. (FSCA); 1 @ 12 mi NW of Steinhatchee, V-24-1983, R. M. Reeves (UNHC); 12 2 Cedar Island, 16 miles NW of Steinhatchee, V-16-1969, H. V. Weems, Jr. (FSCA); 3 2 Cedar Island, 16 miles NW of Steinhatchee, V-16-1969, H. V. Weems, Jr. (CASC). Wakulla County, 1 @ V-2- 1980, L. L. Pechuman (CUIC); 3 2 Och- lockonee River State Park, V-20-1968, H. V. Weems, Jr. (FSCA); 2 2 Ochlockonee River State Park, V-20-1968, H. V. Weems, Jr. (CASC); 1 Ochlockonee River State Park, VI-22-1973, Fairchild & Wilkerson (FSCA). Etymology.—This species is named for its characteristic “‘robust’’ pedicel. Distribution.—West coast of Florida. Biology.—Based of the collection data, most adults have been taken between April—August with a few collected in Sep- tember, and November—January. All speci- mens examined were taken from coastal counties near brackish habitats. Although the larva and pupa are unknown, I suspect this species is limited to salt marshes. Discussion.—This species has been con- fused with C. atlanticus because they both have swollen antennae. Also, I believe that VOLUME 101, NUMBER 1 Pechuman (1949), in his description of C. atlanticus, had before him specimens of this species. He wrote that many specimens ex- amined showed considerable melanism, varying from slight enlargement of abdom- inal spots to extreme enlargement where the tergites were mostly dark with a narrow yellow-brown margin, a pattern I find to be diagnostic C. tumidicornis. In the commonly used keys to species of North American Chrysops (Jones and An- thony 1964, Teskey 1969, Tidwell 1973, Pechuman 1973, 1981) C. tumidicornis will key to C. atlanticus. However, C. tumidi- cornis differs by having a stouter pedicel, frontal callus approximately twice as wide as tall, wing pattern with a distinct hyaline triangle, and an abdominal pattern brown with median orange triangles. Chrysops tumidicornis has an extensive apical spot reaching posterior half of 2nd submarginal cell, and a hyaline triangle that does not reach vein R,,,. Chrysops tumidicornis ap- pears to be more similar to C. pudicus in size, dark mesoscutum and abdominal pat- tern. Variation within C. tumidicornis ranges from specimes with small, slender body, and a more uniformly dark abdomen, to a larger, more robust body, with an abdomi- nal pattern more uniformly yellow-orange. Three specimens from Gulf County, Flori- da, have an entirely dark brown abdomen with a lighter median triangle on the 2nd tergite. Chrysops tumidicornis is easily rec- ognized by its bulbous, barrel shaped ped- icel. ACKNOWLEDGMENTS I acknowledge the assistance of the late G. B. “Sandy” Fairchild, Florida State Col- lection of Arthropods, for assistance with organization of data and critical comments on treatment of species, the late L. L. Pe- chuman, whose distribution records were used to create accurate distribution maps for this paper, John F Burger, University of New Hampshire, for providing critical com- ments, financial support and encouragement 195 throughout this project, Donald S. Chan- dler, University of New Hampshire, for his assistance with development of the data- base, illustrations and editorial comments, the late R. Marcel Reeves, University of New Hampshire, for editorial comments, John Weaver, University of New Hamp- shire, for assistance with illustrations, and finally the individuals responsible for lend- ing specimens held by their respective in- stitutions. This is scientific contribution number 1967 from the New Hampshire Agriculture Experiment Station. LITERATURE CITED Anderson, J. F 1971. Autogeny and mating and their relationship to biting in the saltmarsh deer fly, Chrysops atlanticus (Diptera: Tabanidae). Annals of the Entomological Society of America 64: 1421-1424. Arnett, R. H., Samuelson, G. A., and Nishda, G. M. 1993. The insect and spider collections of the world. Flora and Fauna Handbook No. 11. 306 pp. Bellardi, L. 1859. Saggio di Ditterologia Messicana. Memorie della Reale Academia delle Science di Torino 19: 246-276. Bequaert, J. 1940. The Tabanidae of the (Dipt.) Revista de Entomologia 11: 279. Brennan, J. M. 1935. The Pangoniinae of Nearctic America (Diptera: Tabanidae). The University of Kansas Science Bulletin 22(13): 1—400. Brunetti, E. 1923. Two new species of Tabanidae from Antilles Cuba. Bulletin of Entomological Research 13: 401—402. Burger, J. FE 1995. Catalog of Tabanidae (Diptera) of North America north of Mexico. Contributions to Entomology, International 1(1): 1—100. Fairchild, G. B. 1937. A preliminary list of the Taban- idae (Diptera) of Florida. Florida Entomologist 19: 58-63. Goodwin, J. T. 1972. Immature stages of some eastern nearctic Tabanidae (Diptera). I. Introduction and the genus Chrysops Meigen. Journal of the Geor- gia Entomological Society 7(2): 98-109. . 1976. Immature stages of some eastern nearc- tic Tabanidae (Diptera). VI. Additional species of Chrysops Meigen. Florida Entomologist 59(4): 343-351. Hansens, E. J. 1980. Review: Tabanidae of the east coast as an economic problem. Bulletin of the New York Entomological Society 137(4): 312- 318. Hine, J. S. 1903. Tabanidae of Ohio with a catalogue and bibliography of the species from America 196 north of Mexico. Ohio State Academy of Science, Special Papers 5: 1-63. . 1906. A preliminary report on the horseflies of Louisiana with a discussion of remedies and natural enemies. State Crop Pest Commission of Louisiana. Circular No. 6. 43 pp. International Commission on Zoological Nomencla- ture, 1993. Opinion 1711. Chrysops atlanticus Pe- chuman, 1949 (Insecta, Diptera): specific name conserved. Bulletin of Zoological Nomenclature 50: 83. Jamnback, H. and Wall, W. 1959. The common salt- marsh Tabanidae of Long Island, New York. New York State Museum and Science Service Bulletin Number 375. 1-72. Jones, C. M. 1953. Biology of Tabanidae in Florida. Journal of Economic Entomology 46(6): 1108— 1109. Jones, C. M., and Anthony, D. W. 1964. The Tabanidae (Diptera) of Florida. United States Department of Agriculture Technical Bulletin 1295, 85 pp. Jones, T. H. and Bradley W. G. 1923. Observations on Tabanidae (horseflies) in Louisianna. Journal of Economic Entomology 16: 307-312. Kréber, O. 1926. Die Chrysops Arten Nordamerikas einshl. Mexicos. Stettiner Entomologische Zei- tung 87: 1-353. Macquart, P. J. 1838. Dipteres Exotiques. Nouveaux peu connus. Part I. Tabaniens. Librairie Encyclo- pedique de Roret, Paris. Pages 90-164. Magnarelli, L. A. and Anderson, J. F 1976. Follicular development in salt marsh Tabanidae (Diptera) and incidence of nectar feeding with relation to gonotrophic activity. Annals of the Entomological Society of America 70: 529-533. Orihel, T. C. and Lourie, R. C. 1975. Loa loa: devel- opment to the infective stage in an American deer- fly, Chrysops atlanticus. American Journal of Tropical Medicine and Hygiene 24(4): 610-615. Osten Sacken, C. R. 1875. Prodrome of a monograph of the Tabanidae of the United States. Part I. The genera Pangonia, Chrysops, Silvius, Haematopo- ta, Diabasis. Memoirs of the Boston Society of Natural History 2: 381-382. Pechuman, L. L. 1949. Some notes on Tabanidae (Dip- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tera) and the description of two new Chrysops. The Canadian Entomologist 131(3): 77-84. . 1957. Descriptions of Tabanidae previously known from one sex only (Diptera). Bulletin of the Brooklyn Entomological Society 102(2): 29-— 34. . 1973. Horse flies and deer flies of Virginia (Diptera: Tabanidae). The Insects of Virginia No. 6. 92 pp. . 1974. Two new Tabanidae from southeastern United States (Diptera). Journal of the New York Entomological Society 82(3): 183-188. . 1981. The horse flies and deer flies of New York (Diptera: Tabanidae). Search Agriculture. Cornell University Agriculture Experiment Sta- tion. (Ithaca, N.Y.) 2: 1-65. Pechuman, L. L., Webb, D. W., and Teskey, H. J. 1983. The Diptera, or True Flies, of Illinois, I. Tabani- dae. Illinois Natural History Survey Bulletin 33(1): 1-120. Philip, C. B. 1947. A catalog of the blood-sucking fly family Tabanidae (horseflies and deerflies) of the Nearctic Region North of Mexico. The American Midland Naturalist 37(2): 257-324. . 1955. New North American Tabanidae. IX. Notes on and keys to the genus Chrysops Meigen. Revista Brasileira del Entomologia 3: 47-128. Philip, C. B. 1965. Family Tabanidae, pp. 319-342. 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. U.S. De- partment of Agiculture, Agricultural Research Service Handbook 276, 1696 pp. Teskey, H. J. 1969. Larvae and pupae of some eastern North American Tabanidae (Diptera). The Ento- mological Society of Canada 63: 1-147. . 1990. The horse flies and deer flies of Canada and Alaska (Diptera: Tabanidae). The Insects and Arachnids of Canada. Biosystematics Research Institute, Ottawa 16: 1-381. Tidwell, M. A. 1973. The Tabanidae (Diptera) of Lou- isiana. Tulane Studies in Zoology and Botany 18(1—2): 1-95. Walker, E 1848. List of the specimens of dipterous insects in the collection of the British museum 1: 229 pp. London. Wiedemann, C. R. W. 1821. Diptera exotica (ed. 2), 1-244, 1 fig., 2 plates. Kiliae (= Kiel). PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 197-207 TWO NEW SPECIES OF PHAENOCARPA FOERSTER (HYMENOPTERA: BRACONIDAE: ALYSIINAE) FROM SOUTH AMERICA M. TROSTLE, N. S. CARREJO, I. MERCADO, AND R. A. WHARTON (MT, IM, RAW) Department of Entomology, Texas A&M University, College Station, TX 77843, U.S.A. (e-mail: RAW: rawbaw2 @acs.tamu.edu); (NSC) Departamento de Bio- logia, Seccion Entomologia, Universidad del Valle, A.A. 25360, Cali, Colombia Abstract.—Two new species of Phaenocarpa Foerster are described from South Amer- ica: P. hyalina Trostle and P. pericarpa Wharton and Carrejo. Phaenocarpa pericarpa was reared from Anastrepha distincta Greene in pods of /nga, representing the first host record for Phaenocarpa in South America. A key separating the South American species of Phaenocarpa is included. Key Words: Papp (1969) revised the species of Phaenocarpa Foerster for the Neotropical Region, treating 10 species. Papp (1966, 1969) included Asobara Foerster as a syn- onym of Phaenocarpa in his earlier works, but Fischer (1971) subsequently removed Asobara from synonymy and it has been recognized as a separate genus ever since. Consequently, five of the 10 species treated by Papp (1969) must now be placed in Aso- bara. These are anastrephae (Muesebeck), gahani (Papp), mexicana (Ashmead), pleur- alis (Ashmead), and rubra (Papp). Their identity has been confirmed by one of us (RAW), and the placement of two of these species has been discussed previously (Wharton 1994). A sixth species, delicata Papp, has sev- eral unusual features, and its placement is uncertain (Wharton 1980, Fischer 1994). Papp (1969) divided the four remaining species into those with well-developed no- tauli and those without. Wharton (1994) de- scribed three additional species from the Neotropics. He further defined two distinct species groups to accomodate these, and discussed the differences which set these Anastrepha; parasitoid; Diptera; Neotropical two groups apart from other described spe- cies of Phaenocarpa. Two more species are described here. The first belongs to the cratomorpha Whar- ton species group as defined by Wharton (1994). The second belongs to Papp’s (1969) group with reduced notauli. The genus Phaenocarpa is fairly large, but hosts have been recorded for only about 15% of the 150 described species (Fischer 1974, 1975, 1990, 1993; Shenefelt 1974; Wharton 1984; Vet and van Alphen 1985; Tobias 1986). Holarctic species have been reared most frequently from calypterate Diptera breeding in such habitats as fungus, dung, flower heads and other seed bearing structures. A few have also been recorded from acalypterates such as drosophilids and sciomyzids (Papp 1972, Fischer 1975, Wharton 1984, Vet and van Alphen 1985, van Achterberg 1988). Hosts have not been previously recorded for any of the Neotrop- ical species of Phaenocarpa. MATERIALS AND METHODS Terminology is as in Wharton (1980, 1994) except as follows: venation conforms 198 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON to the standards recently established by Sharkey and Wharton (1997); scutellar sul- cus is used in place of prescutellar pits; the terms mesonotal disc (or simply disc) and anterior declivity are retained in their for- mer meaning, but are referred to as parts of the mesoscutum. Measurements are given to the nearest 0.05. Museum acronyms are as follows: The Natural History Museum, London (BMNH), Texas A&M University Insect Collection (TAMU). Part of the material for this study was sorted from general BMNH accessions by RAW, the remainder was reared by NSC as part of a program to de- termine natural enemies of fruit-infesting tephritid flies of the genus Anastrepha in Colombia. KEY TO NEOTROPICAL SPECIES OF PHAENOCARPA 1. Second flagellomere at least 1.4 times longer thanrhrsta(Fissw12=14) emia rene 2 — Second flagellomere at most 1.1 times longer thankstins (Bot) ieneem rican cute Gy sate cree eas 6 2. Second submarginal cell short, with 2RS near- ly equal to or slightly longer than 3RSa .... 3 — Second submarginal cell longer (Fig. 7), with 3RSa at least 1.25 times longer than 2RS ... 4 3. Notauli distinct posteriorly. Mandible with tooth | extending distinctly distad tooth 3... Ausra ceebtynk eat Phaenocarpa coxalis (Szépligeti) — Notauli absent posteriorly, not reaching midpit on mesoscutum. Mandible with tooth 3 extend- ing, distinctly distadktoothwliys hs 44 = 45. stonatine. ion aie Ores Phaenocarpa anomala Wharton 4. Mandible with broad tooth | separated from tooth 2 by a deep cleft (Fig. 15). Fore wing (RS+M)b distinctly more than half length of m-cu (Fig. 7) Phaenocarpa pericarpa Wharton and Carrejo, n. sp. — Mandibular teeth | and 2 connected by a broad, undulant flange, not separated from each other by a deep cleft (Fig. 10). Fore wing (RS+M)b short to absent, distinctly shorter than half length of m-cu 5. Petiole dark brown, narrowly elongate, about 1.5 times longer than apical width; apex about 1.6 times wider than base. Propodeal areola distinct; transverse carina of propodeum absent lateralliyeniect activi ons Phaenocarpa heynei Papp — Petiole dark yellow, broader, about 1.2 times longer than apical width, apex about twice wid- er than base. Propodeal areola indistinct; trans- verse carina complete to spiracles Duty Sette ies ASS Phaenocarpa subtilistriata Papp 6. Stigma of fore wing broad distally, abruptly narrowing basad junction with r in females (Fig. 5), with posterior margin indistinct on basal half; divided by a hyaline, desclerotized line in males, with portion of stigma posteriad hyaline region broader than portion along an- terior margin of wing (Fig. 6). Mesopleuron, metapleuron, and propodeum dark brown. Metanotum in lateral view with short but dis- tinct spine (Fig. 2) steyeanveids Newent pe Phaenocarpa hyalina Trostle, n. sp. — Stigma of fore wing narrow distally and basal- ly, only gradually narrowing basad junction with r in females, with posterior margin clearly delineated on basal half; males in which the stigma is divided by a desclerotized line have the portion that is posteriorad the desclerotized, hyaline region narrower than the portion along the anterior margin of the wing. Body color and development of metanotal projection var- iable 7. Mesopleuron, metapleuron, and propodeum yellow. Metanotum in lateral view with short, narrow spine-like projection (as in Fig. 2)... Re | te ree cory i tole Phaenocarpa sharkeyi Wharton — Mesopleuron, metapleuron, and propodeum dark brown, contrasting with light brown to yellow mesoscutum. Metanotum in lateral view with broadly triangular projection (distinctly broader than high) WAskoe: cao Phaenocarpa cratomorpha Wharton Phaenocarpa hyalina Trostle, new species (Figs. 1-6) Female.—Head: Moderately transverse in dorsal view, 1.3 wider than long. Face punctate, setiferous but polished, 1.25—1.35 higher than wide. Frontoclypeal suture broad, relatively shallow, unsculptured. Clypeus broad, weakly convex, barely pro- truding. Frons smooth, bare, nearly flat, weakly concave medially. Mandible 1.85 longer than width between tooth | and 3, surface largely smooth; tooth | broad, near- ly orthogonal, with dorsal margin concave; distinct cleft present between tooth | and 2, tooth 2 lacking additional tooth or knob; tooth 2 acutely triangular, short, 0.4—0.45 x apical width of mandible, but extending be- yond tooth | and 3; tooth 3 broadly trian- VOLUME 101, NUMBER 1 199 Figs. 1-4. Phaenocarpa hyalina. 1, Antenna showing relative lengths of basal 3 flagellomeres. 2, Mesosoma, lateral view, showing metanotal spine. 3, Mesosoma, dorsal view. 4, Metasoma, dorsal view. gular. Eye large, 4.25—5 x longer than tem- ple. Anterior tentorial pit large, extending more than 0.5 distance from lateral mar- gin of clypeus to eye. Antenna 25—27 seg- mented; second flagellomere 1.0—1.05X longer than first; first flagellomere 1.1—1.3 longer than third. Eye-antennal sulcus ab- sent. Mesosoma: 1.35—1.5X longer than high. Pronotum predominately smooth, with lon- 200 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5-8. Wings. 5, Phaenocarpa hyalina, female. 6, P. hyalina, male. 7, 8, P. pericarpa fore and hind wing. gitudinal carina separating dorsal and lat- ished; anterior declivity vertical, sparsely eral portions, and fine crenulae laterally ex- setose; notauli extending medially as cari- tending ventrally from this carina; prono- nate ridges along dorsal margin of anterior tum dorsally a narrow band with a shallow, declivity, then proceeding posteriorly as barely perceptible median pit; anterior mar- weak, parallel grooves perpendicular to the gin weakly emarginate. Mesoscutum pol- carinate ridges, notaular grooves evanescent VOLUME 101, NUMBER 1 near anterior margin of midpit; 3—4 pairs of setae extending from transverse portion of notauli to transscutal articulation; midpit well developed, extending anteriorly 0.4 length of disc from transscutal articulation. Scutellar sulcus about 2X broader than long, with medial carina; lateral margins of axilla flangelike. Metanotum with posterior margin a broad, flat plate, elevated medially as a prominent spine, median field with or without carinae radiating from spine. Pro- podeum areolate, as in other members of the cratomorpha species group. Mesopleu- ron with small patch of rugose sculpture on anterior margin medially, otherwise smooth, polished, virtually bare; carinately margined posteriorly. Sternaulus sinuate, crenulate over anterior 0.65, smooth over posterior 0.35. Metapleuron largely un- sculptured, as in P. sharkeyi. Wings: Fore wing stigma 4.5—5.0 lon- ger than broad, solid throughout (1.e., with- out hyaline streak), basal half distinctly nar- rower than distal half with posterior margin of basal half indistinct and somewhat ex- cavated, stigma abruptly widening near junction with r; r short, slightly shorter than and arising slightly distad mid-width of stigma; 3RSa 1.4—1.8 longer than 2RS; 3RSb extending to wing tip, weakly curved at apex; (RS+M)b usually absent; Icu-a of- ten very short, postfurcal by 1.25-—3.5X its length. Hind wing with 1M 2.4—3.5x lon- ger than M+CU; 3 hamuli. Metasoma: Petiole 1.6—1.7 longer than apical width, apex 1.5—1.6< wider than base, dorsal carinae on petiole more or less parallel sided, weakly converging and be- coming less prominent posteriorly; surface sculpture often granular-rugose medially; more finely rugose laterally. Ovipositor sheath with 3 irregular rows of approxi- mately 7 setae each. Ovipositor 0.9 and Ovipositer sheath 0.65 length of mesoso- ma. Color: Generally brown; mesoscutum, metanotum and occasionally propleuron bright orange yellow; mesopleuron, meta- pleuron, propodeum and _ petiole dark 201 brown; legs white at coxae and gradually darkening to yellow distally; maxillary palp white; face variable, dark brown to light or- ange yellow, usually paler near antennae; scape and pedicel bright yellow with pedi- cel slightly paler; basal 16—18 flageliomeres gradually darkening from yellow to brown distally, followed by 4-7 white flagello- meres, and terminating with 1-3 (usually 1.5) brown flagellomeres; wings hyaline. Body length: 2.1—2.4 mm Male.—Similar to female except as fol- lows: head 1.3—1.4 wider than long; face 1.1-1.2 higher than wide; mesosoma 1.45—-1.55X higher than wide; body 2.3—2.9 mm; fore wing stigma longitudinally bi- sected for most of its length with the thick- ened portion along anterior margin of wing almost completely separated from posterior portion by hyaline streak, the two portions only narrowly joined distally, posterior por- tion broader than anterior portion. Biology.—Unknown Material examined.—Holotype 92°: “Brasilien Nova Teutonia 27°11’B. 5093 Baix21935]) Fritz elaumann™= (BMNH). Paratypes (BMNH, TAMU): 2 6, same data as holotype; 2 ¢, 10 6d, same except viil.1935, x1.1935, 6.211:1936, and 15.%-1935:0 1, 2, 2S, same) locality, but country spelled ‘Brazil: and with B.M Accession label ‘“B.M.1937-656’’, collect- edi livin 1937. ande7.v 1937: 2.2 256 same as preceding, but ““B.M. 1938-682”’, collected ZONVAN933; Livi l93s, 23v51938,and.3:v.1938. Diagnosis.—This species is a member of the cratomorpha species group as defined by Wharton (1994). As in other species of this group, P. hyalina has the second fla- gellomere equal to or barely longer than the first (Fig. 1), a median metanotal projection (Fig. 2), and a hyaline streak longitudinally bisecting the male stigma (Fig. 6). The stig- ma is sexually dimorphic, lacking a well- defined streak in females. Phaenocarpa hy- alina differs from other described species of this group in the possession of a broader stigma that narrows more abruptly basad 202 the junction with r. Further, the metanotal projection of P. hyalina is much more spi- nose than in P. cratomorpha, displaying a thinner, finer apex. Obvious color differenc- es exist between P. sharkeyi and P. hyalina. Phaenocarpa sharkeyi is more uniformly pale yellow in coloration, contrasting with the darker brown portions of the head, meso- and metasoma of P. hyalina. Discussion.—This species is named for the hyaline streak that almost completely bisects the stigma in males, resulting in sep- arate anterior and posterior portions. Addi- tional differences in size and shape between males and females were noted by Wharton (1994) in his description of P. cratomor- pha. Detailed scrutiny of males and females of P. hyalina, however, fails to reveal sig- nificant differences between the sexes for most of these same characters. Since so few individuals of P. cratomorpha were avail- able for study, it is quite possible that the differences recorded by Wharton (1994) in the size and shape of the eye and petiole were simply intraspecific rather than spe- cifically intersexual. Measurements of in- dividuals from the much longer series of P. hyalina demonstrate that these quantitative characters are highly variable within each SEX: Although a distinct hyaline streak is pre- sent in males and absent in females, the stigma in females of P. hyalina exhibits modifications suggesting a morphocline leading to the extreme condition expressed in males. The stigma in females has a slight indication of desclerotization in a position comparable to the basal portion of the hy- aline streak in males. Further, the stigma is exceptionally narrow basally, distad the parastigma, then somewhat abruptly widens where it meets r. In most other species of Phaenocarpa, the posterior margin of the stigma is straight to evenly convex between r and the parastigma. Phaenocarpa hyalina and the species described below are excep- tional in this regard. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Phaenocarpa pericarpa Wharton and Carrejo, new species (Figs. 7, 8, 11, 14-16, 19) Female.—Head: Moderately transverse in dorsal view, about 1.5—1.6 wider than long; eyes strongly bulging beyond tem- ples. Face finely punctate, the punctations separated by much more than their diame- ter, otherwise smooth and polished between base of antenna and frontoclypeal suture; frontoclypeal suture deep and very wide, transversely striate; clypeus exceptionally narrow and strongly protruding, lateral mar- gin of clypeus distinctly separated from an- terior tentorial pit, the latter small, set with- in broad concavity between clypeus and lower margin of eye, the concavity trans- versely striate at least in part. Malar space absent. Frons smooth, polished, evenly and weakly convex, bare except for 2—4 short setae along eye margin laterad ocelli. Ver- tex bare. Mandible 1.4—1.5xX longer than apical width, distinctly expanded apically, apex 1.7—1.85X wider than base, surface with strigose sculpture extending from mid- dle of diagonal ridge to dorsal margin at base of tooth 1; tooth | very broad, orthog- onal, separated by deep cleft from narrowly triangular tooth 2; tooth 2 without dorsal knob; tooth 3 extending distally not quite as far as tooth 1, narrower and more round- ed than the latter. Eye very large, 5.2—-6.0X longer than temple, with a few, scattered, minute setae visible at 50. Antenna 39 segmented; second flagellomere 1.5—1.7X longer than first, 1.1 longer than third. Maxillary palp very long, about twice neight of head. Mesosoma: 1.45—-1.55X longer than high. Pronotum dorsally smooth, polished, but uneven: with small, u-shaped depres- sion medially along anterior margin; weak- ly elevated as a small, rounded bump pos- teromedially; flattened laterally. Mesoscu- tum smooth, polished, with setae confined to a short row along notauli on anterior de- clivity, absent on disc; notauli shallow, weakly sculptured, confined to anterior de- VOLUME 101, NUMBER 1 Figs. 9-16. Antennae showing relative lengths of basal 3 flagellomeres (setal pattern shown only on Fig. 13). 12, P. subtil- istriata. 13, P. heynei. 14, P. pericarpa. 15, P. pericarpa, mandible. 16, P. pericarpa, propodeum. clivity; midpit small, shallow, oval. Scutel- lar sulcus twice as wide as long, or nearly so; with a single median ridge; lateral fields polished, unsculptured. Metanotum with thin, longitudinal flange along midline, the flange sloping more precipitously anteriorly than posteriorly, not elevated above level of scutellum. Propodeum smooth, polished, with well-defined pentagonal areola delim- ited by strong carinae; areola narrow, about half as wide as tall, confined to posterior half of propodeum; anterior half with a strong median carina; lateral carina extend- ing from areola to spiracle usually very weak over lateral half. Sternaulus long, nar- row, Sinuate, complete from anterior margin to mid coxa, weaker posteriorly; crenulate Phaenocarpa spp. 9-11, Faces. 9, P. subtilistriata. 10, P. heynei. 11, P. pericarpa. 12-14, anteriorly, the sculpture weakening poste- riorly and usually absent over posterior 0.3— 0.5. Metapleuron finely punctate but oth- erwise polished and unsculptured over most of surface. Wings: Fore wing stigma weakly con- cave basally along posterior margin, grad- ually widening distally towards junction with r, solid throughout, about 5x longer than width at r, r arising from distal 0.7— 0.75; r very short, 0.15—0.20X length of 2RS; 2RS sharply angled near posterior 0.2; 3RSa 1.25—1.35X longer than 2RS, second submarginal cell gradually narrowing dis- tally; 3RSb ending at wing tip, weakly and evenly bowed; m-cu distinctly antefurcal, with (RS+M)b 0.65-—0.8 length of m-cu; 204 lcu-a postfurcal by 2—3X its length; 2CU interstitial: arising directly in line with ICU. Hind wing very narrow, about 6X longer than wide; 1M 2.0—2.5 x longer than M+CU; 2M very short, angled towards posterior margin, m-cu absent. Metasoma: Petiole 1.3—1.45X longer than apical width; apex 1.8-1.9x wider than base; surface strigose, the sculpture distinct medially, often weak laterally; dor- sal carina strong basally, evanescent at level of spiracles, absent posteriorly; dorsope large and deep. Remaining terga without sculpture. Ovipositor of moderate length, Ovipositor sheath not fully exposed in ma- terial available for examination, but about 2.0—2.3X longer than mesosoma; ovipositor finely tapered to apex, without discernible subapical node or notch. Color: Dark brown; mandible, scape, pedicel, propleuron and petiole variously lighter brown or red-brown in most speci- mens; flagellomeres 1—4 brown, 5—12 and 19—29 dark brown, 14—18 white, and 13 bi- colored brown and dark brown; fore and mid legs and hind coxa and trochanter yel- low to dark yellow, hind femur mostly yel- low with dark spot dorsally over apical 0.3— 0.4, hind tibia and tarsus brown; hypopy- gium apically and apical tergite yellow or yellow brown; palps white. Male.—A single male, probably repre- senting this species, fits the above descrip- tion except as follows: face shorter, about 1.5% wider than high; mandible less ex- panded distally, about 1.8 longer than api- cal width; fore wing Icu-a postfurcal by only about 1.4 its length; and fore wing 3RSa about 1.6X 2RS. Flagellomeres 15 and 16 are dirty white, with remaining fla- gellomeres dark brown. Biology.—Reared from puparia of An- astrepha distincta Greene in pods of guama (Inga sp.: Fabaceae). Material examined.—Holotype @: “CO- LOMBIA Dept. del Valle del Cauca Mun. Buenaventura, Corregimiento de Zacarias 23.111.1994 N. Carrejo ex Anastrepha dis- tincta on Guama’”’ Deposited in La Univer- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sidad del Valle, Museo de Entomologia. Paratypes: 3 @, same data as holotype (TAMU; Universidad del Valle; and Insti- tuto de Ciencias Naturales, Universidad Nacional de Colombia). Additional material (not a paratype): 1 36, VENEZUELA, Ar- agua, Parque Nacional Henri Pittier, Ran- cho Grande, 1,100m, 12.1.1996, R. Wharton (TAMU). Diagnosis.—As in both P. heynei and P. subtilistriata, this species has reduced no- tauli, a short first flagellomere, and a large second submarginal cell. It differs from both of these species by the possession of a shorter ovipositor (compare Fig. 19 with Figs. 17, 18) which lacks a subapical node or notch and a broader, more discrete tooth 1 on the mandible (Fig. 15). The ovipositor is more than three times longer than the me- sosoma in P. heynei and P. subéilistriata. The shape of the mandible is sufficient for separating P. pericarpa from all other de- scribed New World species of Phaenocar- pa. Discussion.—Papp (1969) adequately differentiated P. heynei from P. subtilistria- ta, noting especially the difference in shape of the petiole. There are also slight differ- ences in sculpture between the holotypes of P. heynei and P. subtilistriata. The propo- deal areola is essentially obliterated in P. subtilistriata, with the posterior face stri- gose or weakly rugulose below the well- developed and complete transverse carina. A weak areola is present in P. heynei, and the transverse carina is incomplete, not reaching the propodeal spiracle. The ster- naulus is also broader and more heavily sculptured in P. subtilistriata but the scu- tellar sulcus is smooth with a single median carina. In P. heynei, the scutellar sulcus is weakly sculptured on either side of the me- dian sulcus. Although both P. heynei and P. subtilistriata are known only from the holotype, and variation thus cannot be as- sessed, these same sculptural features show little variation in the four specimens of P. pericarpa. Thus, it is likely that these rel- atively minor sculptural differences will be VOLUME 101, NUMBER I 205 Figs. 17-19. carpa. useful for species-level recognition of P. heynei and P. subtilistriata. Several features suggest that P. heynei shares a sister group relationship with P. pericarpa relative to P. subtilistriata. A\- though the clypeus is narrow in all three species, it is more strongly narrowed and protruding in P. heynei and P. pericarpa (Figs. 10, 11), with consequent detachment of the anterior tentorial pit from the lateral margin of the clypeus. Similarly, 2M in the hind wing is shorter and more distinctly de- flected posteriorly in P. heynei and P. peri- carpa than in P. subtilistriata, which retains Meso- and metasoma, lateral view. 17, Phaenocarpa subtilistriata. 18, P. heynei. 19, P. peri- the more plesiomorphic form of a longer, more distally-directed 2M. The fore wing stigma of P. heynei is also more similar to the unusually shaped stigma of P. pericarpa than the more typically shaped stigma of P. subtilistriata. Phaenocarpa subtilistriata appears to form a link between the hey- neit+pericarpa sister group and the more typical Phaenocarpa species from the Hol- arctic Region. This hypothesized relation- ship between P. heynei and P. pericarpa leads the suggestion that the Gnathopleura- like mandible of P. heynei and P. subtilis- triata iS a more primitive feature from 206 which the P. pericarpa configuration was derived. Despite similarities in the mandi- ble, P. heynei and P. subtilistriata can be readily separated from Gnathopleura since the latter has a much smaller second sub- marginal cell. The putative relationship between P. hey- nei and P. pericarpa is based on features which, though unusual, are found elsewhere in Phaenocarpa and related genera. A sim- ilarly narrow hind wing with reduced 2M is found in the Phaenocarpa cratomorpha species group, for example. A narrow, strongly protruding clypeus is found in at least two species of Asobara (one from Bra- zil and one from Papua New Guinea) which otherwise lack shared derived features. In all of these cases, the character states in question are hypothesized as independent derivations. This is the first host record for any of the Neotropical species of Phaenocarpa. The type series of P. pericarpa was reared from the tephritid fly Anastrepha distincta Greene, developing in the pods of guama (Inga sp.). Relatively few tephritids have been recorded as hosts of Alysiinae (Whar- ton 1984). Only one, Asobara anastrephae (Muesebeck), has been verified as a para- sitoid of fruit-infesting tephritids. Asobara orientalis Viereck, originally described from material thought to be reared from da- cine tephritids, is undoubtedly a drosophilid parasitoid. Three other alysiine species have been reared either from tephritids in flower heads or other plant parts, and at least four additional species have been reared from unknown hosts in fruit. Asobara anastre- phae is interesting because of its exception- ally large size and unusual host preferences relative to other species of Asobara, which are primarily drosophilid parasitoids. Aso- bara anastrephae belongs to a group of Neotropical species with typical Asobara fore wing venation, loss of hind wing cu-a, a short, broad petiole, and brightly colored bodies (Muesebeck 1958, Wharton 1994). One of the derived members of this group, as yet undescribed, exhibits the same clyp- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON eal modifications found in P. pericarpa and P. heynei. ACKNOWLEDGMENTS We are grateful to Tom Huddleston (for- merly BMNH) for extending many cour- tesies regarding material in his care, to Frank Koch (Humboldt Museum, Berlin), Jend Papp (Hungarian Natural History Museum), and Paul Marsh and David Smith (Systematic Entomology Laborato- ry, USDA) for the loan of type material and assistance with type collections. Barry Flahey provided the illustrations for P. hy- alina. This work was supported in part by the Texas Agricultural Experiment Station and by the National Science Foundation (DEB: 9300517). LITERATURE CITED Achterberg, C. van. 1988. The genera of the Aspilota- group and some descriptions of fungicolous Aly- siini from the Netherlands (Hymenoptera: Bracon- idae: Alysiinae). Zoologische Verhandelingen 247: 1-88. Fischer, M. 1971. Untersuchungen tber die Euro- paischen Alysiini mit besonderer Berucksichti- gung der Fauna Niederosterreichs. Polskie Pismo Entomologiczne 41: 19-160. . 1974. Die nearktischen Phaenocarpa-Arten. Revision der Gruppe B (Hymenoptera, Braconi- dae, Alysiinae). Polskie Pismo Entomologiczne 44: 103-230. . 1975. Die nearktischen Phaenocarpa-Arten. Revision der Gruppe A (Hymenoptera, Braconi- dae, Alysiinae). Polskie Pismo Entomologiczne 45: 279-356. . 1990. Westpalaarktische Phaenocarpa-Arten: Vorlaufiger Bestimmungsschliissel, Deskriptionen und Redeskriptionen (Hymenoptera, Braconidae, Alysiinae). Annalen Naturhistorische Museum in Wien 91: 105-135. . 1993. Einige Phaenocarpa-Wespen aus der Alten Welt: Redeskriptionen und Stellung in ei- nem vergleichenden System (Hymenoptera, Bra- conidae, Alysiinae). Linzer biologische Beitrage 25: 511-563. . 1994. Beitrag zur Kenntnis der Keiferwespen der Welt (Hymenoptera, Braconidae, Alysiinae: Alysiini). Linzer biologische Beitrage 26: 763- 806. Muesebeck, C. EK W. 1958. New Neotropical wasps of the family Braconidae (Hymenoptera) in the U.S. National Museum. Proceedings of the United States National Museum 107: 405—461. VOLUME 101, NUMBER 1 Papp, J. 1966. New Phaenocarpa Forster species from the Ethiopian Region (Hymenoptera, Braconidae). Acta Zoologica Academiae Scientiarum Hungari- cae 12: 133-144. . 1969. A synopsis of the Phaenocarpa Forst. species of the Neotropic Region (Hymenoptera: Braconidae, Alysiinae). Acta Zoologica Acade- miae Scientiarum Hungaricae 15: 379-389. . 1972. Phaenocarpa impugnata sp. n. (Hym., Braconidae: Alysiinae), a new reared species from Denmark. Zoologischer Anzeiger, Leipzig 188: 52-56. Sharkey, M. J. and R. A. Wharton. 1997. Morphology and terminology, pp. 19-63. Jn Wharton, R. A., P. M. Marsh and M. J. Sharkey, eds., Manual of the New World genera of the family Braconidae (Hymenoptera). Special Publication No. 1 of the International Society of Hymenopterists. Shenefelt, R. D. 1974. Pars 11, Braconidae 7 Alysi- inae, pp. 937-1113. Jn van der Vecht, J. and R. D. Shenefelt, eds., Hymenopterorum Catalogus (nova editio). Dr. W. Junk, The Hague. 207 Tobias, V. I. 1986. Subfamily Alysiinae, pp. 100-231. In Identification of Insects of European USSR, Vol. III, Part V, Hymenoptera, Braconidae. Aka- demia Nauk, Leningrad (in Russian). Vet, L. E. M. and J. J. M. van Alphen. 1985. A com- parative functional approach to the host detection behaviour of parasitic wasps. 1. A qualitative study on Eucoilidae and Alysiinae. Oikos 44: 478-486. Wharton, R. A. 1980. Review of the Nearctic Alysiini (Hymenoptera, Braconidae) with discussion of ge- neric relationships within the tribe. University of California Publications in Entomology 88: 1-112. . 1984. Biology of the Alysiini (Hymenoptera: Braconidae), parasitoids of cyclorrhaphous Dip- tera. Texas Agricultural Experiment Station Tech- nical Monograph 11: 1-39. . 1994. New genera, species, and records of New World Alysiinae (Hymenoptera: Braconi- dae). Proceedings of the Entomological Society of Washington 96: 630—664. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 208-211 THE SMALL MINNOW MAYFLY GENUS CLOEODES TRAVER (EPHEMEROPTERA: BAETIDAE) IN MADAGASCAR C. R. LuUGo-OrtTIz, W. P- MCCAFFERTY, AND J.-L. GATTOLLIAT (CRL, WPM) Department of Entomology, Purdue University, West Lafayette, IN 47907, U.S.A. (e-mail: carlos_lugo-ortiz@entm.purdue.edu); (JLG) Museum of Zoology, P.O. Box 448, CH 1000 Lausanne, Switzerland Abstract.—Cloeodes portabilis, new species, represents the first report of Cloeodes from Madagascar. The species is distinguished in the larval stage by the relatively wide anteromedial emargination of the labrum, presence of tufts of fine, simple setae between the prosthecae and molae of the mandibles, slightly distolaterally acute segment 3 of the labial palps, and abdominal color pattern. The presence of Cloeodes throughout the South- ern Hemisphere suggests a relatively ancient origin among extant Baetidae. Key Words: The distinctive small minnow mayfly ge- nus Cloeodes Traver (Ephemeroptera: Bae- tidae) has been known from the Afrotrop- ics, Neotropics, Orient, and southwestern Nearctic (Traver 1938, Waltz and Mc- Cafferty 1987ab, 1994, Kluge 1991, Flow- ers 1991, Lugo-Ortiz and McCafferty 1993, 1994, 1995, McCafferty and Lugo-Ortiz 1995, McCafferty et al. 1997). Most re- cently, Lugo-Ortiz and McCafferty (1998) reported Cloeodes from Australia, signifi- cantly extending its known range. Three historic biogeographic hypotheses were provided to explain its essentially Pantrop- ical distribution. That Cloeodes was well established throughout Gondwanaland or at least West Gondwanaland (the South American-Afri- can-Malagasy-Indian landmass) prior to 100 million years ago, before Madagascar began to separate from Africa and India, were hypotheses that clearly predicted that Cloeodes would occur in Madagascar. Herein we report Cloeodes from Madagas- car for the first time based on a new species described from larvae. An additional spe- Ephemeroptera, Baetidae, Cloeodes portabilis, new species, Madagascar cies of Cloeodes from the island will be de- scribed elsewhere by one of us (J.-L.G.). The specimens studied are housed in the Purdue Entomological Research Collection, West Lafayette, Indiana. Cloeodes portabilis Lugo-Ortiz and McCafferty, new species (Figs. 1-10) Larva.—Body length: 3.4—4.4 mm. Cau- dal filaments length: 1.8—2.0 mm. Head: Coloration medium brown, with no distinct pattern. Antenna approximately 1.75 length of head capsule. Labrum (Fig. 1) with wide anteromedial emargination, sub- medial pair of long, fine, simple setae, and submarginal row of four to six long, fine, simple setae. [Left and right mandibles (Figs. 2, 3) with incisors worn down.] Left mandible (Fig. 2) with one set of incisors; prostheca robust, apically denticulate; tuft of fine, simple setae present between pros- theca and mola. Right mandible (Fig. 3) with two sets of incisors; prostheca slender, apically bifid; tuft of fine, simple setae pre- sent between prostheca and mola. Maxilla VOLUME 101, NUMBER 1 209 10 Figs. 1-10. Cloeodes portabilis. 1, Labrum (dorsal). 2, Left mandible. 3, Right mandible. 4, Left maxilla. 5, Labium (left-ventral; right-dorsal). 6, Left foreleg. 7, Abdomen (dorsal). 8, Tergum 3 (detail). 9, Gill 3. 10, Paraproct. (Fig. 4) with four small, stout denticles on crown of galealacinia; four to five long, fine, simple setae near medial hump; palp and paraglossa equal in length; palp seg- ment 1 approximately 0.80 length of seg- ments 2 and 3 combined; segment 2 ap- not reaching galealacinia; palp segment | subequal in length to segment 2; segment 2 apically acute. Labium (Fig. 5) with glossa proximately 1.6 length of segment 3; seg- ment 3 falcate apically and slightly pointed distolaterally. Thorax: Coloration yellow 210 brown, with complex markings. Hindwing- pads absent. Legs (Fig. 6) pale yellow brown; femora dorsally with row of five to six medium-sized, robust, simple setae in- termixed with numerous long, fine, simple setae; tibiae ventrally with 10—12 short, stout, simple setae; tarsi ventrally with eight to nine short, stout, simple setae. Abdomen (Fig. 7): Coloration medium to yellow brown; segment | yellow brown; segment 2 yellow brown, with submedial medium brown subtriangular marking and oblique medium brown distolateral markings; seg- ment 3 with submedial medium brown cres- centlike marking posteriorly and oblique medium brown distolateral markings; seg- ment 4 with submedial medium brown cres- centlike marking posteriorly; segment 5 with large, wavy, medium brown marking posteriorly; segment 6 with submedial spikelike medium brown marking and oblique medium brown distolateral mark- ings; segment 7 with small, oblong, medi- um brown anterolateral markings; segment 8 yellow brown, with no markings; seg- ments 9-10 medium brown, with no mark- ings. Terga (Fig. 8) with abundant scale ba- ses, and with posterior marginal spines tri- angular, approximately as long as_ basal width. Sterna yellow brown. Gill (Fig. 9) subtriangular, well tracheated, marginally smooth. Paraproct (Fig. 10) with 10—12 spines, increasing in size apically. Caudal filaments pale yellow brown; medial caudal filament subequal in length to cerci. Adult.—Unknown. Material examined.—Holotype: Larva, MADAGASCAR, Tamatave (= Toamasi- na) Prov., stream at Gri-Gri, RN 2, 17-X- 19713 (G. ES GC; Edmunds; and | Em- manuel. Paratypes: Two larvae, same data as holotype; five larvae, MADAGASCAR, Antsiranana Prov., Djabala R., 11 km NW of Hell-Ville, Nosy Be, 25-X-1971, G. F, C. H. Edmunds, and EF Emmanuel [mouth- parts, left foreleg, tergum 3, gills 3, and paraproct of one larva mounted on slide (medium: Euparal)]. Additional material: Four larvae, same data as holotype; twenty PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON larvae, MADAGASCAR, Antsiranana Prov., Djabala R., 11 km NW of Hell-Ville, Nosy Be, 25-X-1971, G. E, C. H. Edmunds, and K Emmanuel. Etymology.—The specific epithet is a Latin word meaning “that which may be carried.”’ It is an allusion to the species be- ing a drifted representative of the genus. Discussion.—Cloeodes portabilis is dis- tinguished from other members of the ge- nus by the relatively wide anteromedial emargination of the labrum (Fig. 1), pres- ence of tufts of fine, simple setae between the prosthecae and molae of the mandibles (Figs. 2, 3), slightly distolaterally acute seg- ment 3 of the labial palps (Fig. 5), and ab- dominal color pattern (Fig. 7). Cloeodes portabilis is unique among Eastern Hemisphere members of the genus because segment 3 of the labial palps is somewhat falcate, being slightly distolater- ally pointed (Fig. 5); other species have a bulbous segment 3. Only the South Amer- ican species C. hydation McCafferty and Lugo-Ortiz has a similar labial palp mor- phology (McCafferty and Lugo-Ortiz 1995: Fig. 6); we can only assume that it consti- tutes a homoplasy. In addition, the presence of a tuft of setae between the prosthecae and molae of the mandibles in C. portabilis (Figs. 2, 3) is exceptional in Cloeodes, pos- sibly indicating a relatively ancestral posi- tion within the genus, because that charac- teristic is generally associated with plesi- otypic genera in Baetidae (R. D. Waltz, per- sonal communication). ACKNOWLEDGMENTS We thank G. E Edmunds, Jr., Salt Lake City, Utah, for the donation of the material used in this study. This paper has been as- signed Purdue Agricultural Research Pro- gram Journal No. 15655. LITERATURE CITED Flowers, R. W. 1991. Diversity of stream-living in- sects in northwestern Panama. Journal of the North American Benthological Society 10: 322-334. VOLUME 101, NUMBER 1 Kluge, N. 1991. Cuban mayflies of the family Bae- tidae (Ephemeroptera) 1. Genera Callibaetis, Cloeodes, and Paracloeodes. Zoologischeskiy Zhurnal 12: 128—136. [in Russian] Lugo-Ortiz, C. R. and W. P. McCafferty. 1993. Gen- era of Baetidae (Ephemeroptera) from Central America. Entomological News 104: 193-197. . 1994. New records of Ephemeroptera from Mexico. Entomological News 105: 17-26. . 1995. Annotated inventory of the mayflies (Ephemeroptera) of Arizona. Entomological News 106: 131-140. . 1998. First report and new species of the genus Cloeodes (Ephemeroptera: Baetidae) from Australia. Entomological News 109: 122— 128. McCafferty, W. P. and C. R. Lugo-Ortiz. 1995. Cloeodes hydation, n. sp. (Ephemeroptera: Bae- tidae): an extraordinary, drought tolerant may- 211 fly from Brazil. Entomological News 106: 29— 355 McCafferty, W. P., C. R. Lugo-Ortiz, and G. Z. Ja- cobi. 1997. The mayfly fauna of New Mexico. Great Basin Naturalist 57: 283-314. Traver, J. R. 1938. Mayflies of Puerto Rico. Journal of Agriculture of the University of Puerto Rico 22: 5-42. Waltz, R. D. and W. P. McCafferty. 1987a. Generic revision of Cloeodes and description of two new genera (Ephemeroptera: Baetidae). Pro- ceedings of the Entomological Society of Wash- ington 89: 177-184. 1987b. Revision of the genus Cloeodes Traver (Ephemeroptera: Baetidae). Annals of the Entomological Society of America 80: 191— 207. . 1994. Cloeodes (Ephemeroptera: Baetidae) in Africa. Aquatic Insects 16: 165-169. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 212-218 NEW COMBINATIONS, NEW SYNONYMY, AND HYMONOMY IN THE ERIOCOCCIDAE, NEW HOMONOMY AND SYNONYMY IN THE CEROCOCCIDAE, AND TRANSFER OF CANCEROCOCCUS KOTEJA TO THE MARGARODIDAE (HEMIPTERA: COCCOIDEA) DOUGLASS R. MILLER AND MAREN E. GIMPEL Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, Bldg. 046, BARC-W, Beltsville, MD 20705, U.S.A. (e-mail: dmiller@sel.barc.usda.gov) Abstract.—A database and catalog of the eriococcid and cerococcid scale insects of the world is nearly complete and soon will be in press and placed on the World Wide Web. Before this is done, new combinations and other taxonomic changes need to be validated in print. This publication includes Neokaweckia Tang and Hao as a new synonym of Eriococcus, proposal of Neotrichococcus as a new name for Trichococcus Borchsenius, and new combinations in the family Eriococcidae; a new homonym and synonym in the Cerococcidae; and transfer of Cancerococcus from Eriococcidae to Margarodidae. Key Words: felt scales, Coccoidea, Eriococcidae, Cerococcidae, Margarodidae, ScaleNet, catalog, new combinations, Internet We currently are finishing a database and manuscript on the Eriococcidae and Cero- coccidae of the world. This research is part of a larger project called ‘‘ScaleNet”’ to de- velop a systematic database of the Coccoi- dea of the World; see Miller and Gimpel (1996), and Ben-Dov et al. (1997). One of the most controversial subjects in synthe- sizing the systematic data on eriococcids is to make sense out of the genera Acantho- coccus Signoret, Anophococcus Balachow- sky, Eriococcus Targioni Tozzetti, Gossy- paria Signoret, Greenisca Borchsenius, Gregoporia Danzig, Kaweckia Koteja and Zak-Ogaza, Neokaweckia Tang and Hao, and Rhizococcus Signoret. Most European literature recognizes all, or nearly all, of these genera as distinct, e.g., Kosztarab and Kozar (1988), but Hoy (1963) treated many of them as junior synonyms of Eriococcus and Williams (1985) treated all but Gre- goporia as members of Eriococcus. Gre- goporia was first treated as a synonym by Miller and Gimpel (1996) and Neokaweckia is here considered as a new junior synonym of Eriococcus. Although some have criticized the lump- ing of these genera as a reversion to Lin- naean times (Koteja 1997), we believe that the characters used to define these genera are homoplasious and discriminate artificial groups. It is logical to assume that natural groups occur in this assemblage, but it is important to undertake a careful phyloge- netic analysis to discover the groupings. Some cladistic work using molecular and morphological character systems is under- way in this regard by P. J. Gullan and Lyn Cook (Division of Botany and Zoology, Australian National University, Canberra). Their preliminary findings have been quite interesting and suggest the strong possibil- ity that there will be several genera within what is here treated as Eriococcus (Gullan, VOLUME 101, NUMBER 1 personal communication June 17, 1998). Unfortunately, this important research has been underway for only a short period of time and currently includes a small sample of the world eriococcid fauna. For many years, the first author surmised that the unusually large tubular ducts pre- sent on E. buxi (Fonscolombe) and E. eu- calypti Maskell were sufficient to charac- terize Eriococcus as separate from Acantho- coccus (see Miller and Williams 1976). However, it now appears that this is not necessarily the case and for the purposes of this paper we accept the conservative view pending results of the research by Gullan and her colleagues. This conservative view is consistent with the research of Ferris (1957), Hoy (1963), and Williams (1985) and places all or most of the questionable genera in the genus Eriococcus. With this in mind, it is necessary to move several species previously placed in Acanthococcus into the genus Eriococcus for the first time. NEW COMBINATIONS IN THE ERIOCOCCIDAE Eriococcus abaii (Danzig), n. comb. Acanthococcus abaii Danzig 1990: BBE Eriococcus actius (Miller & Miller), n. comb. Acanthococcus actius Miller & Miller 1993: 9. Eriococcus adzharicus (Hadzibejli), n. comb. Acanthococcus adzharicus Hadzibejli 1960: 310. Eriococcus arenariae (Miller & Miller), n. comb. Acanthococcus arenariae Miller & Miller 1993: 13. Eriococcus barri (Miller), n. comb. Acanthococcus barri Miller 1991: 337. Eriococcus beshearae (Miller & Miller), n. comb. Acanthococcus beshearae Miller & Miller 1993: 15. Eriococcus brachypodii (Borchsenius & Danzig), n. comb. i) eo.) Greenisca brachypodii Borchsenius & Danzig 1966: 43. Eriococcus centaureae (Savescu), n. comb. Acanthococcus centaureae Savescu L985; 122: Eriococcus danzigae (Miller & Gimpei), n. comb. Rhizococcus confusus Danzig 1962a: 854 (junior secondary homonym). Acanthococcus danzigae Miller & Gimpel 1996: 600 (replacement name). Remarks: The replacement name A. dan- zigae was given for Rhizococcus confusus Danzig (1962a) when it was transferred to Acanthococcus by Miller & Gimpel (1996) making it a junior secondary homonym of A. confusus (Maskell) (1892). The species epithet danzigae must continue to be used when transferred to Eriococcus, since A. confusus (Maskell) also is placed in Erio- coccus and is the senior homonym. Eriococcus davidsoni (Miller & Miller), n. comb. Acanthococcus davidsoni Miller & Miller 1993: 25. Eriococcus dennoi (Miller & Miller), n. comb. Acanthococcus dennoi Miller & Miller 1993: 27. Eriococcus droserae (Miller, Liu, and How- ell), n. comb. Acanthococcus droserae Miller, Liu, and Howell 1992: 512. Eriococcus epacrotrichus (Miller & Mil- ler), n. comb. Acanthococcus epacrotrichus Miller & Miller 1992: 33. Eriococcus evelinae (Kozar), n. comb. Rhizococcus evelinae Kozar 1983: 144. Acanthococcus evelinae (Kozar); Mill- er & Gimpel 1996: 600. Eriococcus froebeae (Miller), n. comb. Acanthococcus froebeae Miller 1991: 343. Eriococcus herbaceus (Danzig), n. comb. Rhizococcus herbaceus Danzig 1962b: 9) Acanthococcus herbaceus; Nast et al. 199027120; Acanthococcus herbaceus; Terezniko- val 198125295 Eriococcus hoyi (Miller & Miller), n. comb. Acanthococcus hoyi Miller & Miller 1992: 44. Eriococcus iljiniae (Danzig), n. comb. Rhizococcus iljiniae Danzig 1972: 339) Acanthococcus iljiniae; Miller & Gim- pel 1996: 601. Eriococcus istriensis (Kozar), n. comb. Gregoporia istriensis Kozar 1983: 142. Acanthococcus istriensis; Miller & Gimpel 1996: 601. Eriococcus korotyaevi (Danzig), n. comb. Acanthococcus korotyaevi Danzig 1982: 145. Eriococcus laeticoris (Tereznikova), n. comb. _ Greenisca 1965: 975. Kaweckia laeticoris; Koteja & Zak- Ogaza 1981: 507. Neokaweckia laeticoris; Tang & Hao 19953514. Acanthococcus laeticoris; Miller & Gimpel 1996: 601. Eriococcus leptoporus (Miller & Miller), n. comb. Acanthococcus leptoporus Miller & Miller 1993-539: Eriococcus mackenziei (Miller & Miller), n. comb. Acanthococcus mackenziei Miller & Miller 1992: 60. Eriococcus macrobactrus (Miller & Mil- ler), n. comb. Acanthococcus macrobactrus Miller & Miller 1992: 62. Eriococcus matesovae (Miller & Gimpel), n. comb. Acanthococcus multispinosus Mateso- va 1976: 24 Gunior secondary hom- onym). laeticoris Tereznikova PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Acanthococcus matesovae Miller & Gimpel 1996: 600 (replacement name). Remarks: The replacement name A. ma- tesovae was given for Acanthococcus mul- tispinosus Matesova (1976) when it was transferred to Acanthococcus by Miller & Gimpel (1996) making it a junior secondary homonym of A. multispinosus Kuhlgatz (1898). The species epithet matesovae must continue to be used when transferred to Er- iococcus, since A. multispinosus Kulgatz also is placed in Eriococcus and is the se- nior homonym. Eriococcus megaporus (Miller & Miller), n. comb. Acanthococcus megaporus Miller & Miller 1993: 45. Eriococcus mesotrichus (Miller & Miller), n. comb. Acanthococcus mesotrichus Miller & Miller 1993: 48. Eriococcus microtrichus (Miller & Miller), n. comb. Acanthococcus microtrichus Miller & Miller 1992: 65. Eriococcus minimus (Tang & Li), n. comb. Acanthococcus minimus Tang & Li 1988: 71. Rhizococcus minimus; Tang & Hao 19952352: Eriococcus monotrichus (Miller & Miller), n. comb. Acanthococcus microtrichus Miller & Miller 1993: 54. Eriococcus multispinatus (Tang & Hao), n. comb. Rhizococcus multispinatus Tang & Hao 1995: 598. Acanthococcus multispinatus; Miller & Gimpel 1996: 602. Eriococcus oligacanthus (Danzig), n. comb. Rhizococcus oligacanthus Danzig 1972> 341. Acanthococcus oligacanthus; Miller & Gimpel 1996: 602. VOLUME 101, NUMBER 1 Eriococcus oligotrichus (Miller & Miller), n. comb. Acanthococcus oligotrichus Miller & Miller 1993: 57. Eriococcus ophius (Miller & Miller), n. comb. Acanthococcus ophius Miller & Miller 1993559: Eriococcus orientalis (Borchsenius), n. comb. Greenisca orientalis Borchsenius 1956:,676. Kaweckia orientalis (Borchsenius); Tang & Hao 1995: 511. Acanthococcus orientalis (Borchsen- ius); Miller & Gimpel 1996: 602. Eriococcus oxyacanthus (Danzig), n. comb. Acanthococcus oxyacanthus Danzig INES 3). Rhizococcus oxyacanthus; Walter 1985: 75. Eriococcus rubrus (Matesova), n. comb. Greenisca rubra Matesova 1960: 209. Kaweckia rubra; Koteja & Zak-Ogaza 1981: 508. Neokaweckia rubra; Tang & Hao 1995: 515: Acanthococcus rubra; Miller & Gim- pel 1996: 603. Eriococcus salicicola Tang, nomen nudum Kozar & Remarks: Tang (1984) indicated that this species occurs widely over northeastern China on willow. He stated that he would be describing the species as new in the fu- ture, but there is no record of its publica- tion. Eriococcus stauroporus (Miller & Miller), n. comb. Acanthococcus stauroporus Miller & Miller 1992: 82. Eriococcus tosotrichus (Miller & Miller), n. comb. Acanthococcus tosotrichus Miller & Miller 1993: 62. Eriococcus washingtonensis (Miller & Miller), n. comb. Acanthococcus washingtonensis Miller & Miller 1992: 90. 2S Eriococcus whiteheadi (Miller), n. comb. Acanthococcus 1991: 350. Eriococcus zernae (Tereznikova), n. comb. Acanthococcus zernae Tereznikova O72 Sak. whiteheadi Miller NEW GENERIC SYNONYMY IN THE ERIOCOCCIDAE Neokaweckia Tang & Hao 1995: 596, new synonymy Type species: Greenisca rubra Matesova 1960, by monotypy and original designa- tion Remarks: This genus is characterized by having a small anal ring, truncate body se- tae that are restricted to the last abdominal segments, and dorsal cruciform pores. These characters are considered to be with- in the expected range of variation for the genus Eriococcus. NEw GENERIC HOMONOMY AND REPLACEMENT NAME IN THE ERIOCOCCIDAE Neotrichococcus Miller & Gimpel, new replacement name Trichococcus Borchsenius 1948: 503. Type species: Trichococcus filifer Borch- senius 1948, by monotypy and original des- ignation. Remarks: Trichococcus Borchsenius is a junior homonym of Trichococcus Kanda (1941) which is now considered to be a ju- nior synonym of Beesonia Green. Morrison and Morrison (1966) first discovered this homonymy but did not provide a replace- ment name. Neotrichococcus filifer (Borchsenius), n. comb. Trichococcus filifer Borchsenius 1948: 503. FAMILY TRANSFER OF CANCEROCOCCUS TO THE MARGARODIDAE Cancerococcus Koteja 1988: 412, new family assignment 216 Remarks: This monotypic genus was originally placed in the Eriococcidae pre- sumably because of the enlarged setae on the antennae, a character found in many er- iococcid males. However, we have discov- ered that the genus is most closely related to the Pityococcini genera Pityococcus (McKenzie 1942) (including the species P. ferrisi McKenzie, P. deleoni McKenzie, P. rugulosus McKenzie) and Desmococcus (D. captivus McKenzie and D. sedentarius McKenzie) and is here transferred to the Margardodidae. The description of Cancer- ococcus (Koteja 1988) is based on a single wingless male from amber. The illustrations and description provided by Koteja were compared with a single undetermined male of Pityococcus deposited in the collection of National Museum of Natural History, Beltsville, Maryland. The following simi- larities occur in both taxa: Numerous short setae on the antennae; penial sheath that is apically bifurcate and has a broad aedeagus; antennae with short, round segments; eyes apparently numbering 4 (C. apterous) or 5 (Pityococcus sp.) on each side of the head and set on a plate. This combination of characters is unique to the Pityococcini in the Margarodidae. NEw SYNONYMY IN THE CEROCOCCIDAE Asterococcus ramakrishnai (Ramakrishna Ayyar) Cerococcus ramakrishnae Ramachan- ran & Ramakrishna Ayyar 1934: 86. (nomen nudum) Cerococcus ramakrishnae Ramakrish- na Ayyar 1936: 148. Asterococcus ramakrishnai Lambdin 1983: 304-306, new homonymy and synonymy. Remarks: The original combination of Cerococcus ramakrishnae is an unpub- lished manuscript name of Green. Rama- chanran & Ramakrishna Ayyar (1934) cited the name but gave no description thus cre- ating a nomen nudum. Ramakrishna Ayyar (1936) did not realize that he was validating PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the name and describing it after himself, but this was the case. Lambdin (1983) appar- ently knew of Green’s manuscript name, but did not realize that Ramakrishna’s de- scription was valid and described the spe- cies as new. Lambdin also corrected the spelling of the species epithet from “‘ra- makrishnae’’ to ‘‘ramakrishnaw’ ’ and moved the species from Cerococcus to As- terococcus. From the syntypes of Cerococ- cus ramakrishnae, we have chosen and marked as lectotype an adult female labeled in Green’s handwriting: “on Ficus rootlets/ India, (Coimbatore)/ coll. Ramakrishna/ 24/ 31. No. 335.”’ (BMNH). The slide contains three specimens; the center specimen is the lectotype. There are three paralectotypes. ACKNOWLEDGMENTS We thank the following individuals for their comments and suggestions relative to improvements of this manuscript: Douglas J. Williams, Department of Entomology, The Natural History Museum, London, UK and Michael E. Schauff, Systematic Ento- mology Laboratory, Agricultural Research Service, USDA, c/o National Museum of Natural History, Washington, DC. Funding for this paper was partially supported by BARD grant IS-2605. 95CR and is grate- fully acknowledged. LITERATURE CITED Ben-Dovy, Y., C. J. Hodgson, and D. R. Miller. 1997. Changes and comments on the taxonomy and no- menclature of some taxa in the families Coccidae, Eriococcidae and Pseudococcidae (Homoptera: Coccoidea). Phytoparasitica 25: 199-206. Borchsenius, N. S. 1948. On the revision of the genus Eriococcus Sign. (Insecta, Homoptera, Coccoi- dea). (In Russian). Doklady Akademii Nauk SSSR. Moscow (n.s.) 60: 501-503. . 1956. Notes on the Coccoidea of Korea. En- tomologicheskoe Obozrenye 35: 671-679. Borchsenius, N. S. and E. M. Danzig. 1966. A new species of Greenisca Borchs. (Homoptera, Coc- coidea, Eriococcidae) from the USSR, /n Bykhov- skii, B. E., New Species of Insects of the Fauna of the USSR and Adjacent Countries. Trudy Aka- demii Nauk SSR Zoologicheskogo Instituta 37: 41-44. Danzig, E. M. 1962a. Revision of the genus Rhizococ- VOLUME 101, NUMBER 1 cus Signoret (Homoptera, Coccoidea) of the SSR fauna. Entomologicheskoe Obozrenye 41: 839- 860. . 1962b. Addition to the scale insect fauna (Ho- moptera, Coccoidea) of the Leningrad region. Tru- dy Akademii Nauk SSR Zoologicheskogo Insti- tuta. St. Petersburg 31: 22—24. . 1972. Contribution to the fauna of the white flies and scale insects (Homoptera: Aleyrodoidea, Coccoidea) of Mongolia. Insects of Mongolia 1: 325-348. 1975. Species of mealy-bugs (Homoptera, Coccoidea, Pseudococcidae) new for Mongolia. Insects of Mongolia 3: 48-55. . 1982. New species of the scale insects (Ho- moptera, Coccinea) from Mongolia. Insects of Mongolia 8: 140-147. . 1990. New species of coccids (Homoptera, Coccinea) from Iran, Mongolia and Vietnam. En- tomologicheskoe Obozrenye 69: 373-376. Ferris, G. EF 1957. A review of the family Eriococcidae (Insecta: Coccoidea). Microentomology 22: 81— 89. Hadzibejli, Z. K. 1960. New species of coccids (Ho- moptera, Coccoidea) from Georgia. Trudy Aka- demii Nauk Gruzinskoy SSR Instituta Zashchitii Rastenii 13: 299-321. Hoy, J. M. 1963. A catalogue of the Eriococcidae (Ho- moptera: Coccoidea) of the world. New Zealand Department of Scientific and Industrial Research Bulletin 150: 1—260. Kanda, S. 1941. The genus Xylococcus and the genus Trichococcns [sic] N. G. from Japan (Homopteta) [sic]. Insect World 44: 68-72. Kosztarab, M. and F Kozar. 1988. Scale Insects of Central Europe. Akadémiai Kiado, Budapest. 456 Pp. Koteja, J. 1988. Two new eriococcids from Baltic Am- ber. Deutsche entomologische Zeitschrift 35: 405— 416. . 1997. With Microsoft (through Windows) into the XVIII Century. Inclusion Wrostek 26: 17. Koteja, J. and B. Zak-Ogaza. 1981. Kaweckia gen. n. in the Eriococcidae (Homoptera, Coccoidea) and notes on related genera. Acta Zoologica Cracoy- iensia 25: 501-518. Kozar, F 1983. New and little-known scale-insect spe- cies from Yugoslavia (Homoptera: Coccoidea). Acta Zoologica Academiae Scientiarum Hungari- cae 29: 139-149. Kozar, EF and J. Walter. 1985. Check-list of the Pa- laearctic Coccoidea (Homoptera). Folia Entomo- logica Hungarica 46: 63-110. Kuhlgatz. 1898. Schildléuse auf Kakteen nebst Be- schreibung von Rhizococcus multispinosus nov. spec. Monatsschrift fiir Kakteenkunde 8: 166— 170; 185-188. Lambdin, P. L. 1983. A revision of the genus Astero- PG coccus Borchsenius (Homoptera: Cerococcidae). Proceedings of the Entomological Washington 85: 297-308. Maskell, W. M. 1892. Further coccid notes: with de- scriptions of new species, and remarks on coccids from New Zealand, Australia and elsewhere. Transactions and Proceedings of the New Zealand Institute 24: 1—64. Matesova, G. I. 1960. New species of soft scales fam. Pseudococcidae (Homoptera, Coccoidea) of the Kazakhstan fauna. Trudy Instituta Zoologii, Aka- demii Nauk Kazakhskoy SSR, Alma-Ata 11: 205— DAE 1976. Two new species of eriococcids (“‘felt- coccoid”’) (Homoptera, Coccoidea, Eriococcidae) in Kazakhstan. Izvestiya Akademii Nauk Ka- zakhskoi SSR, (Seriya) Biologii 3: 22-26. McKenzie, H. L. 1942. New species of pine-infesting Margarodidae from California and southwestern United States (Homoptera; Coccoidea; Margarod- idae). (Contribution No. 30). Microentomology 7: 1-18. Miller, D. R. 1991. Systematic analysis of Acantho- coccus species (Homoptera: Coccoidea: Eriococ- cidae) infesting Atriplex in western North Amer- ica. Proceedings of the Entomological Society of Washington 93: 333-355. Miller, D. R. and M. E. Gimpel. 1996. Nomenclatural changes in the Eriococcidae (Homoptera: Coccoi- dea). Proceedings of the Entomological Society of Washington 98: 597-606. Miller, D. R., T. Liu, and J. O. Howell. 1992. A new species of Acanthococcus (Homoptera; Coccoi- dea; Eriococcidae) from sundew (Drosera) with a key to the instars of Acanthococcus. Proceedings of the Entomological Society of Washington 94(4): 512-523. Miller, D. R. and G. L. Miller. 1992. Systematic anal- ysis of Acanthococcus (Homoptera: Coccoidea: Eriococcidae) in the western United States. Trans- actions of the American Entomological Society 118 (1): 1-106. 1993. Eriococcidae of the Eastern United States (Homoptera). Contributions of the Ameri- can Entomological Institute 27(4): 1-91. Miller, D. R. and D. J. Williams. 1976. Proposed con- servation of the family-group name Eriococcidae Cockerell, 1899 (Insecta, Homoptera) and the des- ignation of a type-species for Eriococcus Targioni- Tozzetti, 1868 under the plenary powers Z.N.(S.) 2140. Bulletin of Zoological Nomenclature 33: 118-123. Morrison, H. and Morrison, E. R. 1966. An annotated list of generic names of the scale insects (Homop- tera: Coccoidea). Miscellaneous Publication Unit- ed States Department of Agriculture 1015: 1—206. Nast, J., E. Chudzicka, S. M. Klimaszewski, W. Wo}- ciechowski, Czylok, and J. Kotjea. 1990. [Check- Society of list of Animals of Poland.] Wykaz zwierzat Polski, Razowski, J., Ed. Zaklad Narodowy im. Ossolin- skich Wydawnictwo Polskie} Akademii Nauk 158 PPp- Ramachandran, S. and T. V. Ramakrishna Ayyar 1934. Host plant index of Indo-Ceylonese Coccidae. Miscellaneous Bulletin, India Imperial Council of Agricultural Research 4: 1-113. Ramakrishna Ayyar, T. V. 1936. Notes on Coccidae (Homoptera, Rhynchota) from south India. Jour- nal of the Bombay Natural History Society 39: 146-148. Savescu, A. D. 1985. Especes de coccoidées nouvelles pour la science signalées en Roumanie. III. Es- peces appartenant aux genres Pseudococcus Westw., Phenacoccus Ckll., Paroudablis Ckll., Eupeliococcus Savescu et Lepidosaphes Shimer (Homoptera-Coccoidea). Bulletin de 1’ Académie des Sciences Agricoles et Foresti¢res. Bucarest 14: 103-130. Tang, FE T. 1984. Observation on the scale insects in- jurious to forestry of North China. Shanxi Agri- cultural University Press Research Publication 2: 122-133. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Tang, F T. and J. Hao. 1995. The Margarodidae and Others of China. Chinese Agricultural Science Technology Press Beijing, P. R. China, 738 pp. Tang, FE T. and J. Li. 1988. Observations on the Coc- coidea of Inner Mongolia in China. Inner Mon- golia University Press, 227 pp. Tereznikova, E. M. 1965. New species of the genus Greenisca Borchs. (Coccoidea, Eriococcidae). Dopovidi Akademii Nauk Ukrainskoi RSR 7: 957-959. Tereznikova, E. M. 1977. Two new species of Erio- coccidae from the genus of Acanthococcus Sign. (Homoptera, Coccoidea, Eriococcidae). Akademii Nauk Ukrains’koi RSR Dopovidi Seriya B 6: 568-571. . 1981. Scale insects: Eriococcidae, Kermesi- dae and Coccidae. Fauna Ukraini. Akademiya Nauk Ukrainskoi RSR. Institut Zoologii. Kiev 20: 1-215. Williams, D. J. 1985. The British and some other Eu- ropean Eriococcidae (Homoptera: Coccoidea). Bulletin of the British Museum (Natural History) Entomology Series 51: 347-393. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 219-220 NOTE Scaphytopius angustatus (Osborn) (Homoptera: Cicadellidae), a Leafhopper Characteristic of Pitch Pine-Scrub Oak Barrens Scaphytopius angustatus (Osborn), a widely distributed Nearctic leafhopper of the deltocephaline tribe Scaphytopiini, is the only known conifer-feeding member of the genus (Hepner. 1947. University of Kansas Science Bulletin 31: 413-541). Adults are about 4 to 5 mm long, pale greenish fulvous or greenish yellow, with subhyaline forewings; the distinctive male genitalia should be examined for positive identification (Hepner 1947; Berine. 1952. Canadian Entomologist 84: 311—313; 1956. Canadian Entomologist 88(Supplement 2): 1-180). This pine specialist has been re- corded from jack pine (Pinus banksiana Lamb.), red pine (P. resinosa Aiton), and pitch pine (P. rigida Mill.) (Ball. 1932. Ca- nadian Entomologist 64: 251—255; Hepner 1947). Ecological information otherwise is lacking for this infrequently collected spe- cies. During studies of mirids (Wheeler. 1991. Journal of the New York Entomological So- ciety 99: 405—440) and fulgoroids (Wheeler and Wilson. 1996. Proceedings of the En- tomological Society of Washington 98: 100-108) inhabiting pitch pine-scrub oak barrens, I found S. angustatus to be a char- acteristic insect of northeastern pine bar- rens—that is, occurring consistently in, but not restricted to, that community type. It is one of several leafhopper species found on pitch pine in pine barrens. Individuals of the leafhopper were beaten from branches of pines, mainly pitch pine, as described by Wheeler (1991) for mirids occurring on scrub oak (Quercus ilicifolia Wangenh.). Voucher specimens have been deposited in the collections of Cornell University, Itha- ca, N.Y.; National Museum of Natural His- tory, Washington, D.C.; and the Pennsyl- vania Department of Agriculture, Harris- burg. Once considered a “‘distinctly northern species” (DeLong. 1923. pp. 56—163 In Britton, W.E., ed., The Hemiptera or Suck- ing Insects of Connecticut. Connecticut Geological and Natural History Survey Bulletin 34), S. angustatus is now known as far south as Georgia. Other records in- clude Maine, Massachusetts, Minnesota, Missouri, New Hampshire, New Jersey, New York, North Carolina, Ohio, Ontario, Pennsylvania, South Carolina, Virginia, and Wisconsin (Metcalf. 1967. pp. 2,075—2,695 In General Catalogue of the Homoptera, Fascicle VI, Part 10, USDA ARS, Wash- ington, D.C.). Metcalf (1967) also listed Connecticut, but in a treatment of that state’s leafhopper fauna, S. angustatus was only predicted to be found there (DeLong 1923): During 1991-1995, I collected S. angus- tatus at 21 sites, ranging from extensive pitch pine-scrub oak barrens, such as Wa- terboro in Maine, Ossipee in New Hamp- shire, New York’s Shawangunk Mountains, and the New Jersey Pine Barrens, to de- graded, remnant pine barrens. Seven collec- tions involved scattered pitch pines in com- munities other than pine barrens. Collec- tions were made from pitch pine except in the Gadway Barrens, Clinton Co., N.Y., where jack pine was the host. New state records are Connecticut, Rhode Island, and Vermont. Numbers in parentheses refer to adults unless otherwise stated. CONNECTICUT: Hartford Co., Shaker Pines, Enfield, 28 Sept. 1991 (3). MAINE: York Co., Biddeford, 14 Aug. 1993 (2); Kennebunk Plains, 13 Aug. 1993 (1); Wa- terboro Barrens Preserve, 8 Aug. 1995 (1, 15 nymphs). MASSACHUSETTS: Franklin Co., Montague sand plains, 14 Sept. 1991 Ch) 1s Aus. 19935(37 1 nymph). NEW. 220 HAMPSHIRE: Carroll Co., Ossipee Pine Barrens, 7 Aug. 1995 (6 nymphs); Hills- borough Co., Amherst (1) & Nashua (1), 6 Aug. 1995; Brookline, 14 Sept. 1991 (1). NEW JERSEY: Burlington Co., NW. of Warren Grove, 11 Aug. 1991 (2). NEW YORK: Albany Co., remnant barrens near Pine, Bush \Preserve, 22. Aug; 1993, (1); Clinton Co., NE. of Ausable Chasm, 30 Aug. 1992 (1); Gadway Barrens, S. of Can- non Corners, 21 Aug. 1993 (1); West Chazy Barrens, 29 Aug. 1992 (2); Jefferson Co., Plessis, 16 Aug. 1992 (>20 adults and nymphs); Saratoga Co., 3 mi. S. of South Glens Falls, 22 Aug. 1992 (1); Ulster Co., Mohonk Perserve near New Paltz, 29 Sept. 199i (2). 21 Aug. 1992-(3):.20 Aug, 1995 (1). PENNSYLVANIA: Luzerne Co., Hum- boldt Industrial Park SW. of Hazleton, 22 Aug. 1993 (1); Schuylkill Co., near Frack- ville, 6 Oct. 1991 (1). RHODE ISLAND: Providence Co., Slatersville, 19 Sept. 1992 (2); Washington Co., near Arcadia Manage- ment Area, 1 Sept. 1991 @). VERMONT: Chittenden Co., Camp Johnson, Colchester, 28 Aug. 1992 (1). Nymphs were seldom observed, and only the mostly bright green fifth instars were detected, earlier instars perhaps having been overlooked in the beating net. Because nymphs were not observed before August and no adults were taken during extensive spring and early-summer sampling of pitch pine, S. angustatus likely overwinters in the egg stage. Adults of this late-season, apparently univoltine leafhopper were observed in pine barrens from early August until early Oc- tober. In the northern part of its range (OH, NH, NY, WI), S. angustatus has been re- ported only from early August (Sanders and DeLong. 1917. Annals of the Entomologi- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cal Society of America 10: 79-95) to mid- October (Osborn and Knull. 1947. Ohio Journal of Science 46: 329-336), even in New Hampshire, where Lowry (1933. Ohio Journal of Science 33: 59-80) collected leafhoppers nearly throughout the season, beginning in May. In the southern part of the range, adults have been collected in Georgia from June to August (Fattig. 1955. Emory University Museum Bulletin 11: 1— 68). In northeastern pine barrens, I observed S. angustatus mainly on seedling and sap- ling pitch pines and the regrowth from stumps of cut-over trees. Certain other co- nifer-feeding leafhoppers are associated with seedlings, including those of pitch pine (DeLong, 1926. Ohio Journal of Science 26: 69-72). About a third of the collections of S. angustatus were from mature pitch pines, where they often were beaten from bushy growth on the basal whorl of branch- es, some of which touched the ground. This growth habit, which might be a response to light reflected from the sand, typifies pitch pine in pine barrens (Kelley. 1927. Botan- ical Gazette 83: 89-93). I gratefully acknowledge those who ac- companied me in the field or enabled me to find pine barrens: Kenneth Adams, Robert Dirig, Paul Huth, Patrick McCarthy, Alan Nye, Dale Schweitzer, Nancy Sferra, and David VanLuven. Christopher Dietrich kindly identified S. angustatus, The Nature Conservancy and State Heritage Programs allowed access to several pine barrens, and Peter Adler provided useful comments on an early draft of the manuscript. A. G. Wheeler, Jr., Department of Ento- mology, Clemson University, Clemson, SC 29634, U.S.A. (e-mail: awhlr@clemson.edu). PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 221-222 BOOK REVIEW Os Mosquitos de Macau (Diptera: Cu- licidae). Helena Cunha Ramos, Henrique Ribeiro, Maria Teresa Novo, and Emmett R. Easton. 1997. Sociedade Portuguesa de Entomologia, Apartado 8221, P-1800 Lisbon. 201 pp., paper. ISBN 972-97241- 0-5. Among the perquisites of my present po- sition is total access to the world literature on medical entomology. However, despite my databases, I am still occasionally sur- prised, as by the recent receipt of this lavish monograph on the mosquitoes of Macau (English Macao, Mandarin Aomen), Portu- gal’s ancient entrepot on the South China Sea. At the eleventh hour, with Chinese of- ficials poised to take Government House, four widely experienced field biologists from the Centro de Zoologia, Instituto de Investigagao Cientifica Tropical (Cunha Ra- mos), the Disciplina de Entomologia Méd- ica, Instituto de Higiene e Medicina Tropi- cal, Universidade Nova de Lisboa (Ribeiro and Teresa Novo), and the Centro de Es- tudos Pré-Universitarios, Universidade de Macau (Easton, an expatriate American profiled in J. New York Entomol. Soc. 102: 389-391) have joined to forge a faunal sur- vey of continental proportions. Throughout 1994 and 1995, Cunha Ra- mos and colleagues scoured peninsular Ma- cau and its offshore islands of Taipa and Coloane, to which the cidade is linked by bridges and causeways. In an utterly an- thropocentric environment, they secured some 3,000 mosquito specimens represent- ing 28 species (generic and subgeneric ab- breviations follow Reinert, Mosq. Syst. 7: 105-110, 14: 124-126, 23: 209-210): Aedes (Finlaya) togoi (Theobald 1907) Ae. (Stegomyia) albopictus (Skuse 1894) Ae. (Stg.) w-albus (Theobald 1905) Anopheles (Anopheles) sinensis Wiede- mann 1828 Armigeres (Armigeres) subalbatus (Coquil- lett 1898) Ar. (Leicesteria) magnus (Theobald 1908) Culex (Culex) bitaeniorhynchus Giles 1901 Cx. (Cux.) jacksoni Edwards 1934 Cx. (Cux.) pseudovishnui Colless 1957 Cx. (Cux.) quinquefasciatus Say 1823 Cx. (Cux.) sitiens Wiedemann 1828 Cx. (Cux.) tritaeniorhynchus Giles 1901 Cx. (Cux.) vagans Wiedemann 1828 Cx. (Cux.) vishnui Theobald 1901 Cx. (Culiciomyia) pallidothorax Theo- bald 1905 Cx. (Eumelanomyia) foliatus Brug 1932 Cx. (Eum.) malayi (Leicester 1908) Cx. (Lophoceraomyia) infantulus Ed- wards 1922 Cx. (Lop.) rubithoracis (Leicester 1908) Cx. (Lop.) sumatranus Brug 1931 Cx. (Lutzia) fuscanus Wiedemann 1820 Cx. (Lut.) halifaxii Theobald 1903 Mansonia (Mansonioides) uniformis (Theo- bald 1901) Mimomyia (Mimomyia) chamberlaini Lud- low 1904 Toxorhynchites (Toxorhynchites) macaensis Ribeiro 1997 Tripteroides (Rachionotomyia) aranoides (Theobald 1901) Uranotaenia (Uranotaenia) annandalei Barraud 1926 Ur. (Ura.) macfarlanei Edwards 1914 Perusal of the regional literature (their bib- liography, pp. 195-201, contains 208 en- tries) yielded seven additional species: Ae. (Stg.) aegypti (Linnaeus 1762), An. (Cellia) jeyporiensis James 1902, An. (Cel.) karwari (James 1902), An. (Cel.) maculatus Theo- bald 1901, An. (Cel.) minimus Theobald 1901, An. (Cel.) tessellatus Theobald 1901, and Cx. (Cux.) fuscocephala Theobald 1907, all thought to occur in Macau in ‘‘very low densities.”’ For a territorial total of 35 species, including the just-described T. macaensis (J. Am. Mosq. Control Assoc. 13: 213-217), keys are provided to males, females, and 4th instar larvae. The keys are exceptionally well supported with pen-and- ink drawings, paintings, and stunning color photomicrographs (of 179 numbered illus- trations, all but 46 are in full color). The core of the text (pp. 67-193) is a species-by-species account of mosquito tax- onomy, ecology, and distribution. Remark- able in this regard are three full-page, color- coded maps (pp. 68, 84 and 91) that pin- point collecting sites for each species on Taipa, Coloane, and Macau proper. Thus, it is possible to retrace the authors’ steps to particular intersections, ponds, or cramped urban parks, where subsequent generations of the very insects that were the subjects of this study may yet be sought. As guidance to future investigators, photographs of 32 of these sites are included. The vector poten- tial of each species is also assessed for the malarias, 14 arboviruses, human filariases (Brugia malayi, Wuchereria bancrofti) and dirofilariases (Dirofilaria immitis, D._ re- pens). A major goal of this endeavor was to compare the diversity of the Macanese mosquito fauna with that of neighboring Hong Kong and the Oriental Region as a whole. To this end, indices of abundance, association and distribution were calculated and are presented as tables throughout the BOOK The Everglades Handbook: Under- standing the Ecosystem. Thomas E. Lodge, introduction by Marjory Stone- man Douglas. 1994, second printing, 1998:..St."LEucie: Press.\ xix--228"-pp., ‘pa= per. ISBN 1-884015-06-9. $39.95. The one person most closely associated PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON text. These include the distribution of mos- quito species in Macau by principal juris- diction (Table 1); relative breeding indices for each species (Table 2); percentages for utilization of “‘natural’’ versus manmade habitats (Table 3); indices of larval associ- ation and their statistical significance (Ta- bles 4 and 5); the extraterritorial distribu- tion of each species by zoogeographic re- gion (Table 8); and comparative indices of biodiversity for the mosquito faunas of Ma- cau, Hong Kong, Cameroon, and mainland Portugal (Table 9). Such distillations be- speak countless hours of fieldwork—and countless more of analysis. The Portuguese are passing now. On 20 December 1999, they will retrocede to Chi- na the oldest Western enclave in the Orient. How auspicious, for these two peoples have no quarrel with one another. The Lusitani- ans will be remembered not for foisting nar- cotics on a population held hostage, not for having to be forcibly evicted a la Dien Bien Phu, and certainly not for pushing the latest ludicrous cult from the so-called American heartland. Rather, both sides will be left to reflect on over 400 years of mutualism, as evidenced by this oblation. Richard G. Robbins, Armed Forces Pest Management Board, Walter Reed Army Medical Center, Washington, DC 20307- SOOT, U.S.A. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, pp. 222-224 REVIEW with the formation of Everglades National Park was Marjory Stoneman Douglas. Her death earlier this year at 108 years of age marked the end of an important chapter in the history of south Florida. This book pro- vides an introduction to the Everglades, covering all aspects of the region. Mrs. Douglas’ short introduction to the book VOLUME 101, NUMBER 1 serves as a brief memorial to her efforts to establish the Park. The Everglades Hand- book: Understanding the Ecosystem is writ- ten from an environmental protectionist perspective, which is to be expected of any work to which Mrs. Douglas contributed. The book is divided into four parts, each part being divided further into a number of chapters. The first part, Background, con- tains two chapters. Chapter 1, The Ever- glades in Space and Time, describes the geologic history of the Everglades. Chapter 2, An Ecosystem Review, is an important chapter for the reader, for it is here that the author defines many terms referring to wa- ter and hydrology, including what the Ev- erglades are and are not, and more interest- ingly, where they are and where they are not. There is a great difference between what the Everglades historically were and what we now think of them as being. The second part of the book, Environ- ments of the Everglades Region, consists of seven chapters, each detailing one aspect of the Everglades. These chapters are weight- ed toward discussion of plant communities. Chapter 3 describes freshwater marshes, in- cluding the plants, soil, water, and effects of weather and fire on the marshes. Chapter 4 discusses tree islands and their impor- tance to the region. Chapter 5 treats hard- wood hammocks and Chapter 6 pinelands. Both chapters discuss the role of fire in maintenance of the environment. Chapter 7 is a very interesting discussion of mangrove swamps and their importance to wildlife and fisheries. Chapter 8 describes the veg- etation of the coastal lowlands and the ef- fects of hurricanes on south Florida. Final- ly, Chapter 9 deals with the estuaries and marine waters of the coast. This chapter dis- cusses the flora and geology of Florida Bay and the Gulf of Mexico, and the relation- ship between oysters and mangroves. Biogeography of Southern Florida, the third part of the book, probably will be of most interest to entomologists. There are eight chapters in this part of the book, seven of them covering a specific group of ani- 223 mals. Chapter 10 includes such topics as the origin of the biota of southern Florida, and whether south Florida is tropical or sub- tropical. The Everglades region contains both temperate and tropical plant species, and the origins of each component of the flora are reviewed. Chapter 11 treats the in- vertebrates, marine, freshwater, and terres- trial. My bias as an entomologist led me to wonder why so few insect species were mentioned, but in fairness there is only one chapter to deal with all invertebrate ani- mals. Among the organisms mentioned are butterflies, crayfish, lobster, shrimp, snails, and spiders. Chapters 12 and 13 cover the fishes. Chapter 12 deals with freshwater species and Chapter 13 with estuarine and marine species. In Chapter 12 the distinc- tion among primary, secondary, and periph- eral freshwater fishes is explained. The im- portance of these fishes to the food chain is mentioned. Chapter 13 summarizes the di- versity of the marine and estuarine fishes, and their importance both as game fishes and as a food source for birds. Chapter 14 is a very brief (three and one-half pages) mention of the amphibians of the Ever- glades. One interesting facet of this chapter is the impact that introduced frogs and toads have had on the native fauna. Chapter 15 considers the reptiles. Most of the chap- ter deals with crocodilians, although lizards, snakes, tortoises, and turtles are mentioned. Chapter 16 is another very brief chapter that deals with mammals. Almost one entire page is devoted to the Florida panther. Oth- er species receive little attention. The last chapter in this part of the book, Chapter 17, discusses the birds. This chapter devotes much of its space to the large wading birds, which the author admits ‘‘attract much at- tention,’ and so they do here as well. This is one of the longer chapters in the book, comprising 18 pages. There is only brief mention of passerine birds, raptors, and oth- er types of birds. The fourth part of the book, Environ- mental Impacts, contains only one chapter, Chapter 18, that describes the effects that 224 south Florida’s increasing population has had on the Everglades. Specimen collect- ing, off-road vehicles, and introduced spe- cies are all mentioned as having adverse impacts on the Everglades. A great portion of the chapter is devoted to the effect that water control projects have had on the Ev- erglades. There is an interesting attempt to bring the global warming controversy into the discussion. However, the author does point out that there is another opinion with- in the scientific community, that of global cooling. The book ends with speculation on the demise of the Everglades. The book is extensively footnoted, and contains almost 300 references. The index is superb, permitting the reader to locate passages pertinent to any animal or plant by common or scientific name. The illustra- tions are of good quality. One disappoint- ment is that all photos are in black-and- white. The author writes in his preface that it was his intention to include a large num- ber of color photographs, but this became impossible due to realities of the publishing business. Many of the photos in the book were taken by photographer Robert Hamer, and their quality and composition leaves PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON one hoping that the color version will some day see print. The shorter chapters are a bit frustrating to read, because the reader is given only enough information to whet the appetite. The shortness of some of the chap- ters apparently is due to their having been intended originally to accompany the color photos that never were used. The book was not conceived as a textbook, and it is evi- dent that the author did not intend it to be used as one. Readers who want a mathe- matical treatise detailing the population dy- namics of all species in the Everglades will not find that here. For those individuals who have an organismal interest in the Ev- erglades, e.g., naturalists and professional biologists in south Florida, the book will serve aS a convenient introduction to some of the more spectacular animals and plants found in the region. For others, the book makes for pleasant reading, but its coverage of the invertebrate fauna is too limited to be of much use to the professional ento- mologist. Lawrence J. Hribar, Monroe County Mosquito Control District, Marathon, FL 33050) UESZA. PROC. ENTOMOL. SOC. WASH. 101(1), 1999, p. 224 1999 MEETINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The following programs are scheduled for February to May, 1999. Meetings are at 7:30 PM, the first Thursday of each month, in the National Museum of Natural History, Smithsonian Institution, Washington, DC. February 4, 1999—Alan Schroeder (U.S. AID), ‘“‘U.S. AID’s Pest Management Activities in Africa.” March 4, 1999—Dale FE Schweitzer (The Nature Conservancy, Port Norris, NJ), ‘Lepidoptera and Other Insects in the New Jersey Pine Barrens.” April 1, 1999—Ted R. Schultz (Smithson- ian Institution), ‘“‘The Natural History of Fungus-Growing Ants.” May 6, 1999—Joseph V. McHugh (Univer- sity of Georgia), ““A Phylogenetic Analysis of Erotylidae (Coleoptera) with Implications for the Evolution of Their Mycophagy.” 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 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. No. 1. No. 4. No. 5. No. 6. No. 12. No. 22. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi Takahasi. 230 pp. 1969 The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato- pogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. 67 pp. 1984 . Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 pp. 1995 . A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent and W. L. Grogan, Jr. 198 pp. 1995 . The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 _ . Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited by B. B. Norden and A. S. Menke. 216 pp. 1996 . Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and William L. Grogan, Jr. 297 pp. 1997 . Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by Thomas J. Henry. 149 pp. 1997 . The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- gafelter. 118 pp. 1998 . New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. Furth. 110 pp. 1998 Systematics of the North American Species of Trichogramma Westwood (Hymenoptera: Trichogrammatidae), by John D. Pinto. 287 pp. 1999 $10.00 15.00 $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 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 except on prepaid orders. Dealers are allowed a discount of 10 per cent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Custodian, Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. CONTENTS (Continued from front cover) KETH, A. C. and S. C. HARRIS—Two new species of Agarodes Banks (Trichoptera: Sericos- tomatidae):from\southeastern:. United’ Statesnetaacase cae os senate teenie otine eaten ere eee eee KROMBEIN, KARL V. and ARKADY S. LELEJ—Biosystematic studies of Ceylonese wasps, XXII: Bethsmyrmilla, a new genus of mutillid wasps (Hymenoptera: Mutillidae: Myrmillinae) ........ LUGO-ORTIZ, C. R., W. P. McCAFFERTY, and J.-L. GATTOLLIAT—The small minnow mayfly genus Cloeodes Traver (Ephemeroptera: Baetidae) in Madagascar .................... MILLER, DOUGLASS R. and MAREN E. GIMPEL—New combinations, new synonymy, and hymonomy in the Eriococcidae, new homonomy and synonymy in the Cerococcidae, and transfer of Cancerococcus Koteja to the Margarodidae (Hemiptera: Coccoidea) ............. SCHAEFER, CARL W.—The higher classification of the Alydidae (Hemiptera: Heteroptera) ....... SITES, ROBERT W. and BECKY J. NICHOLS—Egg architecture of Naucoridae (Heteroptera): Internal’ and’ external-structure ‘of ‘the \chorionvand mictopylejy a. 23.-9...1.2 epee ese ee tees SLATER, JAMES A. and ALEX SLATER—Ashlockobius, a new genus of Myodochini from Venezuela (Hemiptera: Lygaeoidea: Rhyparochromidae: Myodochini) ........................ TROSTLE, M., N. S. CARREJO, I. MERCADO and R. A. WHARTON—Two new species of Phaenocarpa Foerster (Hymenoptera: Braconidae: Alysiinae) from South America ......... NOTE WHEELER, A. G., JR.—Scaphytopius angustatus (Osborn) (Homoptera: Cicadellidae), a leaf- hopper, characteristic; of pitch) pine=scrub) oak barrens: <22:2. 2.2. sens see ee o- + ead salar BOOK REVIEWS ROBBINS, RICHARD G.—Os Mosquitos de Macau (Diptera: Culicidae) by H. C. Ramos, H. Ribeirov Me sE Novo: andgsEodR Easton us: Foe aie Nels See Serene Iie ac nee HRIBAR, LAWRENCE J.—The Everglades Handbook: Understanding the Ecosystem, by Thomas EL OUR OS ih SEs scat Saas eeRTeh aatviatane opstobt ele Oideatar attests eee te reo ere ese RPI URS area or aor MISCELLANEOUS LOOOUMICEUIN DS isc FO Se cee ee Arya cae rag he Sine Sine ala era cee nla ESS Rep Ae cPe LS ORE Fo ake cenafan ne at 86 143 208 197 219 221 VOL. 101 APRIL 1999 NO. 2 LAL (ISSN 0013-8797) aaa) =’ PROCEEDINGS Hn of the PUBLISHED QUARTERLY CONTENTS ADAMSKI, DAVID—Two Neotropical Hypatopa Walsingham (Gelechioidea: Coleophoridae: Blastobasinae) with retractile labial palpi: A previously unknown lepidopteran feature ..... 438 BATRA, SUZANNE W. T.—Native bees (Hymenoptera: Apoidea) in native trees: Nyssa syl- VRUARETT ANETES OU COVOT GALES TEN [ake a8 WA Sa AW CIA Ge Eee as ts Per mes gr Shae Coit WG UMMA SUN a a te 449 CONTRERAS-RAMOS, ATILANO—List of species of Neotropical Megaloptera (Neuropterida) .. 274 DIETRICH, CHRISTOPHER H., MARK J. ROTHSCHILD, and LEWIS L. DEITZ—Checklist and host plants of the treehoppers (Hemiptera: Membracidae) of North Carolina ............ 242 FREIDBERG, AMNON—A new species of Craspedoxantha Bezzi from Tanzania and a revised PIVIGSeny Once. cenusn(Diptera:, bephritidae) aren ete cos lent cei fee e mate kiejeia shelalactald wad an ils 382 GAGNE, RAYMOND J. and JOHN T. LILL—A new Nearctic species of Lestodiplosis (Diptera: Cecidomyiidae) preying on an oak leaf tier, Psilocorsis quercicella (Lepidoptera: Oeco- jeyYOUCEGEEYE))) Has OSTA Vi Sey Aik ara VEU Ie UG Se Ue ce a a CL a Ae gl Li BaD. GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Life history and description of imma- ture stages of Trupanea wheeleri Curran (Diptera: Tephritidae) on Asteraceae in southern CI RPSYG CUE ye ae is eT a Fea Sh ED PET aa a oe At a eC fe a, 414 HATHORNE, KEVIN T. and PATRICIA A. ZUNGOLI—Ildentification of late-instar nymphs OiMCOGROACHES) CE LattOG Gayie a eau 14 sear ead Se aaah dee Lignan Dae EWN UL TR SU ie Si Neeriei dete Rios oie 316 KIMSEY, LYNN S.—A turn of the century conundrum—reexamination of Aeolothynnus Ash- Meada(HvinenOplerasg Piplitd aes MUMMY INIMITLAG )h \eysnisaleeatie < Saye lee)~ eistoss)-/eie b:njeiefe clclelssaisists(olajaletre eheisy ola catalsyale 263 KONDRATIEFF, BORIS C. and RICHARD W. BAUMANN-—Studies on stoneflies of North Dakota with the description of a new Perlesta species (Plecoptera: Perlidae) .......-........ 325 KORCH, PETER P., STEVEN L. KEFFER, and ELISA WINTER—Description of immature stages of Platyvelia brachialis (Stal) (Heteroptera: Veliidae) .................. eee cece ee cee eens 372 LI, CHUN-LIN and PING-SHIH YANG—A new genus Clypeolontha Li and Yang, for the genus Melolontha Fabricius (Coleoptera: Scarabaeoidea: Melolonthinae) from southeastern (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1999 MICHAEL E. SCHAUFF, President MICHAEL G. POGUE, Treasurer DAVID G. FuRTH, President-Elect JOHN W. Brown, Program Chair STUART H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair Houuis B. WILLIAMS, Corresponding Secretary WARREN E. STEINER, Past President ANDREW S. JENSEN, Custodian DAVID R. SmiTH, Editor Publications Committee THOMAS J. HENRY WAYNE N. MATHIS Gary L. MILLER, Book Review Editor Honorary President LouIsE M. RUSSELL Honorary Members KARL V. KROMBEIN RONALD W. HODGES 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 7:30 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. 599-600 OF THE JULY 1998 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: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC- West, Beltsville, MD 20705. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 30 April 1999 Second Class 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. 101(2), 1999, pp. 225-232 DESCRIPTION OF IMMATURE STAGES OF TRUPANEA IMPERFECTA (COQUILLETT) (DIPTERA: TEPHRITIDAE) JEFFREY A. TEERINK AND RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: rgoeden @ucrac 1 .ucr.edu) Abstract.—The egg, first-, second- and third-instar larvae, and puparium of Trupanea imperfecta (Coquillett), a monophagous, bi- or trivoltine tephritid principally reproducing in flower heads of Bebbia juncea (Bentham) Greene (Asteraceae) in southern California, are described and figured for the first time. The egg pedicel is composed mainly of a single row of large aeropyles. As with the other Trupanea species previously studied, the lateral spiracular complex of the third instar is unique to 7. imperfecta, with a stelex sensillum and two verruciform sensilla on the metathorax, and two verruciform sensilla on the abdominal segments. The third instar of 7. imperfecta very closely resembles T. arizonensis Malloch in general habitus and sensory structures. Key Words: Insecta, Trupanea, Asteraceae, nonfrugivorous Tephritidae, taxonomy of immature stages, egg, larva, puparium The life history of Trupanea imperfecta (Coquillett) (Diptera: Tephritidae) was de- scribed by Goeden (1988) before adoption of our current format incorporating descrip- tion of the immature stages. To correct this deficiency and allow full comparison with the 36 species of southern California non- frugivorous fruit flies for which life histo- ries and descriptions of the immature stages have now been published, this paper de- scribes the immature stages of 7. imperfec- td. MATERIALS AND METHODS One-liter samples of excised, immature and mature flower heads from the main host of T. imperfecta, Bebbia juncea (Bentham) Greene (Asteraceae), potentially containing eggs, larvae, and puparia were transported in cold-chests in an air-conditioned vehicle to the laboratory and stored under refrig- eration for subsequent dissection, photog- raphy, description, and measurement. Twenty-two eggs, 23 first-, 14 second-, and nine third-instar larvae, and nine puparia dissected from flower heads were preserved in 70% EtOH for scanning electron micros- copy (SEM). Specimens for SEM were hy- drated to distilled water in a decreasing se- ries of acidulated EtOH. They were osmi- cated for 24 h, dehydrated through an in- creasing series of acidulated EtOH and two, l1-h immersions in Hexamethlydisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, and studied with a Philips XL30-FEG SEM in the In- stitute of Geophysics and Planetary Phys- ics, University of California, Riverside. Plant names used in this paper follow Munz (1974); tephritid names follow Foote et al. (1993). Terminology and telegraphic format used to describe the immature stages follow Knio et al. (1996), Goeden and Teer- inke 5(L997-4 1998;. 11999) .Goedeniyet val: (1998a, b), and Teerink and Goeden (1998), and our earlier works cited therein. Means 226 + SE are used throughout this paper. Voucher specimens of 7. imperfecta eggs, larvae and puparia are stored in a collection of immature Tephritidae acquired by JAT and now maintained by RDG. RESULTS AND DISCUSSION Taxonomy Immature stages.—The egg and pupari- um of JT. imperfecta were described and photographs of these stages and larvae were provided by Goeden (1988), but detailed descriptions based on scanning electron mi- croscopy heretofore have not been pub- lished. Egg: The egg (Fig. 1A) of 7. imperfecta has a short pedicel circumscribed by a sin- gle row of subrectangular aeropyles so large that they uniquely occupy more than half of this structure (Fig. 1B). The micro- pyle is located on the anterior end of the pedicel (Fig. 1C-1). First instar: White, elongate-cylindrical, rounded anteriorly and posteriorly, minute acanthae circumscribe intersegmental lines (Fig. 2A); gnathocephalon smooth, lacking rugose pads (Fig. 2C); dorsal sensory organ a dome-shaped papilla (Fig. 2B-1); subdor- sal sensillum located laterad of dorsal sen- sory organ (Fig. 2B-2); anterior sensory lobe bears terminal sensory organ (Fig. 2B- 3), lateral sensory organ (Fig. 2B-4) and su- pralateral sensory organ (Fig. 2B-5); stomal sense organ ventrad of anterior sensory lobe (Fig. 2C-1); mouth hooks bidentate (Fig. 2C-2); median oral lobe laterally flat- tened (Fig. 2C-3); a pair of integumental petals dorsad of mouth hooks (Fig. 2C-4); pit sensillum laterad of mouth lumen (Fig. 2C-5); minute acanthae ventrad of mouth lumen (Fig. 2C-6); anterior spiracle absent; abdominal lateral spiracular complex con- sists of a spiracle and two verruciform sen- silla; caudal segment with two stelex sen- silla dorsad and ventrad of posterior spirac- ular plates (Fig. 2D-1); two verruciform sensilla dorsolaterad of posterior spiracular plates (Fig. 2D-2); posterior spiracular plate PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON AccV SpotMagn WD /—-——————— 299 im 100kV 30 168x 145 wee Om x 146 Spot Magn. = Fig. 1. Egg of Trupanea imperfecta: (A) habitus, pedicel to left; (B) egg pedicel; (C) pedicel, anterior view, |—micropyle, 2—aeropyle. bears two ovoid rimae, ca. 0.008 mm in length (Fig. 2D-3), and four interspiracular processes, each with 1—3 branches, longest measuring 0.010 mm (Fig. 2D-4); inter- mediate sensory complex consists of a ste- lex sensillum (Fig. 2D-5) and a medusoid sensillum (Fig. 2D-6). VOLUME 101, NUMBER 2 Acc V pot Magn 48) 100kV30 198x 145 100 yum ?, és ¥ ‘ oy; AccV “Spot Magn = WD 10.0 kV3.0,3323K_ 14.1% Fig. 2. AccV_ Spot Magn 100kV 30 8281x First instar of Trupanea imperfecta: (A) habitus, anterior end to right; (B) anterior sensory lobe, 1— dorsal sensory organ, 2—subdorsal sensillum, 3—terminal sensory organ, 4—lateral sensory organ, 5—supra- lateral sensory organ; (C) gnathocephalon, anterior view, 1—stomal sense organ, 2—mouth hook, 3—median oral lobe, 4—integumental petal, 5—pit sensillum, 6—minute acanthae; (D) caudal segment, 1—stelex sensillum, 2—-verruciform sensillum, 3—rima, 4—interspiracular process, 5—intermediate sensory complex, stelex sensil- lum, 6—intermediate sensory complex, medusoid sensillum. Second instar: White, elongate-cylindri- cal, tapering anteriorly, rounded posteriorly, minute acanthae circumscribe intersegmen- tal lines (Fig. 3A); gnathocephalon conical; rugose pads laterad of anterior sensory lobe (Fig. 3B-1); dorsal sensory organ a dome- Shaped papilla (Fig. 3B-2, 3C-1); anterior sensory lobe bears terminal sensory organ (Fig. 3C-2), pit sensory organ (Fig. 3C-3), lateral sensory organ (Fig. 3C-4), and su- pralateral sensory organ (Fig. 3C-5); stomal sense organ ventrolaterad of anterior sen- sory lobe (Fig. 3B-3, 3C-6); mouth hooks bidentate (Fig. 3D-1); median oral lobe lat- erally flattened (Fig. 3D-2); labial lobe at- tached to median oral lobe (Fig. 3D-3); six pit sensilla circumscribe gnathocephalon (Fig. 3B-4); minute acanthae circumscribe anterior margin of prothorax (Fig. 3E-1); rugose pads (Fig. 3E-2) and two rows of verruciform sensilla circumscribe prothorax (Fig. 3E-3); anterior thoracic spiracles bear 3 rounded papillae (Fig. 3E-4); lateral spi- racular complex not seen; caudal segment with minute acanthae dorsally (Fig. 3F-1); two stelex sensilla dorsad and ventrad of posterior spiracular plates (Fig. 3F-2); two verruciform sensilla dorsolaterad of poste- rior spiracular plates (Fig. 3F-3); posterior spiracular plate bears three ovoid rimae, ca. 0.021 mm in length (Fig. 3F-4), and four interspiracular processes, each with 1-2 branches, longest measuring 0.013 mm (Fig. 3F-5); intermediate sensory complex 228 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON AccV Spot Magn WD /———— 4 200um 100kV30 112x 143 AccV Spot Magn B10.0kV30 3808K . 15.9 Re ee, ee OY oht Maa Acc Vagpot M3 WO ag. WD * 10.0 KVS0\. 401x —" ‘ ; ic.“ 16.0 Fig. 3. Second instar of Trupanea imperfecta: (A) habitus, anterior to left; (B) gnathocephalon, anterior view, 1—rugose pad, 2—dorsal sensory organ, 3—stomal sense organ, 4—pit sensillum; (C) anterior sensory lobe, 1—dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4—lateral sensory organ, 5— supralateral sensory organ, 6—stomal sense organ; (D) gnathocephalon, ventral view, 1—mouth hooks, 2— median oral lobe, 3—labial lobe; (E) gnathocephalon, prothorax, anterolateral view, 1—minute acanthae, 2 rugose pads, 3—verruciform sensillum, 4—anterior thoracic spiracle; (F) caudal segment, 1—minute acanthae, 2—stelex sensillum, 3—verruciform sensillum, 4—rima, 5—interspiracular process, 6—intermediate sensory complex, medusoid sensillum, 7—intermediate sensory complex, stelex sensillum. VOLUME 101, NUMBER 2 consisting of a medusoid sensillum (Fig. 3F-6) and a stelex sensillum (Fig. 3F-7). Third instar: White, elongate-cylindri- cal, tapering anteriorly, rounded posteriorly, minute acanthae circumscribe intersegmen- tal lines (Fig. 4A); gnathocephalon conical (Fig. 4B), rugose pads laterad of anterior sensory lobe (Fig. 4B-1), those laterad of mouth lumen serrated on ventral margin (Fig. 4C-1); dorsal sensory organ a dome- shaped papilla (Fig. 4B-2, 4C-2); subdorsal sensillum laterad of dorsal sensory organ; anterior sensory lobe (Fig. 4B-3, 4C) bears terminal sensory organ (Fig. 4C-3), pit sen- sory organ (Fig. 4C-4), lateral sensory or- gan (Fig. 4C-5), and supralateral sensory organ (Fig. 4C-6); stomal sense organ ven- trolaterad of anterior sensory lobe (Fig. 4B- 4, 4C-7); mouth hooks tridentate (Fig. 4B- 5); median oral lobe laterally flattened, ta- pering anteriorly (Fig. 4B-6); prothorax cir- cumscribed anteriorly with minute acanthae (Fig. 4D-1); rugose pads circumscribe pro- thorax posteriorad to minute acanthae (Fig. 4D-2); two rows of verruciform sensilla cir- cumscribe prothorax posteriorad to rugose pads (Fig. 4D-3); stelex sensillum located dorsomedially (Fig. 4D-4); anterior thoracic spiracle bears three rounded papillae (Fig. 4D-5); mesothorax and metathorax circum- scribed anteriorly with verruciform sensilla; metathoracic lateral spiracular complex consists of a spiracle (Fig. 4E-1), a stelex sensillum (Fig. 4E-2), and two verruciform sensilla (Fig. 4E-3); abdominal lateral spi- racular complex consists of a spiracle (Fig. 4F-1) and two verruciform sensilla (Fig. 4F-2); caudal segment circumscribed by minute acanthae; two stelex sensilla dorsad and ventrad of posterior spiracular plates (Fig. 4G-1); two verruciform sensilla dor- solaterad of posterior spiracular plates (Fig. 4G-2); posterior spiracular plate bears three ovoid rimae, ca. 0.03 mm in length (Fig. 4G-3), and four interspiracular processes, each with 2—4 branches, longest measuring 0.02 mm (Fig. 4G-4); intermediate sensory complex consists of a medusoid sensillum 229 (Fig. 4H-1), and a stelex sensillum (Fig. 4H-2). Puparium: Black, elongate-cylindrical (Fig. 5A); anterior end bears the invagina- tion scar (Fig. 5B-1), and anterior spiracles (Fig. SB-2); caudal segment circumscribed by minute acanthae (Fig. 5C-1), two stelex sensilla dorsad and ventrad of posterior spi- racular plates (Fig. 5C-2); two verruciform sensilla dorsolaterad of posterior spiracular plates (Fig. 5C-3); posterior spiracular plate bears three ovoid rimae (Fig. 5C-4), and four interspiracular processes, each with 2— 4 branches (Fig. 5C-5); intermediate sen- sory complex consists of a medusoid sen- sillum and a stelex sensillum (Fig. 5C-6). Discussion The egg of Trupanea imperfecta is elon- gate-ellipsoidal, with a reduced peg-like an- terior pedicel (Goeden 1988). It is similar in size and shape to T. signata Foote (Goe- den and Teerink 1997), longer than 7. ac- tinobola (Loew) (Goeden et al. 1998b), 7. californica Malloch (Headrick and Goeden 1991), and T. pseudovicina Hering (Goeden and Teerink 1998), and shorter but wider than T. arizonensis (Goeden and Teerink 1999). The pedicel is mainly composed of large aeropyles (Figure 1B, 1C). The pedi- cels of 7. arizonensis, T. jonesi Curran, T. nigricornis (Coquillett), and 7. pseudovici- na are similar in shape, but have smaller aeropyles spaced farther apart (Goeden and Teerink 1998, 1999; Goeden et al. 1998a; Knio et al. 1996). The first instar of 7. imperfecta is similar in general habitus to previously studied Trupanea species (Goeden and Teerink 1998, 1999; Goeden et al. 1998a, b; Knio et al. 1996; Teerink and Goeden 1998). The gnathocephalon is smooth, lacking rugose pads. The pit sensory organ is indistinct on the anterior sensory lobe, and the stomal sense organ is also indistinct. Minute acan- thae are limited to the ventral margin of the prothorax. The interspiracular processes, each with 1—3 blade-like processes, are sim- ilar to T. actinobola, T. bisetosa (Coquil- 230 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Acc. V Spot Magn WD 10.0kV 3.0558 > 165 . YY Spot Magn LoS kV 3.0 _1025x AccV SpotMagn WD /——4 54m , 1O0kV30 3748x 17.1 Fig. 4. Third instar of Trupanea imperfecta: (A) habitus, anterior to left; (B) gnathocephalon, anterior view, I—rugose pads, 2—dorsal sensory organ, 3—anterior sensory lobe, 4—stomal sense organ, 5—mouth hook, 6—median oral lobe; (C) anterior sensory lobe, |—serrated rugose pad, 2—dorsal sensory organ, 3—terminal sensory organ, 4—pit sensory organ, 5—lateral sensory organ, 6—supralateral sensory organ, 7—stomal sense 2—rugose pad, 3—verruciform sen- organ; (D) gnathocephalon, prothorax, anterior view, 1—minute acanthae, thoracic spiracle; (E) metathorax, 1l—spiracle, 2—stelex sillum, 4—stelex sensillum, 5—anterior VOLUME 101, NUMBER 2 AccV SpotMagn WD es 10.0kV 3.0 36x 149 AX oe: & ‘ 0 RE . a1): NS SS ; : “yas ate Ace V—SpotMagn = WD) = 0:0-kV-3.0~272x Fig. 5. Puparium of Trupanea imperfecta: (A) habitus, anterior end to left; (B) anterior end, 1—in- vagination scar, 2 anterior thoracic spiracle; (C) cau- dal segment, 1—minute acanthae, 2—stelex sensillum, 3—-verruciform sensillum, 4—rima, 5—interspiracular process, 6—intermediate sensory complex. 231 lett), and 7. pseudovicina (Goeden and Teerink 1998; Goeden et al. 1998b; Knio et all 996). The second instar of 7. imperfecta differs from the first instar in possessing rugose pads laterad of the mouth lumen, and the anterior margin of the prothorax is circum- scribed by minute acanthae, rugose pads, and two rows of verruciform sensilla. All four of the anterior sensory lobe sensilla, as well as the stomal sense organ, are distinct in the second instar. The anterior thoracic spiracle is present in the second instar, and the posterior spiracular plates possess three ovoid rimae rather than two as in the first instar. The caudal segment in the second in- star, unlike in the first instar, is circum- scribed by minute acanthae. The third instar differs from the second instar in possessing serrated rugose pads laterad of the mouth lumen, and the mouth hooks tridentate. The third instar of T. im- perfecta is very similar in general habitus to T. arizonensis and T. pseudovicina, being more elongate-cylindrical than barrel- shaped (Goeden and Teerink 1998, 1999). Trupanea imperfecta and T. arizonensis are also similar, in that the meso- and metatho- rax are circumscribed by verruciform sen- silla and the anterior thoracic spiracle bears three ovoid papillae (Goeden and Teerink 1998). However, there are slight differences between these two species in that the rugose pads laterad of the mouth lumen are serrat- ed in 7. imperfecta, but not in T. arizonen- sis. Moreover, the metathoracic lateral spi- racular complex is slightly different, with a stelex sensillum and two verruciform sen- silla in 7. imperfecta, and three verruciform sensilla in 7. arizonensis (Goeden and Teer- ink 1999). Trupanea pseudovicina is simi- lar to 7. imperfecta in having serrated ru- gose pads laterad of the mouth lumen (Goe- a sensillum, 3—verruciform sensilla; (F) sixth abdominal segment, |—spiracle, 2—verruciform sensilla: (G) cau- dal segment, 1—stelex sensillum, 2—verruciform sensillum, 3—rima, 4—interspiracular process; (H) interme- diate sensory complex, |—medusoid sensillum, 2—stelex sensillum. 232 den and Teerink 1998). It differs from T. imperfecta by having four, not three, papil- lae on the anterior spiracle; the meso- and metathorax not circumscribed by verruci- form sensilla, and a stelex sensillum and two, not one, verruciform sensilla in the metathoracic lateral spiracular complex (Goeden and Teerink 1998). As with the other Trupanea species studied by us to date, the lateral spiracular complex is unique to 7. imperfecta (Goeden and Teer- ink 1997, 1998, 1999; Goeden et al. 1998a, b; Headrick and Goeden 1991; Knio et al. 1996, Teerink and Goeden 1997). The puparium of 7. imperfecta is larger than T. actinobola and T. californica (Goe- den et al. 1998b; Headrick and Goeden 1991), wider but shorter than T. arizonensis and 7. pseudovicina (Goeden and Teerink 19985 1999): ACKNOWLEDGMENTS We are grateful to David Headrick for his helpful comments on an earlier draft of this paper. LITERATURE CITED Foote, R. H., E L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York. Goeden, R. D. 1987. Life history of Trupanea con- jJuncta (Adams) on Trixis californica Kellogg in southern California (Diptera: Tephritidae). Pan- Pacific Entomologist 63: 284-291. 1988. Life history of Trupanea imperfecta (Coquillett) on Bebbia juncea (Bentham) Greene in the Colorado Desert of southern California (Diptera: Tephritidae). Pan-Pacific Entomologist 64: 345-351. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Goeden, R. D. and J. A. Teerink. 1997. Life history and description of immature stages of Trupanea signata Foote (Diptera: Tephritidae) on Gnaphal- ium luteo-album L. in southern California. Pro- ceedings of the Entomological Society of Wash- ington 99: 748-755. . 1998. Life history and description of imma- ture stages of Trupanea pseudovicina Hering (Diptera: Tephritidae) on Porophyllum = gracile Bentham (Asteraceae) in southern California. Pro- ceedings of the Entomological Society of Wash- ington 100: 361-372. . 1999. Life history and description of imma- ture stages of Trupanea arizonensis Hering (Dip- tera: Tephritidae) on Trixis californica Kellogg var. californica (Asteraceae) in southern Califor- nia. Proceedings of the Entomological Society of Washington 101: 75-85. Goeden, R. D., J. A. Teerink, and D. H. Headrick. 1998a. Life history and description of immature stages of Trupanea jonesi Curran (Diptera: Te- phritidae) on native Asteraceae in southern Cali- fornia. Proceedings of the Entomological Society of Washington 100: 126-140. . 1998b. Life history and description of im- mature stages of Trupanea actinobola (Loew) (Diptera: Tephritidae) on Acamptopappus sphaer- ocephalus (Harvey and Gray) Gray (Asteraceae) in southern California. Proceedings of the Ento- mological Society of Washington 100: 674-688. Headrick, D. H. and R. D. Goeden. 1991. Life history of Trupanea californica Malloch (Diptera: Te- phritidae) on Gnaphalium spp. in southern Cali- fornia. Proceedings of the Entomological Society of Washington 93: 559-570. Knio, K. M., R. D. Goeden, and D. H. Headrick. 1996. Descriptions of immature stages of Trupanea ni- gricornis and T. bisetosa (Diptera: Tephritidae) from southern California. Annals of the Entomo- logical Society of America 89: 1-11. Munz, P. A. 1974. A Flora of Southern California. Uni- versity of California Press, Berkeley and Los An- geles. Teerink, J. A. and R. D. Goeden. 1998. Description of immature stages of Trupanea conjuncta (Adams) (Diptera: Tephritidae). Proceedings of the Ento- mological Society of Washington 100: 431—438. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 233-241 THE MICROCADDISFLY GENUS ITHYTRICHIA EATON (TRICHOPTERA: HYDROPTILIDAE) IN NORTH AMERICA STEPHEN R. MOULTON II, STEVEN C. HARRIS, AND JOSEPH P. SLUSARK (SRM, JPS) U.S. Geological Survey, National Water Quality Laboratory, Biological Unit, PO. Box 25046, MS 407, Denver, CO 80225, U.S.A. (e-mail, SRM: smoulto@usgs.gov); (SCH) Department of Biology, Clarion University, Clarion, PA 16214, U.S.A. Abstract.—The distribution and taxonomy of the microcaddisfly genus /thytrichia Eaton in North America is reviewed. Males and females of /. clavata Morton, [. mazon Ross, and I. mexicana Harris and Contreras-Ramos are illustrated, and a key is provided for their separation. Females of /. mazon and I. mexicana are described for the first time; the female of J. clavata is redescribed. Key Words: The holarctic genus /thytrichia (Eaton 1873) is a small group of microcaddisflies with six species worldwide (Marshall 1979, Morse 1993) belonging to the subfamily Hydroptilinae, tribe Orthotrichiini. Three species, [. clavata Morton, J. mazon Ross, and J. mexicana Harris and Contreras-Ra- mos, are found in North America (Morse 1993). Adults are distinguished from those of other nearctic hydroptilid genera by the presence of ocelli, a 0-3-4 tibial spur count, mesoscutellum without transverse suture, and posterodorsal margin of mesoscutellum separated from posterior margin by a nar- row strap (Moulton and Stewart 1996). The laterally compressed abdomen having dor- sal and ventral membranous lobes easily identifies larvae of I/thytrichia; the larval case is composed entirely of silk and is purse-like with a small circular anterior opening (Wiggins 1996). Morphological characters have not been discovered to dis- tinguish the larvae to species. Before this study, only the female of /. clavata was known. In this paper, we describe for the first time the females of /. mazon and I. mexicana. Males of the three species are Trichoptera, Hydroptilidae, /thytrichia, taxonomy, nearctic distribution reillustrated with accompanying distribu- tional notes. Keys for separating males and females of the three North American spe- cies are also provided. Material examined in this study is depos- ited at the Arkansas State University Muse- um of Zoology, Jonesboro (ASUMZ), the California Academy of Sciences, San Fran- cisco (CAS), the C. P. Gillette Museum of Arthropod Diversity, Colorado State Univer- sity, Fort Collins (CSU), the Illinois Natural History Survey, Champaign (INHS), the Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. (NMNH), the Ohio Biological Survey, Co- lumbus (OBS), the University of Minnesota, St. Paul (UM), the University of North Tex- as, Denton (UNT), and in the research col- lection of the senior author (SRM). Al- though most specific characters may be dis- cerned by using a dissecting microscope (60—100X), it is necessary to view some of the female genitalic characters (e.g., sper- mathecal sclerite) by using a compound mi- croscope (100—400). Morphological ter- minology follows that of Marshall (1979). Length is measured from the tip of the head to the posterior tip of the forewings. 234 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ithytrichia clavata Morton (Figs; 174. 7) Ithytrichia clavata Morton 1905:67. Ross (1944) described the female of J. clavata, however, the discovery of the fe- males of [. mazon and I. mexicana neces- sitates a redescription for comparison. Female description.—Length 2.9-—3.5 mm. 21 antennal segments. Light brown in alcohol. Sternite VI with short acute ventro- mesal process. Sternite VII in ventral view dome-shaped. Sternite VIII in ventral view parallel-sided, posterior margin with series of stout setae, each arising from a membra- nous tubercle; ventral sclerite widening posteriorly, posterior margin with broad concavity, bi-lobed, each lobe with a mem- branous pocket; two pairs of lateral apo- demes. Segment IX bullet-shaped in ventral view; one pair of lateral apodemes. Seg- ment X button-like with a pair of short cer- ci. Apodemes slender and rod-like, one pair extending from anterior end of segment X apodemes, other pair from anterior edge of segment VIII, both pairs extending to seg- ment VI, with anterior apices gently curv- ing mesad. Spermathecal sclerite in ventral view with anterior membranous and circu- lar; basal one-third sclerotized on lateral margins, anterior apices angled laterad, gradually tapering posteriorly to pointed, incurved apices, middle portion of sclero- tized apparatus arrowhead-shaped; posterior two-thirds widest at base, narrowing in middle to tubular apex, middle portion with lateral patches of minute spines. Material examined.—USA: CALIFOR- NIA, Colusa Co., Bear Creek, 26 km E Clearlake Highlands, 28-VII-1974, P. Peter- son, 6 d (CAS); Lake Co., elev. 402 m, 16- IX-1949, H. P. Chandler, 1 6 (CAS); Napa Co., Capell Cr., 7-VI-1952, 1 d (CAS); Big Canyon Creek, 13 km NE Middletown, 23- VI-1974, P. Peterson, 1 6 (CAS); FLORI- DA, Jackson Co., Florida Caverns State Park, 4-V-1970, 2 6 (NMNH); ILLINOIS, Galena River, Council Hill, 26-VI-1940, Mohr and Riegel, 10 ¢ (INHS); MAINE, Ashland, 29-VII-1924, 1 ¢ (CAS); Orono, Lake Pushaw, 1-3-VIII-1966, W. W. Wirth, 3 36 (NMNH); Oxbow (T9RS), 22-VII- 1961, A. Brower, 2 6 (NMNH); Allagash, 29-VII-1959, 32 ¢ (NMNH); same but, 30- VII-1959, 130 36 (NMNH); same but, 5- VIII-1959, 8 36 (NMNH); MISSOURI, Gasconade Co., Gasconade River, Held’s Island Access, 28-VIII-1990, B. C. Poulton, 3 6 (UNT); Maries Co., Gasconade River, Island Ford Resort @ Hwy 42, 7-VIII, 1990, B. C. Poulton, | d (UNT); same but, Paydown Access, SW Belle, 15-VII-1990, 1 6 (UNT); Osage Co., Gasconade River, Hwy 89 @ Dallas Ferry Access, 27-IX- 1990, B. C. Poulton, 2 6 (UNT); PENN- SYLVANIA, Presque Isle, 15-VIII-1947, 1 3 (CAS); Chemung River, Athens, 8-VII- 1937, J. Eddleston, 23 ¢ (INHS); TEXAS, Brewster Co., county park, 8 km S Mara- thon, 22-VI-1994, B. Kondratieff, 5 6, (CSU); Edwards Co., South Llano River @ Paint Rock Springs, Hwy 337, 12-VI-1992, B. Kondratieff, 1 ¢ (CSU); Hays Co., Blan- co River @ Post Rd., 14-V-1991, S. Tie- mann, 3 6 (UNT); Palo Pinto Co., Brazos River, TX Hwy 4, 23-III-1972, Stark and Rhame, | 6, 1 2 (NMNH), same but, 6- V1-1995;" Ds C. Houghton, 27, 6-592 (UNT); Randall Co., Prairie Dog Town Fork of Red River, Palo Duro Canyon State Park, water crossing No. 5, 12-IX-1997, S. R. Moulton and G. W. Easley, 2 6, 1 @ (SRM); WISCONISIN, Door Co., Egg Har- bor, 13-VIII-1940, C. O. Mohr, 21 6, 15 9 (INHS); same but, Ephraim, WI, ?-?-1957, 1 2 (NHS); CANADA: MANITOBA, Lake Manitoba, 5 km W Delta, 3-VIII- 1967, D. Webb, 9 d (INHS); QUEBEC, Ot- tawa River, Quyon, 1-VIII-1976, O. S. Flint, Jr, 4 d, 1 2 (NMNH); Norway Bay, 4-VIII-1973, O. S. Flint, Jn, 4 6, 4 2 (NMNH). Discussion.—Ithytrichia clavata has a holarctic distribution (Fischer 1961); it is widely distributed throughout the United States and southern Canada (Wiggins and Parker 1997). Houghton and Stewart (1998) reported on the seasonal flight periodicity VOLUME 101, NUMBER 2 ote: 220 Ta TIER tN Fig. 1. Ithytrichia clavata, male genitalia. A, Left lateral. B, Dorsal. C, Ventral. D, Phallus. Nn 236 of J. clavata from the Brazos River in north-central Texas. Specimens of /. clavata from the eastern United States determined before 1944 should be checked to ensure they are not misidentifications of J. mazon. We found this to be true in some of the material we examined. Ithytrichia mazon Ross (igs: 2;,5;.'8) Ithytrichia mazon Ross 1944:124. Female description.—Length 3.3—4.0 mm. 21 antennal segments. Light brown in alcohol. Sternite VI with short acute ventro- mesal process. Sternite VII in ventral view dome-shaped. Sternite VIII in ventral view parallel-sided, posterior margin with series of stout setae, each arising from a membra- nous tubercle; ventral sclerite widening posteriorly, apex truncate. Segment IX bul- let-shaped in ventral view, apex membra- nous. Segment X button-like with a pair of short cerci. Apodemes slender and rod-like, one pair extending from anterior end of seg- ment X apodemes, other pair from anterior edge of segment VIII, both pairs extending to segment VI, with anterior apices gently curving laterad. Spermathecal sclerite in ventral view with anterior a membranous funnel-shape; basal one-third sclerotized laterally and bowed, middle portion of sclerotized apparatus arrowhead-shaped; posterior two-thirds widest at base, narrow- ing in middle to tubular apex, middle por- tion with lateral patches of minute spines. Material examined.—USA: ARKAN- SAS, Logan Co., Sixmile Creek, 23-V- 1986, H. W. Robison, 1 ¢d (INHS); ILLI- NOIS, Mazon, along Mazon Creek, 16-VI- 1938, B. D. Burks, holotype ¢ (INHS); Se- rena, Indian Creek, 16-VI-1939, B. D. Burks, 1 6 (INHS); OHIO, Adams Co., Hills Fork-Eagle Creek, SR 125, W West Union, OH, 27-VI-1993, B. Armitage, UV trap, 37°¢3.52 2 (OBS); OKLAHOMA, Latimer Co., 10-VI-1931, R. D. Bird, 3 6, 3 ¢ (1 vial each sex, INHS). Discussion.—This species appears to be PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON restricted to small streams in the Ohio and middle Mississippi River drainages. New State records are presented here for Ohio and Oklahoma. In addition to the states list- ed above, /. mazon has also been recorded from the Salt River drainage, Spencer Co., Kentucky (Resh 1975). Ithytrichia mexicana Harris and Contreras-Ramos (Figs. 3, 6, 9) Ithytrichia mexicana Harris and Contreras- Ramos 1989:176. Female description.—Length 3.5 mm. 20 antennal segments. Brown in alcohol. Ster- nite VI with short acute ventro-mesal pro- cess. Sternite VII in ventral view subrect- angular. Sternite VIII in ventral view par- allel-sided, posterior margin with series of stout setae, each arising from a membra- nous tubercle; ventral sclerite with antero- lateral flaps, each bearing a series of short setae, narrowing to truncate posterior mar- gin, posterior margin with a short mesal process. Segment IX bullet-shaped in ven- tral view. Segment X button-like with a pair of short cerci. Apodemes slender, rod-like, extending from segment IX to VII, anterior apices straight, second pair of lateral apo- demes branching from mesal pair at ap- proximately the anterior margin of segment VIII. Spermathecal sclerite in ventral view with anterior portion forming a funnel- shaped collar constricted in middle; basal one-half sclerotized laterally with anterior apices acutely produced mesad, gradually tapering posteriorly to pointed, incurved apices, middle portion of sclerotized appa- ratus arrowhead-shaped; posterior one-half widest at base with lateral patches of mi- nute spines, narrowing in middle to tubular apex, middle portion with short mesal in- cision; anterior and posterior portions join- ing with lateral membranous lobes. Material examined.—USA: ARIZONA, Coconino Co., Oak Creek at Sterling Spring Fish Hatchery, U.S. Hwy 894A, elev. 1,829 m, 4-5-VI-1993, S. Moulton and K. Alex- VOLUME 101, NUMBER 2 N Fig. 2. Ithytrichia mazon, male genitalia. A, Left lateral. B, Dorsal. C, Ventral. D, Phallus. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Ithytrichia mexicana, male genitalia. A, Left lateral. B, Dorsal. C, Ventral. D, Phallus. VOLUME 101, NUMBER 2 239 Figs. 4-6. ander, 2 6 (UM); NEW MEXICO, Sandov- al Co., Rito de los Frijoles @ Bandelier Na- tional Monument, 10.4 km S Los Alamos, elev. 1,839 m, 2-VIII-1994, L. FE Carter, 1 3 (INHS), same but 1-3-VIII-1997, J. Slu- sark and B. Richards, UV trap, 1 2 (NMNH). MEXICO: TAMAULIPAS, Municipio de Gomez Farias, Rio Frio @ headwaters, La Poza Azul, 6 km S Gomez Ithytrichia female genitalia, ventral. 4, 1. clavata. 5, I. mazon. 6, I. mexicana. Farias, 7-VIII-1988, A. Contreras and A. Moreno, blacklight, holotype ¢ (NMNH). Discussion.—This species was originally described by Harris and Contreras-Ramos (1989) based on a single male collected from the headwaters of the Rio Frio, Ta- maulipas, Mexico. The Arizona record list- ed above was erroneously reported as a new Arizona state record for /. clavata by Moul- 240 Figs. 7-9. ton et al. (1994). Ithytrichia mexicana is re- ported herein from the United States for the first time and the species is now represented by a total of five specimens (4 males, | fe- male). On the basis of known collection rec- ords, it appears to have an affinity for small cold mountain streams in the southwestern United States and northern Mexico at about 1,800 m in elevation. Undetermined /thytrichia Material We examined several larvae and pupae in this study that could not be positively determined to species. However, specula- tion as to their probable identity is indicated 8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 9 Ithytrichia spermathecal sclerites, ventral. 7, /. clavata. 8, I. mazon. 9, I. mexicana. in brackets for a few records based on dis- tributional information. Material examined.—ARIZONA, Gila Co., Christopher Creek, AZ 260, ca. 40 km NE Payson, elev. 1,792 m, 14-VI-1985, A. R. Brigham, 2 larvae (INHS) [/. mexicana]; Christopher Creek, ?-?-1985, A. Brigham, 8 larvae (INHS) [/. mexicana]; Lower Hor- ton Creek, 12-VIII-1937, Tazwell, 1 larva (INHS); ARKANSAS, Randolph Co., Jane’s Creek, AR Hwy 90, S Ravenden Springs, III-1985, S. R. Moulton, | larva (ASUMZ); MAINE, Washington Co., Nar- raguagus River, island, 6-VIH-1973, T. Min- go, 2 pupae, 5 larvae (NMNH) [/. clavata]; VOLUME 101, NUMBER 2 TEXAS, Pecos River, Sheffield, 6-I-1976, J. Davis, 6 larvae (NMNH) [/. clavata]; WISCONSIN, Madison, Fox River, Lake Winnebago, ?-?-1954, K. M. Mackenthun, 1 larva (INHS) [/. clavata]. Key to the North American Species of [thytrichia Pe alex (Ete S213) aoe toga ke aoe crea © Sen 2 Remialex(Gios'e4—6))eaaten concrete ce) so yan) t= 4 2. Inferior appendages in ventral view tapering posterad, apices rounded (Figs. IC, 3C) .... 3 — Inferior appendages in ventral view rectangu- lar, apices truncate (Fig. 2C) 3. Posterolateral margins of tergum IX sclerotized, hooked laterad (Fig. 3B); apex of subgenital plate emansinate (ries3 6). sates ae I. mexicana — Posterolateral margins of tergum IX rounded, not hooked (Fig. 1B); apex of subgenital plate dome-shaped (Fig. 1C) 4. Ventral sclerite of VIII gradually widening pos- teriorly (Figs. 4, 5); lateral sclerites of sper- mathecal sclerite less than one-half length of entire apparatus, ending anterolaterally in an- eledvorroundedvapices (Figs: 75.8) {2.2 422: 5 — Ventral sclerite of VII narrowing posteriorly (Fig. 6); lateral sclerites of spermathecal sclerite about one-half length of entire apparatus, an- terior apices curving inward to join arrowhead- shaped mesal process (Fig. 9) 5. Ventral sclerite of VIII with concave posterior margin (Fig. 4); lateral processes of sperma- thecal sclerite short, anterolaterally angled out- ward (Fig. 7) I. clavata — Ventral sclerite of VIII with truncated posterior margin (Fig. 5); lateral processes of sperma- thecal sclerite elongate, rounded anteriorly (Fig. 8) I, mazon I, mazon I. clavata eRe oe I. mexicana ACKNOWLEDGMENTS We thank Lisa Carter, Oliver Flint, Jr. (NMNH), George Harp (ASUMZ), David Houghton (UM), Vincent Lee (CAS), Boris Kondratieff (CSU), and Kathy Zieders (INHS) for loaning specimens from their collections. Brian Armitage (OBS) provid- ed us with light trap material from which the female of J. mazon was discovered. Bra- dy Richards (USGS) assisted JPS in the col- lection of the J. mexicana female. Brian Ja- cobs (Bandelier National Monument, NM) greatly facilitated the completion of this 241 study by providing a scientific collector permit. Brian Armitage, Gregg Easley (USGS), Oliver Flint, Jr., Boris Kondratieff, Jon Raese (USGS), and John Sandberg (USGS) reviewed drafts of the manuscript. LITERATURE CITED Eaton, A. E. 1873. On the Hydroptilidae, a family of the Trichoptera. Transactions of the Entomological Society of London 1873: 125-150. Fischer, F C. J. 1961. Trichopterorum Catalogus Vol. Il: Philopotamidae, Hydroptilidae, Stenopsychi- dae. iv + 190 pp. Amsterdam. Harris, S. C. and A. Contreras-Ramos. 1989. /thyrichia mexicana (Trichoptera: Hydroptilidae), a new spe- cies of caddisfly from Mexico. Entomological News 100: 176-178. Houghton, D. C. and K. W. Stewart. 1998. Seasonal flight periodicities of six microcaddisflies (Trichop- tera: Hydroptilidae, Glossosomatidae) in the Brazos River, Texas, with notes on larval biology and site records. Entomological News 109: 103—109. Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bulletin of the British Museum of Natural History (Entomology) 39: 135-239. Morse, J. C. 1993. A checklist of the Trichoptera of North America, including Greenland and Mexico. Transactions of the American Entomological So- ciety 119: 47-93. Morton, K. J. 1905. New American Hydroptilidae. New York State Museum Bulletin 86: 63-85. Moulton, S. R., If, K. W. Stewart, and K. L. Young. 1994. New records, distribution, and taxonomic status of some northern Arizona caddisflies (Tri- choptera). Entomological News 105: 164-174. Moulton, S. R., Hand K. W. Stewart. 1996. Caddisflies (Trichoptera) of the Interior Highlands of North America. Memoirs of the American Entomologi- cal Institute Vol. 56, 313 pp. Resh, V. H. 1975. A distributional study of the cad- disflies of Kentucky. Transactions of the Kentucky Academy of Science 36: 6—16. Ross, H. H. 1944. The caddisflies, or Trichoptera, of Illinois. Bulletin of the Illinois Natural History Survey 23: 1-326. Wiggins, G. B. 1996. Larvae of the North American caddisfly genera (Trichoptera), 2nd Edition. Uni- versity of Toronto Press, Toronto. 457 pp. Wiggins, G. B. and C. R. Parker. 1997. Caddisflies (Trichoptera) of the Yukon, with analysis of the Beringian and holarctic species of North America. pp. 787-866. Jn Danks, H. V. and J. A. Downes, eds., Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa. 1034 pp. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 242-262 CHECKLIST AND HOST PLANTS OF THE TREEHOPPERS (HEMIPTERA: MEMBRACIDAE) OF NORTH CAROLINA CHRISTOPHER H. DIETRICH, MARK J. ROTHSCHILD, AND LEwIs L. DEITZ (CHD) Center for Biodiversity, Illinois Natural History Survey, 607 East Peabody Drive, Champaign, IL 61820, U.S.A. (e-mail: dietrich @ denr1.igis.uiuc.edu); (MJR) Mary- land Department of Agriculture, Plant Protection Section, 27722 Nanticoke Road, Unit 2, Salisbury, MD 21801, U.S.A.; (LLD) Department of Entomology, North Carolina State University, Raleigh, NC 27695-7613 (e-mail: Idietz@ppentl.ppath.ncsu.edu) Abstract.—Based on recent collecting and an examination of museum specimens, at least 89 treehopper species (Hemiptera: Membracidae) occur in North Carolina, of which 26 species represent new state records. The presence of 13 species previously recorded from North Carolina could not be verified based on available material. Three previous North Carolina records were found to be based on misidentifications. The known distribution (by county) and host plants in North Carolina are given for each species. Photographs of rep- resentative taxa and a host plant index are included. Stictocephala bisonia Kopp and Yonke is reinstated as a valid name (and not a junior synonym of Ceresa alta Walker). Key Words: The family Membracidae (Figs. 1—10) in- cludes more than 3,000 described treehopper species worldwide (McKamey 1998). About 260 are known to occur in temperate North America. Many of these species are restrict- ed to the mixed hardwood forests and sa- vannas of the eastern United States, where they exploit a variety of woody and herba- ceous plants as hosts for oviposition, feed- ing, or both. Most North American treehop- per species are univoltine, solitary, and cryp- tic as both immatures and adults, and, hence are seldom noticed or collected. A few spe- cies, however, are multivoltine, gregarious (Figs. 7, 8), ant-mutualistic (Figs. 8, 10), or aposematic (Fig. 7), and are therefore some- what conspicuous. Three kinds of life cycles are common among North American tree- hoppers (Table 1). Many members of cate- gory III that feed and oviposit on oaks (Figs. 5, 8) are usually found as adults for only a few weeks in May or June, depending on the location within the state. Membracidae, treehopper, taxonomy, biogeography, insect-plant interactions Records of North Carolina treehoppers were summarized by Metcalf (1915), Brim- ley (1938, 1942), Wray (1950, 1967), Met- calf and Wade (1965), and Kopp and Yonke (1973a—c, 1974: distribution maps). Kopp and Yonke’s series provided keys to many species in eastern North America. Deitz et al. (1976), McGiffen and Neunzig (1985), and Hargrove (1986) gave further records of North Carolina treehoppers associated with soybeans, grapes, and black locust, re- spectively. The objectives of the present work were to document the species richness of North Carolina treehoppers, summarize the known distributions (Fig. 11: county map) and host plant associations within the state, and pro- vide an up-to-date checklist following cur- rent nomenclature. Although host records for numerous species have been published (e.g., Funkhouser 1917; Ball 1931; Kopp and Yonke 1973a—c, 1974), the extent to VOLUME 101, NUMBER 2 243 Figs. 1-6. Micrutalis calva. 4, Campylenchia latipes. 5, Glossonotus univittatus. 6, Stictocephala militaris. which host associations vary geographical- ly is poorly documented and most pub- lished records have not been verified through rearing of immatures. MATERIALS AND METHODS The species records below are based on specimens in the following collections: the North Carolina State University Insect Col- lection, Raleigh [NCSU: includes recent material from the authors]; Mark J. Roths- Representative treehoppers of North Carolina. 1, Microcentrus caryae. 2, Acutalis tartarea. 3, child Collection, % Maryland Department of Agriculture, Salisbury [MJRC]; Florida State Collection of Arthropods, Gainesville [FSCA]; North Carolina Department of Ag- riculture, Raleigh [NCDAJ]; and the Nation- al Museum of Natural History, Smithsonian Institution, Washington, D.C. [USNM]. C. S. Brimley’s historically important material is housed at the collections of NCDA (spec- imens and card files) and NCSU (speci- mens). 244 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-10. Representative treehoppers of North Carolina (continued). 7, Platycotis vittata, aggregation of teneral adults. 8, Vanduzea arquata, aggregation of adults and nymphs attended by ants. 9, Ophiderma evelyna: left, female (green), right, male (brown). 10, Entylia carinata (left) and Publilia concava (right, attended by ant). VOLUME 101, NUMBER 2 245 Table 1. Three major kinds of life cycles in treehoppers of eastern North America (modified from Kopp and Yonke 1973a). Category: Taxa Overwintering I: Polyglyptini, Platycotis vittata, probably Cam- pylenchia latipes, some Vanduzea and some Ceresini II: Acutalini, most Ceresini, some Micrutalis eggs overwinter under bark in young twigs woody hosts Il: Smiliini, Microcentrus spp., Enchenopa binota- ta complex, some Mi- crutalis eggs overwinter under bark in young twigs woody hosts Records are based on adult specimens except as noted. Among species of Ceresi- ni, males are generally required for positive identification (Kopp and Yonke 1979); con- sequently, females without associated males often could not be identified. Each entry in- cludes a list of counties in North Carolina from which the species has been recorded, the seasonal distribution (earliest and latest calendar date of collection of adults), and North Carolina host records. A few speci- mens bore labels indicating a locality situ- ated on the border of two or three counties; these were considered to occur in all of the counties involved. Except as noted, only North Carolina host associations based on specimens ex- amined are reported here. Hosts marked with an asterisk (*) are those from which both nymphs and adults have been collect- ed. Other plants listed are those from which only adults have been collected, so some may not be true hosts. Botanical nomencla- ture follows Kartesz (1994), Liberty Hyde Bailey Hortorium (1976), and Radford et al. (1968). To conserve space, botanical com- mon names and the authors of plant scien- tific names are given only in the alphabet- ical host index. To facilitate comparisons with The In- adults overwinter in litter nymphs feed and develop of of Development Generations Per Year 2 (most are bivoltine) on herbaceous or woody host plants nymphs feed and develop 1 (univoltine) on herbaceous host plants (many females require preoviposition period before laying eggs) nymphs feed and develop | (most are univoltine) on same woody hosts used for oviposition sects of North Carolina (Brimley 1938, 1942, Wray 1950, 1967), and The Treehop- pers of Missouri (Kopp and Yonke 1973a— c, 1974), names from those works that dif- fer from current nomenclature are given for each entry in square brackets (occasionally with other notes on synonymy). For addi- tional synonymy, see Fascicle 1, Membra- cidae, and its supplements, in the General Catalogue of the Hemiptera (Funkhouser 1927, Metcalf and Wade 1965, McKamey 1998) and the associated bibliographies (Metcalf and Wade 1963, Deitz and Kopp 1987, Deitz 1989). For convenience, the checklist is ar- ranged alphabetically by genus and species. Table 2 summarizes the placement of the included genera into tribes and subfamilies. RESULTS Recent collecting in North Carolina yielded numerous new state, county, and host records. Figure 11 indicates the num- ber of treehopper species recorded for each of North Carolina’s 100 counties. The high- er species richness recorded near Raleigh (Wake County, 68 species), Asheville (Bun- combe County, 39 species), Boone (Watau- ga county, 30 species), and Charlotte PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 246 A, V5; SY A COd|Wed = a at s@AOURH MON 4A Japuad ‘AJUNOS YORI WOIJ Papsoda1 saisads saddoysen jo saquinu yM dew vurporeg yWON ps4 AN PRS a Wise 7 puejjoos uolun Binquayoeyy uosiapua}y ‘TL Sld eiuen|Asuel | meant’ Powe 419d \e uosyoer, Zseier) VOLUME 101, NUMBER 2 Table 2. Yonke 1979, and Deitz and Dietrich 1993). Subfamily Stegaspidinae: Tribe Microcentrini: Microcentrus Stal 1869 (Fig. 1). Subfamily Membracinae: Tribe Hoplophorionini: Playcotis Stal 1869 (Fig. 7). 247 Summary of the classification of North Carolina Membracidae (based on Deitz 1975, Kopp and Tribe Membracini: 7ylopelta Fowler 1894; Campylenchia Stal 1869 (Fig. 4); Enchenopa Amyot & Serville 1843. Subfamily Smiliinae: Tribe Acutalini: Acutalis Fairmaire 1846 (Fig. 2). Tribe Micrutalini: Micrutalis Fowler 1895 (Fig. 3). Tribe Ceresini: Hadrophallus Kopp and Yonke 1979; Stictolobus Metcalf 1916; Tortistilus Caldwell 1949: Stictocephala Stal 1869 (Fig. 6); Spissistilus Caldwell 1949. Tribe Amastrini: Vanduzea Goding 1892 (Fig. 8). Tribe Smiliini: Archasia Stal 1867; Carynota Fitch 1851; Glossonotus Butler 1877 (Fig. 5); Heliria Stal 1867: Telamona Fitch 1851; Thelia Amyot and Serville 1843; Atymna Stal 1867; Cyrtolobus Goding 1892; Ophi- derma Fairmaire 1846 (Fig. 9); Smilia Germar 1833; Xantholobus Van Duzee 1908. Tribe Polyglyptini: Publilia Stal 1866 (Fig. 10, right); Entylia Germar 1833 (Fig. 10, left). (Mecklenburg County, 29 species) reflects greater collecting effort in those areas. With at least 89 species, North Carolina ranks third among the few states for which treehopper checklists have been published. New York ranks first (100 currently recog- nized species: Leonard 1928) and Ohio sec- ond (93 currently recognized species, in- cluding two presumed present based on rec- ords in neighboring states: Osborn 1940), but dubious identifications may have inflat- ed the accuracy of counts for those states. Among treehoppers, males, females, or both may be polymorphic with respect to pronotal shape and coloration (Figs. 7, 9). Frequently, previous workers incorrectly identified a single polymorphic species as two or more distinct species. Thirteen species previously reported from North Carolina whose occurrence in the state we could not verify and three pre- Table 3. Comparison of treehopper diversity in the world, the Nearctic Region, and North Carolina (based on McKamey 1998 and the present work). Geographic Sub- Area Species Genera Tribes families World Sida 397 49 12 Nearctic Region 258 62 13 6 North Carolina 89 26 9 3 viously misidentified species—Hadrophal- lus constans (Walker), Spissistilus unifor- mis (Fairmaire), and Telamona concava Fitch, are included below in square brack- ets. Five additional species reported from ad- jacent states may eventually be found in North Carolina, but are not included in the checklist: Palonica pyramidata (Uhler), Stictocephala albescens (Van Duzee), and Thelia uhleri Stal, all reported from Virgin- ia (Kopp and Yonke 1973b, 1974: distri- bution maps); Helonica excelsa (Fairmaire), reported from South Carolina (Kopp and Yonke 1974: map); and Telamona compac- ta Ball (specimen [USNM] from Rocky Bottom, South Carolina [<10 miles from Transylvania County, North Carolina]). Table 3 shows the species richness of North Carolina’s treehopper fauna com- pared to the Nearctic Region and the world. Especially well represented in North Caro- lina are the genera Cyrtolobus (21 species), Telamona (10), Ophiderma (7), and Heliria (6). All species recorded are native to North Carolina and nearly all are endemic to the eastern U.S. North Carolina treehoppers have been collected on at least 45 plant genera in 19 families, including 12 genera of Asteraceae, 248 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 9 of Fabaceae, and 3 of Fagaceae. More- over, 39 species (43 percent of the state’s membracid fauna) may be found on oaks (Fagaceae), with 16 species of Quercus list- ed below as hosts. CHECKLIST OF NORTH CAROLINA MEMBRACIDAE Acutalis tartarea (Say 1830) [in part as Acutalis tartarea var. semicrema (Say 1830) in Brimley 1938]. Fig. 2. Counties: Alamance, Alexander, Anson, Ashe, Beaufort, Bladen, Buncombe, Burke, Ca- barrus, Caldwell, Carteret, Chatham, Da- vidson, Duplin, Edgecombe, Forsyth, Granville, Haywood, Henderson, Hyde, Jackson, Johnston, Mecklenburg, Nash, Onslow, Orange, Pamlico, Pasquotank, Person, Polk, Rutherford, Stanly, Surry, Tyrrell, Wake, Warren, Washington, Wa- tauga, Wayne, Wilkes, Wilson. Seasonal distribution: 24 May—23 November. Host associations: Ambrosia artemisiifolia, Arundinaria sp., Bidens coronata, Eu- patorium capillifolium, Helianthus sp., Solidago sp. Archasia auriculata (Fitch 1851) [as A. gal- eata (Fabricius 1803), a preoccupied name, in Brimley 1938]. Counties: Blad- en, Buncombe, Columbus, Moore, Wake. Seasonal distribution: 9 May—27 August. Host associations: Quercus nigra, Q. ve- lutina*. Archasia_ belfragei Stal 1869. Counties: Bladen, Buncombe, Columbus, Hender- son, Macon, Wake. Seasonal distribution: 12 May—11 September. Host associations: Quercus alba*, Q. phellos. Notes: Brim- ley’s (1938) record of A. belfragei from Southern Pines (Moore County, June), actually represents a specimen (NCDA) of A. auriculata. Atymna castaneae (Fitch 1851) [as Cyrto- lobus castaneus (Fitch) in Brimley 1938]. Counties: Ashe, Bladen, Buncombe, Burke, Graham, Haywood, Jackson, Sampson, Surry, Swain, Watauga, Yan- cey. Seasonal distribution: 23 May-—31 August. Host associations: Castanea den- tata*, C. pumila*. [Atymna inornata (Say 1830)]. Records of this species in North Carolina (Metcalf 1915, Van Duzee 1917, Brimley 1938 [as Cyrtolobus inornata]) were not verified by the present authors. Atymna querci (Fitch 1851). Counties: Al- leghany, Ashe, Buncombe, Cabarrus, Cleveland, Duplin, Forsyth, Haywood, Macon, Martin, Nash, Orange, Vance, Wake, Warren, Washington, Watauga, Wayne, Yadkin. Seasonal distribution: 14 April-8 September. Host associations: Quercus alba*, Q. stellata*, Vitis rotun- difolia. Campylenchia latipes (Say 1824). Fig. 4. Counties: Alamance, Alexander, Allegh- any, Ashe, Avery, Bladen, Brunswick, Buncombe, Burke, Cabarrus, Caldwell, Chatham, Cleveland, Davidson, Davie, Duplin, Durham, Forsyth, Graham, Gran- ville, Guilford, Halifax, Haywood, John- ston, Jones, Macon, Madison, Martin, McDowell, Mecklenburg, Mitchell, Montgomery, Moore, New Hanover, Onslow, Orange, Pender, Polk, Ruther- ford, Sampson, Scotland, Stanly, Swain, Transylvania, Wake, Warren, Washing- ton, Watauga, Wayne, Wilkes, Yadkin. Seasonal distribution: [? 4 May (NCSU)], 3 June—21 November. Host associations: Medicago sativa, Solidago sp. Carynota marmorata (Say 1830). Counties: Haywood, Henderson, Jackson, Moore, Stanly, Wake, Watauga. Seasonal distri- bution: 6 June—30 July. Host associa- tions: Betula sp. Carynota mera (Say 1830). Counties: Blad- en, Currituck, Hyde, Mecklenburg, New Hanover, Pitt, Wake. Seasonal distribu- tion: [18 May, nymphs], 21 May—late Oc- tober. Host associations: Carya_ illino- inensis, C. sp*. Cyrtolobus arcuatus (Emmons 1854). Counties: Greene, Harnett, Hertford, Wake. Seasonal distribution: 24 April—12 May. Host associations: Quercus falca- ta*, Q. phellos. VOLUME 101, NUMBER 2 Cyrtolobus auroreus Woodruff 1924. NEW STATE RECORD. Counties: Ashe, Du- plin, Guilford, Macon, Mecklenburg, Wake, Wayne. Seasonal distribution: 23 April—19—20 June. Host associations: Quercus alba*, Q. prinus. Cyrtolobus celsus Van Duzee 1917 [as C. celsis (sic) in Brimley 1938]. Counties: Moore. Seasonal distribution: 22 May. Host associations: no data for North Car- olina. Cyrtolobus clarus Woodruff 1924. NEW STATE RECORD. Counties: Wake. Sea- sonal distribution: 4—25 May. Host as- sociations: Quercus palustris*, Q. phel- los*. [Cyrtolobus discoidalis (Emmons 1854)]. Brimley’s (1938) record of this species in North Carolina (Balsam, Jackson Coun- ty) could not be verified, but may be the basis for Kopp and Yonke’s (1973c: map) record. Cyrtolobus dixianus Woodruff 1924. NEW STATE RECORD. Counties: Cabarrus, Duplin, Franklin, Wake. Seasonal distri- bution: 26 April-26 May. Host associa- tions: Quercus alba, Q. falcata, Q. pal- ustris, Q. stellata*. Cyrtolobus fenestratus (Fitch 1851). Coun- ties: Bladen, Dare, Edgecombe, Forsyth, Greene, Hertford, Hoke, Johnston, Le- noir, Nash, Pasquotank, Robeson, Rock- ingham, Wake, Washington, Wilson. Sea- sonal distribution: [15 April, nymph], 24 April-19 May. Host associations: Quer- cus falcata*, Q. laevis, Q. margarettiae, Q. marilandica*, Q. nigra*, Q. palustris, Q. phellos*, Q. rubra*, Q. velutina, Q. virginiana. Notes: Brimley’s (1938) June record of C. fenestratus from Blowing Rock (Watauga County) was not verified by the present authors, while his July rec- ord from Blowing Rock may represent a specimen (NCDA) of C. puritanus Woodruff. Cyrtolobus flavolatus Woodruff 1924. NEW STATE RECORD. Counties: Bun- combe, Vance, Wake. Seasonal distribu- 249 tion: 18 May—18 June. Host associations: no data for North Carolina. Cyrtolobus fuliginosus (Emmons 1854). Counties: Cabarrus, Franklin, Greene, Harnett, Hertford, Johnston, Lenoir, Mar- tin, Mecklenburg, Pasquotank, Randolph, Wake, Warren, Washington, Wayne. Sea- sonal distribution: 23 April—26 May. Host associations: Quercus coccinea, Q. falcata*, Q. palustris, Q. phellos, QO. stel- lata, Q. velutina*. Cyrtolobus funkhouseri Woodruff 1924. NEW STATE RECORD. Counties: Mecklenburg, Stokes, Wake, Wilkes, Yadkin. Seasonal distribution: 13 May— 11 June. Host associations: Quercus pal- uSTriS. Cyrtolobus fuscipennis Van Duzee 1908. Counties: Ashe, Buncombe, Burke, Jack- son, Macon, Mecklenburg, Wake. Sea- sonal distribution: 24 April—22 July. Host associations: Quercus alba*, Q. nigra. Cyrtolobus griseus Van Duzee 1908. NEW STATE RECORD. Counties: Ashe, Burke, Montgomery, Vance, Wake. Sea- sonal distribution: 29 April—6 July. Host associations: Quercus alba, Q. stellata. Cyrtolobus inermis (Emmons 1854). NEW STATE RECORD. Counties: Bladen, Hoke, Rutherford, Wake. Seasonal distri- bution: 3 May—10 June. Host associa- tions: Quercus falcata*, Q. marilandica*, Q. stellata. Cyrtolobus maculifrontis (Emmons 1854). Counties: Ashe, Bladen, Duplin, Gran- ville, Vance, Wake, Watauga, Wayne. Seasonal distribution: [? March (NCSU)], 23 April—27 June. Host associations: Quercus alba*, Q. falcata. Cyrtolobus ovatus Van Duzee 1908. Coun- ties: Bladen, Hoke, Johnston, Moore, Richmond, Sampson, Wake. Seasonal distribution: 7 May—28 June. Host asso- ciations: Quercus laevis*, Q. marilandi- Ca: Cyrtolobus pallidifrontis (Emmons 1854). NEW STATE RECORD. Counties: Bun- combe, Currituck, Wake, Warren, Watau- ga. Seasonal distribution: 29 April—26 250 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON July. Host associations: Quercus alba, Q. stellata*. Cyrtolobus parvulus Woodruff 1924. NEW STATE RECORD. Counties: Bladen, Moore, Richmond. Seasonal distribution: 23 May-—17 June. Host associations: no data for North Carolina. Notes: Hosts re- ported elsewhere include two species of Quercus (Kopp and Yonke 1973c). Cyrtolobus pulchellus Woodruff 1924. NEW STATE RECORD. Counties: Ashe. Seasonal distribution: 19-20 June. Host associations: Quercus rubra. Cyrtolobus puritanus Woodruff 1924. Counties: Buncombe, Wake, Watauga, Yancey [as “Black Mountains,” which is on the Buncombe—Yancey border (USNM)]. Seasonal distribution: 26 May-—20 July. Host associations: no data for North Carolina. Notes: Brimley’s (1938) record of C. puritanus from Lake Toxoway (Transylvania Co.) was not ver- ified by the present authors. Hosts re- ported elsewhere include five species of Quercus (Kopp and Yonke 1973c). [Cyrtolobus sculptus (Fairmaire 1846)]. Rec- ords of C. sculptus in North Carolina (Goding 1893, Van Duzee 1917, Brimley 1938) were not verified by the present au- thors. Cyrtolobus togatus Woodruff 1924. NEW STATE RECORD. Counties: Buncombe, Cabarrus, Durham, Franklin, Harnett, Mecklenburg, Nash, Rockingham, Vance, Wake, Yadkin. Seasonal distribution: 23 April—23—30 June. Host associations: Quercus nigra*, Q. phellos*, Q. stellata. Cyrtolobus tuberosus (Fairmaire 1846). Counties: Bladen, Cabarrus, Camden, Caswell, Columbus, Craven, Franklin, Hertford, Hoke, Lenoir, Mecklenburg, Montgomery, Nash, Northampton, Robe- son, Rockingham, Wake, Warren, Wayne, Wilson. Seasonal distribution: 16 April— 1 June. Host associations: Quercus alba*, Q. margarettiae, Q. marilandica, Q. ni- gra, Q. prinus, Q. rubra, Q. stellata, Q. virginiana. Cyrtolobus vau (Say 1830). Counties: Al- leghany, Ashe, Avery, Buncombe, Ca- swell, Columbus, Duplin, Franklin, Hay- wood, Henderson, Jackson, Macon, Mecklenburg, Moore, Rockingham, Tran- sylvania, Vance, Wake, Yancey. Seasonal distribution: late March—5 September. Host associations: Quercus alba*, Q. pri- nus, Q. stellata. Enchenopa binotata (Say 1824) complex (see Notes, below, for discussion of the complex). Counties: Alamance, Allegh- any, Ashe, Buncombe, Carteret, Chat- ham, Gates, Graham, Guilford, Hay- wood, Iredell, Macon, Madison, McDow- ell, Moore, New Hanover, Richmond, Sampson, Stanly, Transylvania, Wake, Yancey. Seasonal distribution: [1 May, nymph], 15 May—3 October. Host asso- ciations: Carya sp., Cercis canadensis*, Juglans nigra*, Liriodendron tulipifera, Robinia pseudoacacia*, Viburnum prun- ifolium*. Notes: The Enchenopa binotata complex is thought to include nine bio- logically distinct North American spe- cies, each of which is associated with a different genus or species of deciduous woody host plant: (1) Carya spp., (2) Ce- lastris scandens, (3) Cercis canadensis, (4) Juglans cinerea, (5) J. nigra, (6) Lir- iodendron tulipifera, (7) Ptelea trifoliata, (8) Robinia pseudoacacia, and (9) Vibur- num spp. (Pratt and Wood 1992, 1993). Three published names are currently available for species within this complex, but the corresponding original descrip- tions lack host plant data as well as mor- phological criteria useful for distinguish- ing either the nymphs or adults from oth- er species in the complex (Pratt and Wood 1992). Pratt and Wood (1992) de- scribed the fifth instar nymphs of species in the complex and provided a key for their identification, but did not attempt to resolve the nomenclatural problems. Based on host data, specimens examined from North Carolina appear to represent five of the nine species in the complex: VOLUME 101, NUMBER 2 Cercis canadensis*: Buncombe and Wake Counties (10 June—5 July). Juglans nigra*: Ashe and Sampson County ({23 May, nymph] 5 July). Liriodendron tulipifera: Chatham County (2 July). Robinia pseudoacacia*: Macon County (5 August). Viburnum prunifolium*: Wake County (27 May). Entylia carinata (Forster 1771) [in part as E. concisa Walker 1851, and as E. sinu- ata (Fabricius 1798) in Brimley 1938; as E. bactriana Germar 1835, in Kopp and Yonke 1973b; E. carinata (Forster) in Remes-Lenicov 1973]. Fig. 10 (left). Counties: Alamance, Alexander, Allegh- any, Anson, Ashe, Avery, Beaufort, Ber- tie, Bladen, Buncombe, Burke, Cabarrus, Caldwell, Camden, Catawba, Chatham, Cherokee, Clay, Cleveland, Cumberland, Dare, Davidson, Davie, Duplin, Durham, Edgecombe, Gaston, Gates, Graham, Granville, Harnett, Haywood, Henderson, Hertford, Hoke, Hyde, Iredell, Jackson, Johnston, Lincoln, Macon, Madison, Mc- Dowell, Mecklenburg, Montgomery, Moore, Nash, New Hanover, Onslow, Orange, Pasquotank, Perquimans, Pitt, Polk, Rockingham, Sampson, Scotland, Stanly, Surry, Swain, Transylvania, Vance, Wake, Warren, Washington, Wa- tauga, Wayne, Yadkin, Yancey. Seasonal distribution: 13 March—19 December. Host associations: Ambrosia artemisiifol- ia*, Ambrosia sp., Aster sp., Bidens bi- pinnata, B. coronata, B. sp., Conyza can- adensis (as Erigeron canadensis), Dahlia sp., Erechtites hieraciifolia*, Erigeron sp., Eupatorium capillifolium, E. pilos- um, E. sp., Glycine max, Helianthus an- nuus, H. tuberosus*, H. sp., Quercus pal- ustris, Silphium sp., Solanum tuberosum, Solidago sp., Verbesina alternifolia (as Actinomeris alternifolia), Vitis rotundi- folia, V. sp. Notes: Deitz et al. (1976) re- ported an Entylia from Glycine max in Columbus Co., however, the voucher ma- 251 terial could not be located for the present study. Also, Brimley’s (1938) material from Hendersonville (Henderson County) and Willard (Pender County) could not be located and his specimen (NCDA) from “‘Spruce”’ is actually labelled ‘“‘Sun- burst’”” (Haywood County). Glossonotus acuminatus (Fabricius 1775). Counties: Duplin, Wake. Seasonal distri- bution: 6 May—14 June. Host associa- tions: Quercus falcata*. Glossonotus turriculatus (Emmons 1854). Counties: Buncombe-Yancey [as “Black Mountains,’ which is on the border of these counties (USNM)]. Seasonal distri- bution: 15—20 June. Host associations: no data for North Carolina. Notes: Hosts re- ported elsewhere include Crataegus and Quercus (Kopp and Yonke 1974). Glossonotus univittatus (Harris 1841). Fig. 5. Counties: Bladen, Burke, Mecklen- burg, Rockingham. Seasonal distribution: 16 May—4 August. Host associations: Quercus alba, Q. rubra. Hadrophallus borealis (Fairmaire 1846) [as Ceresa borealis Fairmaire in Brimley 1938; as Spissistilus borealis (Fairmaire) in Kopp and Yonke 1973b]. Counties: Buncombe, Stanly, Swain, Wake, Watau- ga. Seasonal distribution: 3 June—29 Au- gust. Host associations: no data for North Carolina. Notes: One female specimen from Waynesville, Haywood County, 14 Sept., probably H. borealis-was formerly misidentified as Ceresa constans (Walk- er) by Z.P. Metcalf (NCDA). Hosts re- ported elsewhere include species in sev- eral plant families (Kopp and Yonke 1973b): [Hadrophallus constans (Walker 1851), misidentification]. Brimley’s (1938) rec- ords of Ceresa constans (Walker) refer to H. borealis (Raleigh [Wake County], Waynesville [Haywood County]), Sticto- cephala militaris (Havelock), or S. brev- itylus (Newton); Kopp and Yonke’s (1973b: map) North Carolina record of this species (as S. constans) was probably based on Brimley’s publication. Heliria cornutula Ball 1925. NEW STATE RECORD. Counties: Bladen, Mecklen- burg, Randolph, Wake. Seasonal distri- bution: 5 July—15 November. Host asso- ciations: no data for North Carolina. Heliria cristata (Fairmaire 1846). Counties: [as ‘‘Eastern N.C.’’ on data label (NCSU)]. Seasonal distribution: mid July. Host associations: no data for North Carolina. Notes: Elsewhere, Kopp and Yonke (1974) reported Quercus macro- carpa Michx. Heliria gemma Ball 1925. NEW STATE RECORD. Counties: Graham. Seasonal distribution: 1 September. Host associa- tions: no data for North Carolina. Heliria gibberata Ball 1925. NEW STATE RECORD. Counties: Burke, Wake. Sea- sonal distribution: 23 May-—late June. Host associations: no data for North Car- olina. Notes: Elsewhere, Ball (1931) re- corded Celtis occidentalis L. as a host. [Heliria mexicana Stal 1869]. Records of H. mexicana in North Carolina (Ball 1931, Brimley 1938) were not verified by the present authors. Heliria molaris (Butler 1877). Counties: Wake, Watauga. Seasonal distribution: August. Host associations: no data for North Carolina. Notes: Elsewhere, Kopp and Yonke (1974) reported Quercus and Populus as hosts. Heliria scalaris (Fairmaire 1846). Counties: Wake. Seasonal distribution: 21 June. Host associations: no data for North Car- olina. Notes: Elsewhere, Kopp and Yon- ke (1974) reported Crateagus as a host. Microcentrus caryae (Fitch 1851). Fig. 1. Counties: Buncombe, Forsyth, Haywood, Martin, Pitt, Sampson, Wake. Seasonal distribution: [7 May, nymph], 25 June—4 December. Host associations: Carya illi- noinensis*, Carya sp. Microcentrus perditus (Amyot and Serville 1843). Counties: Moore, Stokes, Wake. Seasonal distribution: 20 May—20 Octo- ber. Host associations: Carya illinoinen- SiS. Micrutalis calva (Say 1830) [in part as Mi- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON crutalis calva var. illinoiensis (Goding 1893) in Brimley 1938]. Fig. 3. Counties: Alamance, Alexander, Alleghany, Avery, Bertie, Bladen, Brunswick, Buncombe, Cabarrus, Caswell, Chatham, Cherokee, Columbus, Cumberland, Dare, Davidson, Davie, Duplin, Durham, Forsyth, Gra- ham, Granville, Guilford, Haywood, Hoke, Jackson, Johnston, Lincoln, Meck- lenburg, Mitchell, Moore, New Hanover, Onslow, Person, Richmond, Rutherford, Scotland, Stanly, Surry, Swain, Wake, Warren, Wayne, Wilkes. Seasonal distri- bution: late April—2 November. Host as- sociations: Conyza canadensis, Erigeron annuus, Gleditsia triacanthos*, Glycine max, Robinia pseudoacacia, Salix nigra, Solidago sp., Vitis rotundifolia, V. sp. ‘French hybrid’. Notes: Brimley’s (1938) records from Tin City and Willard (Pen- der County) were not verified by the pre- sent authors. Micrutalis dorsalis (Fitch 1851). Counties: Alleghany, Burke, Caldwell, Haywood, Swain, Watauga, Yancey. Seasonal distri- bution: 11 July—21 August. Host associ- ations: no data for North Carolina. Micrutalis malleifera Fowler 1895. NEW STATE RECORD. Counties: Dare. Sea- sonal distribution: 14 June. Host associ- ations: Physalis sp. Notes: Mead (1986) reviewed information on this treehopper, the only known vector of pseudo-curly top virus (a minor disease of tomatoes). Ophiderma definita Woodruff 1919. NEW STATE RECORD. Counties: Alamance, Beaufort, Bladen, Buncombe, Cabarrus, Edgecombe, Franklin, Harnett, Mecklen- burg, Nash, Northampton, Pitt, Polk, Randolph, Wake, Warren, Watauga. Sea- sonal distribution: 23 April—16 July, [11 Sept. at UV-light (NCSU)]. Host associ- ations: Fagus grandifolia, Quercus fal- cata, Q. laurifolia, Q. nigra*, Q. palus- tris, Q. phellos*. Ophiderma evelyna Woodruff 1919. Fig. 9. Counties: Alamance, Bladen, Cabarrus, Greene, Harnett, Johnston, Lenoir, Meck- lenburg, Montgomery, Nash, Pitt, Ran- VOLUME 101, NUMBER 2 dolph, Wake, Washington, Wayne, Wil- son. Seasonal distribution: late March— 23-30 June. Host associations: Quercus alba, Q. falcata*, Q. marilandica*, Q. ni- gra, Q. palustris, Q. phellos*. Ophiderma flava Goding 1893. Counties: Ashe, Buncombe-Haywood [as “‘Mt. Pis- gah,’ which is on the border of these counties], Macon, Watauga. Seasonal dis- tribution: | June—31 August. Host asso- ciations: Quercus alba*, Q. rubra. Ophiderma flavicephala Goding 1893. Counties: Alamance, Beaufort, Bun- combe, Cabarrus, Durham, Franklin, Hoke, Lenoir, Nash, Pitt, Rutherford, Wake, Wayne, Wilkes, Wilson, Yadkin. Seasonal distribution: 23 April—18 June. Host associations: Quercus alba, Q. coc- cinea, Q. falcata*, Q. nigra*, Q. palus- tris, Q. phellos*, Q. rubra var. ambigua (as borealis), Q. stellata. Ophiderma grisea Woodruff 1919. Coun- ties: Buncombe, Yancey [as “Valley of Black Mountains,” which is on the Bun- combe—Yancey border (USNM)]. Sea- sonal distribution: 18 June—9 August. Host associations: no data for North Car- olina. Notes: Elsewhere, Kopp and Yon- ke (1973c) reported four species of Quer- cus as hosts. Ophiderma pubescens (Emmons 1854) [in part as Ophiderma flaviguttula Goding 1893, and as Ophiderma pubescens vat. australis Woodruff 1919, in Brimley 1938]. Counties: Ashe, Hoke, Jackson, Johnston, Moore, Scotland, Wake. Sea- sonal distribution: 24 April—26 July. Host associations: Quercus falcata*, Q. mari- landica, Q. stellata. Notes: Brimley’s (1938) record of O. flaviguttata [misiden- tification] from Raleigh [Wake County], May, was based on a specimen of Cyr- tolobus flavolatus (NCDA). Ophiderma salamandra Fairmaire 1846. Counties: Ashe, Burke, Greene, Harnett, Johnston, Lee, Nash, Pasquotank, Wake, Warren, Watauga, Wayne, Wilson. Sea- sonal distribution: 18 April—-31 August. Host associations: Quercus falcata, Q. ni- 253 gra, Q. palustris, Q. phellos*, O. rubra, Q. stellata, O. velutina*. Platycotis vittata (Fabricius 1803) [as Pla- tycotis vittata var. quadrivittata (Say 1830) in Brimley 1938]. Fig. 7. Counties: Alamance, Alexander, Alleghany, Anson, Ashe, Avery, Bladen, Buncombe, Burke, Cabarrus, Caswell, Catawba, Chatham, Cherokee, Chowan, Clay, Columbus, Craven, Cumberland, Dare, Durham, Forsyth, Franklin, Gaston, Graham, Granville, Guilford, Halifax, Haywood, Henderson, Hertford, Hoke, Iredell, Jack- son, Lee, Lenoir, Lincoln, Macon, Mar- tin, Mecklenburg, Montgomery, Moore, Nash, New Hanover, Northampton, Ons- low, Pasquotank, Pitt, Randolph, Robe- son, Rockingham, Sampson, Stanly, Stokes, Surry, Swain, Vance, Wake, War- ren, Watauga, Wilkes, Wilson, Yadkin, Yancey. Seasonal distribution: 16 Janu- ary—29 December. Host associations: Bet- ula sp.*, Castanea dentata*, Fagus sp.*, Quercus alba*, Q. falcata*, Q. incana*, Q. laevis, Q. margarettiae, Q. nigra*, Q. palustris*, Q. phellos, Q. rubra*, Q. rub- ra var. ambigua, Q. stellata*, Q. velutina, Q. virginiana*. Publilia concava (Say 1824). Fig. 10 (right). Counties: Ashe, Avery, Bun- combe, Burke, Cherokee, Graham, Hay- wood, Henderson, Jackson, Macon, Mad- ison, McDowell, Mitchell, Swain, Watau- ga, Wilkes, Yadkin. Seasonal distribu- tion: late May-—7 October. Host associations: Ambrosia artemisiifolia*, A. sp.*, Eupatorium sp.*, Helianthus sp.*, Solidago sp.* Notes: Brimley’s (1938) specimen from “‘Spruce”’ is actually la- belled ‘Sunburst’? (Haywood County: NCDA). Publilia reticulata Van Duzee 1908. Coun- ties: Ashe, Avery, Buncombe, Caldwell, Haywood, Henderson, Jackson, Transyl- vania, Wake, Watauga, Yancey (as ‘‘Black Mountains,” which is on the Buncombe—Yancey border (USNM)]. Seasonal distribution: 29 April—4 Octo- ber. host associations: no data for North 254 Carolina. Notes: Hosts reported else- where include members of the Asteracae (Kopp and Yonke 1973b). Smilia camelus (Fabricius 1803). Counties: Burke, Greene, Hoke, Mecklenburg, Moore, Pender, Robeson, Sampson, Wake, Watauga. Seasonal distribution: 24 April—25 September. Host associations: Quercus falcata*, Q. laevis*, Q. nigra*, Q. stellata. Notes: Brimley’s (1938) re- cord of this species from Linville Falls [Caldwell County] was not verified by the present authors. Smilia fasciata Amyot and Serville 1843. NEW STATE RECORD. Counties: Alamance, Beaufort, Cabarrus, Chatham, Craven, Davidson, Franklin, Gaston, Harnett, Johnston, Lenoir, Mecklenburg, Moore, Nash, Orange, Pasquotank, Pen- der, Rockingham, Wake, Warren, Wayne, Wilson. Seasonal distribution: 2 April—5 September. Host associations: Carya il- linoinensis, Quercus coccinea, Q. falca- ta*, O: nigra*, QO: palusiris*, QO. phellos*, Q. stellata*, Q. velutina. [Spissistilus femoratus (Fairmaire 1846)]. Records of this species [as Ceresa an- gulata (Walker 1851)] in North Carolina (Van Duzee 1917, Brimley 1938) were not verified by the present authors. Spissistilus festinus (Say 1830) [as Sticto- cephala festina (Say) in Brimley 1938]. Counties: Alexander, Anson, Beaufort, Bertie, Bladen, Brunswick, Buncombe, Cabarrus, Caldwell, Camden, Carteret, Catawba, Chatham, Chowan, Cleveland, Columbus, Craven, Cumberland, Curri- tuck, Dare, Davie, Duplin, Durham, Edge- combe, Franklin, Gaston, Gates, Gran- ville, Greene, Halifax, Harnett, Hay- wood, Henderson, Hoke, Iredell, Jackson, Johnston, Jones, Lee, Lincoln, Madison, Martin, Mecklenburg, Mitchell, Mont- gomery, Moore, Nash, New Hanover, Onslow, Orange, Pamlico, Pasquotank, Pender, Perquimans, Polk, Randolph, Robeson, Rockingham, Rowan, Samp- son, Scotland, Stanly, Swain, Transylva- nia, Tyrrell, Union, Vance, Wake, War- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ren, Washington, Wayne, Wilkes, Wilson. Seasonal distribution: 19 February—19 December. Host associations: Arachis hy- pogaea*, Aster ericoides, Glycine max*, Helianthus sp., Lespedeza sp.*, Medica- go sativa*, Phaseolus vulgaris*, Sarra- cenia flava, Solidago sp. Notes: This spe- cies is commonly known as the “‘three- cornered alfalfa hopper.’’ Hargrove’s (1986) record of S. festinus from Cow- eeta, Macon County (on Robinia pseu- doacacia), was not verified by the present authors. [Spissistilus rotundata Stal 1869]. Brim- ley’s (1938) records of Stictocephala ro- tundata Stal in North Carolina (Raleigh [Wake County] and Southern Pines [Moore County]) were not verified by the present authors. [Spissistilus uniformis (Fairmaire 1846), misidentification]. Brimley’s (1938) rec- ords of Ceresa uniformis Fairmaire ac- tually refer to Hadrophallus borealis; these include one female in NCDA from Balsam [Jackson County], 14-18 Sep- tember. Stictocephala bisonia Kopp and Yonke 1977 [as Ceresa bubalus (Fabricius 1794), misidentification, in Brimley 1938; as Stictocephala bubalus (Fabri- cius), misidentification, in Kopp and Yonke 1973b; Stictocephala bisonia Kopp and Yonke 1977 (Kopp and Yonke 1977); as S. alta (Walker 1851) in An- drade 1997 (questionable synonymy, see ‘Notes,’ below)]. Counties: Alleghany, Ashe, Avery, Beaufort, Buncombe, Hay- wood, Madison, Mitchell, Montgomery, Stanly, Surry, Wake, Warren, Watauga. Seasonal distribution: 16 July—30 Sep- tember. Host associations: Glycine max. Notes: This species is commonly known as the “buffalo treehopper.”’ Recently, Andrade (1997) considered S. bisonia to be a junior synonym of S. alta (Walker 1851). We believe this synonymy is in- sufficiently justified in light of the need to examine the male genitalia to reliably identify many species of the tribe Cere- VOLUME 101, NUMBER 2 sini. The holotype of Ceresa alta Walker is of ambiguous identity, being a female specimen from an unspecified locality. On the other hand, the identity of the male holotype of S. bisonia Kopp and Yonke is definitive, and this type is ac- companied by 45 paratypes (males and females), all from Columbia, Missouri, USA. [Stictocephala brevicornis (Fitch 1856)]. Records of this species in North Carolina were not verified by the present authors. Brimley’s (1938) record of Ceresa brev- icornis Fitch (Swannanoa, Buncombe County, May) probably referred to Stic- tocephala brevitylus (Van Duzee); Kopp and Yonke’s (1973b: map) record may be based on Brimley’s publication. Hargrove (1986) also listed this treehopper from Coweeta, Macon County, on Robinia pseudoacacia. [Stictocephala brevis (Walker 1851)]. Notes: Brimley’s (1938) record of this species in North Carolina (as Ceresa brevis Walker) has not been verified by the present authors. One specimen (Swannanoa, Buncombe County, 21 May: NCDA) identified as Ceresa brevis Walker by Brimley is actually Stictoceph- ala brevitylus, however, Brimley’s pub- lished record gave no locality or date. Stictocephala brevitylus (Van Duzee 1908) [as Ceresa brevitylus Van Duzee in Brim- ley 1938]. Counties: Alleghany, Anson, Avery, Brunswick, Buncombe, Cabarrus, Caswell, Catawba, Chatham, Cumber- land, Currituck, Dare, Durham, Forsyth, Granville, Harnett, Haywood, Henderson, Hertford, Jackson, McDowell, Mecklen- burg, Moore, New Hanover, Pasquotank, Randolph, Stanly, Swain, Wake, Wash- ington, Watauga, Wilkes, Yadkin, Yan- cey. Seasonal distribution: 5 April—2 July. Host associations: Aster sp., Cea- nothus sp., Chrysanthemum leucanthe- mum, Helianthus sp.*, Morus sp., Quer- cus falcata, Robinia pseudoacacia, Ru- bus argutus*, Sarracenia flava, Smilax sp.*, Solanum tuberosum, Vaccinium sp. 255 (as Polycodium sp.), Vitis sp. ‘French hy- brid’. Stictocephala diceros (Say 1824) [as Cer- esa diceros (Say) in Brimley 1938]. Counties: Ashe, Buncombe, Cabarrus, Caldwell, Durham, Granville, Haywood, Iredell, McDowell, Mecklenburg, Wake, Watauga. Seasonal distribution: 8 June—6 October. Host associations: Sambucus canadensis. [Stictocephala diminuta Van Duzee 1908]. North Carolina records of this species were not verified by the present authors. Brimley’s (1938) record of S. diminuta in Raleigh [Wake County], July, probably represents S. brevitylus. Stictocephala lutea (Walker 1851). Coun- ties: Alleghany, Avery, Bladen, Bruns- wick, Buncombe, Burke, Carteret, Cas- well, Chatham, Cumberland, Duplin, Haywood, Hoke, Jackson, Johnston, Madison, Mecklenburg, Moore, New Hanover, Onslow, Pender, Sampson, Scotland, Stanly, Transylvania, Vance, Wake, Washington, Watauga. Seasonal distribution: 24 February—16 September. Host associations: Quercus falcata (suck- ers). Stictocephala militaris (Gibson and Wells 1917). NEW STATE RECORD. Fig. 6. Counties: Bladen, Craven, Hyde, John- ston, Mecklenburg, Pitt, Wake. Seasonal distribution: 19 June—16 October. Host associations: Cercis canadensis, Prunus serotina. Stictocephala palmeri (Van Duzee 1908) [as Ceresa palmeri Van Duzee in Brim- ley 1938]. Counties: Avery-Caldwell- Watauga [as ‘Grandfather Mountain,”’ which is on the border of these three counties (NCSU)], Buncombe, Graham, Macon, McDowell-Yancey [as ‘“‘Buck Creek Gap,” which is on the border of these counties (NCSU)], Stanly, Wake. Seasonal distribution: 21 July—29 Sep- tember. Host associations: no data for North Carolina. Notes: Elsewhere, Carya is the oviposition host (Funkhouser 256 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1917)—feeding occurs also on other woody hosts (Kopp and Yonke 1973b). Stictocephala stimulea (Van Duzee 1914). NEW STATE RECORD. Counties: Ca- barrus, Pender, Wake. Seasonal distribu- tion: 1-31 May. Host associations: Vitis prob. vulpina [ovipositing in canes]. [Stictocephala substriata (Walker 1851)]. Brimley’s (1938) records of this species in North Carolina (Raleigh [Wake Coun- ty] and Southern Pines [Moore County]) were not verified by the present authors. Stictocephala taurina (Fitch 1856) [as Cer- esa taurina (Fitch) in Brimley 1938]. Counties: Ashe, Avery, Buncombe, For- syth, Madison, Mitchell, Surry, Wake, Watauga. Seasonal distribution: 19—20 June—18 September. Host associations: Helianthus sp., Rubus sp., Sambucus can- adensis, Smilax sp.* Stictocephala tauriniformis Caldwell 1949. NEW STATE RECORD. Counties: Hay- wood. Seasonal distribution: 14 Septem- ber. Host associations: no data for North Carolina. Notes: Elsewhere, Kopp and Yonke (1973b) reported Quercus and Til- ia as hosts. Stictolobus minutus (Funkhouser 1915) [as Stictolobus subulatus (Say 1830) in Brimley 1938 and Kopp and Yonke 1973b]. Counties: Pender, Wake. Season- al distribution: early July. Host associa- tions: no data for North Carolina. Notes: Elsewhere, Kopp and Yonke (1973b) re- ported Taxodium distichum (L.) L. Ri- chard as a host. Telamona ampelopsidis (Harris 1841). Counties: unknown, as “‘NC”’ on data la- bel (USNM). Seasonal distribution: un- known. Host associations: no data for North Carolina. Notes: Early reports of this species in North Carolina (Ball 1931; Brimley 1938) apparently refer to 7. ma- culata Van Duzee, based on a specimen misidentified by C. S. Brimley as T. am- pelopsidis (NCDA). Kopp and Yonke (1974: map), who also reported the spe- cies in North Carolina, listed its host as Parthenocissus quinquefolia (L.) Plan- chon. Telamona collina (Walker 1851). NEW STATE RECORD. Counties: Wake. Sea- sonal distribution: 18—22 May. Host as- sociations: Platanus occidentalis. [Telamona concava Fitch 1851, misidenti- fication]. Records of Telamona concava from Linville Falls (Caldwell County, June 1920: NCDA) (Brimley 1942, Wray 1967), actually refer to Heliria gibberata. Telamona decorata Ball 1903. NEW STATE RECORD. Counties: Ashe, Hen- derson, Watauga. Seasonal distribution: 19-20 June—13 September. Host associ- ations: Castanea dentata, Quercus alba, Q. rubra. Telamona dubiosa Van Duzee 1916. NEW STATE RECORD. Counties: Bladen. Seasonal distribution: 21 September—1 1 October. Host associations: no data for North Carolina. Funkhouser (1917) listed Quercus alba as a host. Telamona extrema Ball 1903. NEW STATE RECORD. Counties: Carteret, Wake, Yadkin. Seasonal distribution: 13—29 May. Host associations: no data for North Carolina. Notes: Hosts reported else- where include two species of Quercus (Kopp and Yonke 1974). Telamona maculata Van Duzee 1908. NEW STATE RECORD. Counties: Ashe, Wake. Seasonal distribution: 10 May—18 July. Host associations: no data for North Carolina. Notes: Hosts elsewhere include three species of Quercus (Kopp and Yon- ke 1974). Telamona monticola (Fabricius 1803). Counties: Ashe, Avery, Buncombe, Burke, Caldwell, Columbus, Hoke, John- ston, Macon, Robeson, Surry, Vance, Wake, Watauga. Seasonal distribution: 20 April—18 October. Host associations: Quercus falcata, Q. margarettiae, Q. ni- gra*, Q. rubra*, Q. stellata, Vitis rotun- difolia. Telamona reclivata Fitch 1851. Counties: Ashe, Buncombe, Macon, Mecklenburg, VOLUME 101, NUMBER 2 Wake. Seasonal distribution: 2 June—31 August. Host associations: Quercus alba. [Telamona salvini Distant 1879]. Records (Ball 1931, Brimley 1938) of T. salvini in North Carolina (Black Mountains) were not verified by the present authors, but the species is known to occur in Charleston County, South Carolina (NCSU, 1 specimen). [Telamona tiliae Ball 1925]. Records of T. tiliae in ‘“‘North Carolina’ (Ball 1931, Brimley 1938) seem to be based, at least in part, on a specimen (Blowing Rock [Watauga County], 13 Sept., NCDA) identified by C. S. Brimley as T. tiliae which is actually 7. decorata. Presence of this species in North Carolina remains unconfirmed. Telamona unicolor Fitch 1851. Counties: Bertie, Bladen, Buncombe—Yancey [as ‘““Black Mountains” which is on the Bun- combe—Yancey border], Columbus, Hoke, Macon, Mecklenburg, Wake. Sea- sonal distribution: 29 April—27 October. Host associations: Carya illinoinensis*, Caspe: Telamona westcotti Goding 1893. Counties: Henderson, Union. Seasonal distribution: 25-30 June—S September. Host associa- tions: no data for North Carolina. Notes: Hosts reported elsewhere include Quer- cus, Tilia, and Ulmus (Kopp and Yonke 1974). Thelia bimaculata (Fabricius 1794). Coun- ties: Alexander, Ashe, Avery, Buncombe, Graham, Haywood, Jackson, Lincoln, Macon, McDowell, Mecklenburg, Swain, Wake, Watauga. Seasonal distribution: 10 June—19 October. Host associations: Ro- binia pseudoacacia*. Tortistilus abnormus (Caldwell 1949). NEW STATE RECORD. Counties: Dur- ham, Stanly. Seasonal distribution: 13—21 July. Host associations: no data for North Carolina. [Tortistilus inermis (Fabricius 1775)]. Brimley’s (1938) record of 7. inermis (as Stictocephala inermis) in North Carolina was not verified by the present authors, D5, but the species has also been recorded from two adjoining states, Virginia and Tennessee (Kopp and Yonke 1973b). Tortistilus lateralis (Funkhouser 1936). NEW STATE RECORD. Counties: Blad- en. Seasonal distribution: 8-22 Septem- ber. Host associations: no data for North Carolina. Tylopelta gibbera (Stal 1869) [as Tylopelta brevis Van Duzee 1908, in Brimley 1938; as Tylopelta americana (Goding 1893) in Kopp and Yonke 1973a]. Counties: Bun- combe, Haywood, New Hanover, Swain, Wake. Seasonal distribution: 16 April—27 September. Host associations: Desmo- dium sp. Vanduzea arquata (Say 1830) [as V. arcu- ata (sic) in Brimley 1938]. Fig. 8. Coun- ties: Alamance, Alexander, Alleghany, Ashe, Avery, Bladen, Buncombe, Cald- well, Caswell, Catawba, Chatham, Cher- okee, Clay, Durham, Forsyth, Graham, Haywood, Jackson, Lincoln, Macon, Madison, Mecklenburg, Mitchell, Orange, Polk, Richmond, Rockingham, Sampson, Surry, Swain, Union, Wake, Watauga, Wilkes, Yancey. Seasonal dis- tribution: [? late March (NCSU)], 9 May-—27 October. Host associations: Ro- binia pseudoacacia*. Vanduzea triguttata (Burmeister 1836). Counties: Hoke, Johnston, Lee, Moore, New Hanover, Pender, Stanly, Wake. Sea- sonal distribution: 17 June—27 Septem- ber. Host associations: Lespedeza sp.* Xantholobus intermedius (Emmons 1854). Counties: Ashe, Buncombe—Yancey [as “Valley of Black Mountains,” which is on the Buncombe—Yancey border (USNM)], Wake. Seasonal distribution: 4 May—7 July. Host associations: Betula alleghan- iensis (as B. lutea), Quercus phellos*. Xantholobus lateralis (Van Duzee 1908) [as Cyrtolobus lateralis Van Duzee in Brim- ley 1938]. Counties: Ashe, Burke, Cald- well, Lee, Mecklenburg, Watauga. Sea- sonal distribution: 3 May—6 July. Host associations: Betula sp., Quercus alba. Xantholobus muticus (Fabricius 1777) [as 258 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cyrtolobus muticus (Fabricius) in Brim- ley 1938]. Counties: Camden, Chatham, Duplin, Forsyth, Franklin, Gates, Greene, Hertford, Lee, Moore, Nash, Northamp- ton, Randolph, Rutherford, Wake, War- ren, Wayne, Wilson. Seasonal distribu- tion: 18 April—14 July. Host associations: Quercus alba*, Q. laevis, Q. prinus, Q. stellata*. Xantholobus nitidus (Van Duzee 1908). Counties: Bladen, Forsyth, Johnston, Moore, Sampson, Wake, Wilkes. Season- al distribution: 17 May—19 June. Host as- sociations: no data for North Carolina. Host PLANT INDEX FOR NORTH CAROLINA TREEHOPPERS Plant scientific name, common name(s) (Family): associated treehopper species. Ambrosia artemisiifolia L., ragweed (Aster- aceae): Acutalis tartarea, Entylia cari- nata*, Publilia concava*. Ambrosia sp., ragweed (Asteraceae): Enty- lia carinata, Publilia concava*. Arachis hypogaea L., peanut, common pea- nut, goober, groundnut, grass nut, earth nut, monkey nut, pindar (Fabaceae): Spis- sistilus festinus*. Arundinaria sp., bamboo, cane (Poaceae): Acutalis tartarea. Aster ericoides L., heath aster (Asteraceae): Spissistilus festinus. Aster sp., aster, Michaelmas daisy, starwort, frost flower (Asteraceae): Entylia cari- nata, Stictocephala brevitylus. Betula alleghaniensis Britton, yellow birch, gray birch (Betulaceae): Xantholobus in- termedius. Betula sp., birch (Betulaceae): Carynota marmorata, Platycotis vittata*, Xantho- lobus lateralis. Bidens bipinnata L., Spanish needles (As- teraceae): Entylia carinata. Bidens coronata (L.) Britton, beggar ticks, beggar’s ticks (Asteraceae): Acutalis tar- tarea, Entylia carinata. Bidens sp., beggar ticks, beggar’s ticks, bur marigold, water marigold, pitchforks, Spanish needles, stick-tights, tickseed (Asteraceae): Entylia carinata. Carya illinoinensis (Wangenh.) K. Koch, pecan (Juglandaceae): Carynota mera, Microcentrus caryae*, M. perditus, Smi- lia fasciata, Telamona unicolor*. Carya sp., hickory (Juglandaceae): Cary- nota mera*, Enchenopa binotata com- plex, Microcentrus caryae, Telamona unicolor*. Castanea dentata (Marshall) Borkh., Amer- ican chestnut (Fagaceae): Atymna casta- neae*, Platycotis vittata*, Telamona de- corata. Castanea pumila (L.) P. Miller, chinquapin (Fagaceae): Atymna castaneae*. Ceanothus sp., redroot (Rhamnaceae): Stic- tocephala brevitylus. Cercis canadensis L., eastern redbud, red- bud, Judas tree (Fabaceae): Enchenopa binotata complex*, Stictocephala mili- taris. Chrysanthemum leucanthemum L., ox-eye daisy, white daisy, marguerite, white- weed (Asteraceae): Stictocephala brevi- tylus. Conyza canadensis (L.) Cronquist, horse- weed, hogweed, butterweed (Asteraceae): Entylia carinata, Micrutalis calva. Dahlia sp., dahlia (Asteraceae): Entylia carinata. Desmodium sp., beggar lice, beggar ticks, beggar’s ticks, tick trefoil, tick clover (Fabaceae): Tylopelta gibbera. Erechtites hieraciifolia (L.) Raf. ex DC., fireweed (Ranuculaceae): Entylia cari- nata*. Erigeron annuus (L.) Persoon, daisy flea- bane, sweet scabious, white-top, fleabane (Asteraceae): Micrutalis calva. Erigeron sp., fleabane (Asteraceae): Entylia carinata. Eupatorium capillifolium (Lam.) Small, dog-fennel (Asteraceae): Acutalis tarta- rea, Entylia carinata. Eupatorium pilosum Walter, thoroughwort, boneset (Asteraceae): Entylia carinata. Eupatorium sp., thoroughwort, boneset a VOLUME 101, NUMBER 2 (Asteraceae): Entylia carinata, Publilia concava*. Fagus grandifolia J. FE Ehrhart, beech, American beech (Fagaceae): Ophiderma definita. Fagus sp., beech (Fagaceae): Microcentrus caryae, Platycotis vittata. Gleditsia triacanthos L., honey locust, sweet locust, honeyshuck (Fabaceae): Micrutalis calva*. Glycine max (L.) Merrill, soybean, soja bean, soya bean (Fabaceae): Entylia car- inata, Micrutalis calva, Spissistilus fes- tinus*, Stictocephala alta. Helianthus annuus L., sunflower, common sunflower, mirasol (Asteraceae): Entylia carinata. Helianthus sp., sunflower (Asteraceae): Acutalis tartarea, Entylia carinata, Pub- lilia concava, Spissistilus festinus, Stic- tocephala brevitylus*, S. taurina. Helianthus tuberosus L., Jerusalem arti- choke, girasole (Asteraceae): Entylia car- inata*. Juglans nigra L., black walnut (Juglanda- ceae): Enchenopa binotata complex*. Lespedeza sp., bush clover (Fabaceae): Spissistilus festinus*, Vanduzea trigutta- fa. Liriodendron tulipifera L., tulip tree, tulip poplar, whitewood, yellow poplar (Mag- noliaceae): Enchenopa binotata complex. Medicago sativa L., alfalfa, lucerne (Faba- ceae): Campylenchia latipes, Spissistilus festinus*. Morus sp., mulberry (Moraceae): Stictoce- phala brevitylus. Phaseolus vulgaris L., common bean, kid- ney bean, green bean, snap bean, haricot, French bean, frijol, runner bean, string bean, salad bean, wax bean (Fabaceae): Spissistilus festinus*. Physalis sp., ground cherry (Solanaceae): Micrutalis malleifera. Platanus occidentalis L., eastern sycamore, sycamore, button wood, buttonball, American plane-(Platanaceae): Telamona collina. Prunus serotina J. FE Ehrhart, black cherry, 259 wild black cherry, rum cherry, (Rosa- ceae): Stictocephala militaris. Quercus alba L., white oak (Fagaceae): Ar- chasia auriculata*, A. belfragei*, Atym- na querci*, Cyrtolobus auroreus*, C. dixianus, C. fuscipennis*, Cyrtolobus gri- seus, C. maculifrontis*, C. pallidifrontis, C. tuberosus*, C. vau*, Glossonotus uni- vittatus, Ophiderma evelyna, O. flava*, O. flavicephala, Platycotis vittata*, Te- lamona decorata, T. reclivata, Xantho- lobus lateralis, X. muticus*. Quercus coccinea Muenchh., scarlet oak (Fagaceae): Cyrtolobus fuliginosus, Ophiderma flavicephala, Smilia fasciata. Quercus falcata Michaux, southern red oak, Spanish oak, Spanish red oak (Fagaceae): Stictocephala lutea, Telamona monticola, Cyrtolobus arcuatus*, C. dixianus, C. fe- nestratus*, C. fuliginosus*, C. inermis*, C. maculifrontis, Glossonotus acumina- tus*, Ophiderma definita, O. evelyna*, O. flavicephala*, O. pubescens*, O. sala- mandra, Platycotis vittata*, Smilia ca- melus*, S. fasciata*, Stictocephala brev- itylus, S. lutea, Telamona monticola. Quercus incana Bartram, bluejack oak, up- land willow oak, bluejack, turkey oak, high-ground willow oak, sand jack (Fa- gaceae): Platycotis vittata*. Quercus laevis Walter, turkey oak, Catesby oak (Fagaceae): Cyrtolobus fenestratus, C. ovatus*, Platycotis vittata, Smilia ca- melus*, Xantholobus muticus. Quercus laurifolia Michaux, laurel oak, Darlington oak, laurel-leaved oak (Faga- ceae): Ophiderma definita. Quercus margarettiae Ashe ex Small, scrubby post oak (Fagaceae): Cyrtolobus fenestratus, C. tuberosus, Telamona mon- ticola, Platycotis vittata. Quercus marilandica Muenchh., blackjack oak, blackjack, jack oak (Fagaceae): Cyr- tolobus fenestratus*, C. inermis, C. ova- tus*, C. tuberosus, Ophiderma evelyna*, O. pubescens. Quercus nigra L., water oak, possum oak (Fagaceae): Archasia auriculata, Cyrtolo- bus fenestratus*, C. fuscipennis, C. toga- 260 tus*, C. tuberosus, Ophiderma definita*, O. evelyna, O. flavicephala*, O. salaman- dra, Platycotis vittata*, Smilia camelus*, S. fasciata*, Telamona monticola*. Quercus palustris Muenchh., pin oak, Spanish oak (Fagaceae): Cyrtolobus cla- rus*, C. dixianus, C. fenestratus, C. fu- liginosus, C. funkhouseri, Entylia cari- nata, Ophiderma definita, O. evelyna, O. flavicephala, O. salamandra, Platycotis vittata, Smilia fasciata*. Quercus phellos L., willow oak (Fagaceae): Archasia belfragei, Cyrtolobus arcuatus, C. clarus*, C. fenestratus*, C. fuligino- sus, C. togatus*, Ophiderma definita*, O. evelyna*, O. flavicephala*, O. salaman- dra*, Platycotis vittata, Smilia camelus, S. fasciata*, Xantholobus intermedius*. Quercus prinus L., chestnut oak, rock chestnut oak, basket oak (Fagaceae): Cyr- tolobus auroreus, C. tuberosus, C. vau, Xantholobus muticus. Quercus rubra L. (see also var. ambigua, gray oak or northern red oak, below), red oak (Fagaceae): Cyrtolobus fenestratus*, C. pulchellus, C. togatus, Glossonotus univittatus, Ophiderma flava, O. sala- mandra, Platycotis vittata*, Telamona decorata, T. monticola. Quercus rubra var. ambigua (Gray) Fern., gray oak, northern red oak (Fagaceae): Ophiderma flavicephala, Platycotis vit- tata. Quercus stellata Wangenh., post oak (Fa- gaceae): Atymna querci*, Cyrtolobus dix- tanus=, Gs juliginosuss «G.ioriseuss «C. inermis, C. pallidifrontis*, C. togatus, C. tuberosus, C. vau, Ophiderma flavicepha- la, O. pubescens, O. salamandra, Platy- cotis vittata*, Smilia camelus, S. fascia- ta*, Telamona monticola, Xantholobus muticus*. Quercus velutina Lam., black oak, yellow- bark oak, quercitron (Fagaceae): Archas- ia auriculata*, Cyrtolobus fenestratus, C. fuliginosus, Ophiderma salamandra*, Platycotis vittata, Smilia fasciata. Quercus virginiana P. Miller, live oak, southern live oak (Fagaceae): Cyrtolobus PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fenestratus, C. tuberosus, Platycotis vit- tata*. Robinia pseudoacacia L., black locust, false acacia, yellow locust (Fabaceae): Enchenopa binotata complex*, Micrutal- is calva, Stictocephala brevitylus, Thelia bimaculata*, Vanduzea arquata*. Rubus argutus Link, blackberry (Rosa- ceae): Stictocephala brevitylus*. Rubus sp., bramble (Rosaceae): Stictoceph- ala taurina. Salix nigra Marshall, black willow (Sali- caceae): Micrutalis calva. Sambucus canadensis L., elderberry, Amer- ican elderberry, sweet elderberry, (Capri- foliaceae): Stictocephala diceros, S. taur- ina. Sarracenia flava L., yellow pitcher plant, trumpets, watches, biscuit-flower, trum- petleaf, umbrella-trumpets, huntsman’s horn (Sarraceniaceae): Spissistilus festin- us, Stictocephala brevitylus. Silphium sp., rosinweed (Asteraceae): En- tylia carinata. Smilax sp., greenbrier, catbrier (Liliaceae): Stictocephala brevitylus*, S. taurina*. Solanum tuberosum L., potato, Irish potato, white potato (Solanaceae): Entylia cari- nata, Stictocephala brevitylus. Solidago sp., goldenrod (Asteraceae): Acu- talis tartarea, Campylenchia latipes, En- tylia carinata, Micrutalis calva, Publilia concava*, Spissistilus festinus. Vaccinium sp., blueberry, huckleberry, cranberry, bilberry (Ericaceae): Sticto- cephala brevitylus. Verbesina alternifolia (L.) Britton ex Kear- ney, wingstem, yellow ironweed (Aster- aceae): Entylia carinata. Viburnum prunifolium L., black haw, sweet haw, sheepberry, nanny-berry, stagbush (Caprifoliaceae): Enchenopa binotata complex. Vitis rotundifolia Michaux, muscadine grape, scuppernong grape, bullace grape (Vitaceae): Atymna querci, Entylia cari- nata, Micrutalis calva, Telamona monti- cola. Vitis sp., grape (Vitaceae): Entylia carinata. VOLUME 101, NUMBER 2 Vitis sp. “French hybrid,’ grape ‘French hy- brid’ (Vitaceae): Micrutalis calva, Stic- tocephala brevitylus. Vitis prob. vulpina L., frost grape, winter grape, chicken grape (Vitaceae): Sticto- cephala stimulea. ACKNOWLEDGMENTS The authors are grateful to the following individuals: for collecting treehoppers listed in this work—J. R. Baker, C. H. Bartlett, R. L. Blinn, B. K. Cassel, P. J. Crawford, J. R. Cryan, B. A. Harrison, H. H. Neunzig, W. L. Nicholson, R. L. Regnery, D. L. Ste- phanss. J. Toth, Jr, J. Dp. Trexler, M. S. Wallace, and B. M. Wiegmann; for prepar- ing many specimens for the North Carolina State University Insect Collection—M. P. Sobanski; for allowing us to examine spec- imens in their care—S. E. Halbert and F W. Mead (FSCA); K. R. Ahlstrom (NCDA); R. L. Blinn (NCSU); and T. J. Henry, S. H. McKamey, Systematic Entomology Labo- ratory, USDA, and R. C. Froeschner, Smithsonian Institution (USNM); for assis- tance with plant identifications and botani- cal nomenclature—J. W. Floyd and J. W. Hardin (Department of Botany, North Car- olina State University); and for helpful comments on manuscript—R. L. Blinn, J. RoGryan Ww. + Gimpel,, Jr, (S: i. Me- Kamey, H. H. Neunzig, D. J. Voegtlin, B. M. Wiegmann, and D. W. Webb. Y. W. Chan and S. J. Toth, Jr., assisted in the prep- aration of Fig. 11; photographs are by C. H. Dietrich, except Fig. 2, which is by R. L. Blinn. This research was supported by the North Carolina Agricultural Research Service and NSF grant DEB-9726282. LITERATURE CITED Andrade, G. S. de. 1997. Stictocephala alta (Walker 1851) sp. rev., comb. n., the correct name for the “buffalo treehopper,” with S. bisonia Kopp & Yonke, 1977 as a new synonym, and notes on Hadrophallus bubalus (Fabricius 1794) comb. n. (Homoptera: Membracidae). Transactions of the American Entomological Society 123: 289-295. Ball, E. D. 1931. A monographic revision of the tree- 261 hoppers of the tribe Telamonini of North America. Entomologica Americana 12: !—69. Brimley, C. S. 1938. The insects of North Carolina, being a checklist of the insects of North Carolina and their close relatives. Division of Entomology, North Carolina Department of Agriculture, Ra- leigh, North Carolina. 560 pp. Brimley, C. S. 1942. Supplement to insects of North Carolina. Division of Entomology, North Carolina Department of Agriculture, Raleigh, North Caro- lina. 39 pp. Deitz, L. L. 1975. Classification of the higher catego- ries of the New World treehoppers (Homoptera: Membracidae). North Carolina Agricultural Ex- periment Station Technical Bulletin 225: [i-iv], 1— Nae Deitz, L. L. 1989. Bibliography of the Membracoidea (Homoptera: Aetalionidae, Biturritiidae, Membra- cidae, and Nicomiidae) 1981-1987. North Caro- lina Agricultural Research Service Technical Bul- letin 290: 1-31. Deitz, L. L. and C. H. Dietrich. 1993. Superfamily Membracoidea (Homoptera: Auchenorrhyncha). I. Introduction and revised classification with new family-group taxa. Systematic Entomology 18: 287-296. Deitz, L. L. and D. D. Kopp. 1987. Bibliography of the Membracoidea (Homoptera: Aetalionidae, Bi- turritiidae, Membracidae, and Nicomiidae) 1956— 1980. North Carolina Agricultural Research Ser- vice Technical Bulletin 284: [i-ii], 1-39. Deitz ES J. WwW. Van Duyn. Jt R: Bradley: Jr RoE Rabb, W. M. Brooks, and R. E. Stinner. 1976. A guide to the identification and biology of soybean arthropods in North Carolina. North Carolina Ag- ricultural Experiment Station Technical Bulletin 238: [i-vi], 1-264. Funkhouser, W. D. 1917. Biology of the Membracidae of the Cayuga Lake Basin. Cornell University Ag- ricultural Experiment Station Memoir 11: 177— 445. Funkhouser, W. D. 1927. Membracidae. General Cat- alogue of the Hemiptera. Fascicle 1. Smith Col- lege, Northampton, Massachusetts. 581 pp. Goding, E W. 1893. Bibliographical and synonymical catalogue of the described Membracidae of North America. Bulletin of the Illinois State Laboratory of Natural History 3: 391—482. Hargrove, W. W. 1986. An annotated species list of insect herbivores commonly associated with black locust, Robinia pseudoacacia, in the southern Ap- palachians. Entomological News 97: 36—40. Kartesz, J. T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Vol 2. Thesaurus. The biota of North America. 2nd edition. The University of North Carolina Press, Chapel Hill, North Carolina. vii + 816 pp. 262 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Kopp, D. D. and T. R. Yonke. 1973a. The treehoppers of Missouri: Part 1. Subfamilies Centrotinae, Ho- plophorioninae, and Membracinae (Homoptera: Membracidae). Journal of the Kansas Entomolog- ical Society 46: 42—64. Kopp, D. D. and T. R. Yonke. 1973b. The treehoppers of Missouri: Part 2. Subfamily Smiliinae; tribes Acutalini, Ceresini, and Polyglyptini (Homoptera: Membracidae). Journal of the Kansas Entomolog- ical Society 46: 233-276. Kopp, D. D. and T. R. Yonke. 1973c. The treehoppers of Missouri: Part 3. Subfamily Smiliinae; tribe Smiliini. Journal of the Kansas Entomological So- ciety 46: 375-421. Kopp, D. D. and T. R. Yonke. 1974. The treehoppers of Missouri: Part 4. Subfamily Smiliinae; tribe Te- lamonini (Homoptera: Membracidae). Journal of the Kansas Entomological Society 47: 80—130. Kopp, D. D. and T. R. Yonke. 1977. Taxonomic status of the buffalo treehopper and the name Ceresa bubalus. Annals of the Entomological Society of America 70: 901—905. Kopp, D. D. and T. R. Yonke. 1979. A taxonomic re- view of the tribe Ceresini (Homoptera: Membra- cidae). Miscellaneous Publications of the Ento- mological Society of America 11(2): 1-97. Leonard, M. D. 1928. Membracidae, pp. 147—156. In Leonard, M. D., ed., A list of 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. Liberty Hyde Bailey Hortorium. 1976. Hortus Third. A concise dictionary of plants cultivated in the United States and Canada. Macmillian, New York. xiv + 1290 pp. McGiffen, K. C. and H. H. Neunzig. 1985. A guide to the identification and biology of insects feeding on muscadine and bunch grapes in North Caroli- na. North Carolina Agricultural Research Service Bulletin 470: [i-iv], 1-93. McKamey, S. H. 1998. Taxonomic catalogue of the Membracoidea (exclusive of leafhoppers): second supplement to Fascicle I—Membracidae of the General Catalogue of the Hemiptera. Memoirs of the American Entomological Institute 60: 1—377. Mead, F. W. 1986. Micrutalis treehoppers and pseudo- curly top in Florida (Homoptera: Membracidae). Florida Department of Agriculture & Consumer Services, Division of Plant Industry, Entomology Circular 283: [i-iv]. Metcalf, Z. P. 1915. A list of the Homoptera of North Carolina. Journal of the Elisha Mitchell Scientific Society 31: 35-60. Metcalf, Z. P. and V. Wade. 1963. A Bibliography of the Membracoidea and Fossil Homoptera (Ho- moptera: Auchennorhyncha [sic]). North Carolina State [University], Raleigh. iv + 200 pp. Metcalf, Z. P. and V. Wade. 1965. General Catalogue of the Homoptera. A Supplement to Fascicle I— Membracidae of the General Catalogue of the He- miptera. Membracoidea. In Two Sections. North Carolina State University, Raleigh. 1552 pp. Osborn, H. 1940. The Membracidae of Ohio. Ohio Bi- ological Survey Bulletin 7: 51-101. Pratt, G. and T. K. Wood. 1992. A phylogenetic anal- ysis of the Enchenopa binotata species complex (Homoptera: Membracidae) using nymphal char- acters. Systematic Entomolology 17: 351—357. Pratt, G. and T. K. Wood. 1993. Genitalic analysis of males and females in the Enchenopa_ binotata (Say) complex (Membracidae: Homoptera). Pro- ceedings of the Entomological Society of Wash- ington 95: 574-582. Radford, A. E., H. E. Ahles, and C. R. Bell. 1968. Manual of the vascular flora of the Carolinas. Uni- versity of North Carolina Press, Chapel Hill, North Carolina. Ixi + 1183 pp. Remes-Lenicov, A. M. M. de. 1973. Nota sinonimica sobre Entylia carinata (Forster) (Homoptera: Membracidae). Revista de la Sociedad Entomo- logica Argentina 34: 89-94. Van Duzee, E. P. 1917. Catalogue of the Hemiptera of America north of Mexico excepting the Aphidi- dae, Coccidae and Aleurodidae [sic]. California Agricultural Experiment Station Entomological Technical Bulletin 2: [i—xiv], 1—902. Wray, D. L. 1950. Insects of North Carolina. Second supplement. Division of Entomology, North Car- olina Department of Agriculture, Raleigh, North Carolina. 59 pp. Wray, D. L. 1967. Insects of North Carolina. Third supplement. Division of Entomology, North Car- olina Department of Agriculture, Raleigh, North Carolina. 181 pp. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 263-269 A TURN OF THE CENTURY CONUNDRUM—REEXAMINATION OF AEOLOTHYNNUS ASHMEAD (HYMENOPTERA: TIPHIIDAE: THYNNINAE) LYNN S. KIMSEY Bohart Museum of Entomology, Department of Entomology, University of California, Davis, CA 95616, U.S.A. (e-mail: Iskimsey @ucdavis.edu) Abstract.—The identity of the Australian thynnine genus Aeolothynnus Ashmead has been the source of nomenclatural confusion since the early 1900’s. Its identity is reevaluated relative to other related genera, species placements are reconsidered, and one new species, Aeolothynnus caliventer, from South Australia, is described. Asthenothynnus is discovered to be a new junior synonym of Aeolothynnus, and Turner’s concept of Aeolothynnus is in reality synonymous with Thynnoturneria Rohwer. New combinations of species in Aeolothynnus include: Thynnus beatrix Turner, Asthenothynnus deductor Turner, Thynnus generosus Turner, Asthenothynnus kurandensis Turner, Thynnus lactarius Turner, Asthen- othynnus lilliputianus Turner, Asthenothynnus maritimus Turner, Asthenothynnus minutis- simus Turner, Asthenothynnus perkinsi Turner, Asthenothynnus pleuralis Turner, Thynnus pulcherrimus Turner, and Asthenothynnus vicarius Turner. Key Words: tralia Many genera of Thynninae were origi- nally based on one or a small number of species, and the original characterizations were obscure at best. One of the finest ex- amples of the confusion resulting from these inadequate descriptions can be seen in the taxonomic literature near the turn of the century, published by taxonomists at the British Museum and at the U.S. National Museum. This confusion was largely the re- sult of miscommunications, misinterpreta- tions and a bit of transatlantic competition. Two generic names, Aeolothynnus Ash- mead and Asthenothynnus Turner, were pro- posed for the same group of species. The valid generic name for these species is Aeo- lothynnus and Turner’s mistaken concept of the genus Aeolothynnus is a very different entity later renamed Thynnoturneria Roh- wer, which in turn has been confused with Iswaroides Ashmead. Tiphiidae, Hymenoptera, Thynninae, Aeolothynnus, Asthenothynnus, Aus- Ashmead originally based Aeolothynnus on the new species multiguttatus. He de- scribed the species and genus simultaneous- ly in one of his notorious keys (1903). As a result of his habit of describing new gen- era and species in very brief keys, most subsequent authors confused the identity of Aeolothynnus. Although Ashmead did not clearly indicate the species as new, the ge- nus was monotypic, therefore as stated in the International Zoological Code (1985), Article 12, the generic description also ap- plied to the species. As a result, Turner’s 1908 statement that “Ashmead gives Aeo- lothynnus multiguttatus Ashm., as the type of his genus but, as he has not given any description of that species, his name cannot stand.” is incorrect. To further confuse the situation, Rohwer (1910a) subsequently recognized Ashmead’s designation of mul- tiguttatus and gave a new name to Turner’s 264 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON concept of Aeolothynnus as Turnerella, (1903) having given mere key distinc- stating that: tions. Rohwer (1910a) stated that T. “The characters given by Ashmead in his table of the genera of Thynnidae, are sufficient to satisfy the technical re- quirements so this species [Aeolothyn- nus multiguttatus Ashmead (nec Turn- er)] should date from that time and be accredited to Ashmead.” “Turner considering that Aelothynnus (sic!) multiguttatus Ashm, was unde- scribed named Thynnus cerceroides Sm. as the type of Aelothynnus. Aelothyn- nus multiguttatus Sm. and Thynnus cerceroides are not congeneric, which leaves Aelothynnus Turn, without a name. For this genus the name Tur- nerella may be used.” Unfortunately the name Turnerella ran into problems of homonymy, and according to Turner (911): “My identification of Ashmead’s genus, of which the type was undescribed, was incorrect, as has been pointed out by Mr. Rohwer, who renamed the genus Turnerella. That name, however, was used by Professor Cockerell for a ge- nus of bees; his paper was published in London on the same day as Mr. Rohwer’s paper was published in America, and I believe the name should be retained for the bee. I there- fore have to propose a new name for the genus.” Turner never made a new generic descrip- tion for his concept of Aeolothynnus. The uncertainty of the situation caused Given (1959) to lament: “The genus Aeolothynnus was erected by Ashmead in 1903 with the genotype A. multiguttatus. The genus was then very poorly defined and has been fre- quently misinterpreted by subsequent workers.” “Rohwer (1910a) published the first description of the genotype, Ashmead cerceroides and A. multiguttatus were not congeneric and therefore the genus Aeolothynnus Turner was left without a name as that name was valid for the genotype multiguttatus of Ashmead. Rohwer (1910a) proposed the name Turnerella for Turner’s genus. How- ever, this generic title was preoccu- pied, and both Turner (1911) and Roh- wer (1910b) appreciated this at about the same time. Turner (1911) then pro- posed the name Eurohweria for his ge- nus, but he was forestalled by Rohwer (1910b) who proposed the name Thyn- noturneria.” When Rohwer (1910b) renamed Aeolothyn- nus, as treated by Turner, he established Turner’s concept of the group as a valid ge- nus, particularly since none of the species placed by Turner (1910a) in Aeolothynnus under cerceroides were congeneric with multiguttatus. Thus the name Thynnotur- neria Rohwer applied to the cerceroides group of species. Turner (1912) was not en- tirely pleased by this situation: “Tam by no means sure that the name Aeolothynnus should not be used for this genus. Ashmead in describing the genus Aeolothynnus took an undescribed species for the type. In my work on the Thynnidae I accepted Ashmead’s ge- nus, but treated the species as a nomen nudum. Mr. Rohwer, on the other hand, holds that the description of the genus covers the species also; but I cannot agree with this opinion, as Ash- mead evidently did not intend the de- scription for a specific one, and a de- scription to be recognized should be at least intended by the author for a de- scription of a species.” ‘‘Unfortunately, A. cerceroides, Sm., selected by me as the type of the genus, does not appear to belong to the same genus as Ashmead’s type. Yet if Ash- mead’s specific name is treated as a VOLUME 101, NUMBER 2 nomen nudum, A. cerceroides must be treated as the type of the genus. It is bad enough to have to recognized the very insufficient descriptions of some authors as valid, but if we are also to accept what were never intended for descriptions things would be still worse. For the present, pending some decision on the subject, I am using Rohwer’s name, but do not consider that it can stand. The whole confusion is due to a want of editing in Ash- mead’s paper, as no editor should pub- lish a description of a genus with an undescribed species taken for the type.” Turner’s argument that Ashmead did not in- tend to describe multiguttatus as new at the same time as he described the genus was not accepted by other systematists. Simultaneously, Turner (1910a) also named a new genus Asthenothynnus, based on Thynnus pulchellus Klug. Upon exami- nation, Zhynnus pulchellus turns out to be congeneric with multiguttatus. Therefore Asthenothynnus also becomes synonymous with Aeolothynnus. In light of this generic confusion all of the species placed various- ly in Aeolothynnus, Thynnoturneria and As- thenothynnus need to be reevaluated for their correct placement. Some of this re- placement of species has been done below, showing new combinations as indicated. Types that have been seen and the generic placement confirmed are indicated by an as- terisk (*). However, those without either in- dication are placed in the appropriate genus based on whether Turner himself placed them in his concept of Aeolothynnus or one of the subsequent generic names for that en- tity, or placed them in Asthenothynnus, which is the equivalent of Ashmead’s Aeo- lothynnus. To further clarify Aeolothynnus Ashmead the genus is rediagnosed and discussed be- low, species placements are reassigned, and a new species, caliventer, is described, 265 which exhibits some very unusual thoracic modifications in the male. MATERIALS AND METHOD Specimens were studied in situ or were borrowed from the following institutions: the Natural History Museum, London, S. Lewis; Hope Museum, Oxford University, C. O’ Toole; and the Australian National In- sect Collection, CSIRO, Canberra, ACT, I. Naumann and J. Cardale. Aeolothynnus Ashmead (Figs. 1—10) Aeolothynnus Ashmead 1903: 101. Type species: Aeolothynnus multiguttatus Ash- mead 1903: 101. Monobasic. Asthenothynnus Turner 1910a: 34. Type species: Thynnus pulchellus Klug 1842: 20. Original designation. New synony- my. Aelothynnus Rohwer 1910a. Invalid emen- dation of Aeolothynnus Ashmead 1903. Male.—Body length 3-10 mm. Head: face flattened in profile, interantennal area nearly planar with clypeus; antennal sockets with dorsal antennal lobe highly reduced without carinae or transverse welt; suban- tennal area highly polished and impunctate; apical 7 to 9 flagellomeres somewhat ar- cuate or bulging along inner margin, with two small linear tyloids; clypeus nearly flat or slightly convex in profile, apical margin narrowly truncate; labrum smali, medially emarginate with narrow neck-like base; tongue short unmodified, prementum with apical brush of long setae reaching back to occipital foramen; maxillary stipes with short stipal fringe originating on inner mar- gin and extending transversely across stipes to outer margin; mandibles slender with small subapical tooth on inner margin; oral fossa extending to inner margin (Fig. 8) of mandible, widely separated from occipital carina; venter of head with large flattened and polished area, margined by carinae and surrounding the oral fossa. Thorax: prono- tum with transverse anterior carina well-de- 266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 5. caliventer 6. 3. caliventer - westwoodi 4. caliventer Figs. 1-10. i) ( a yy - my a 9 iii A multiguttatus 8. multiguttatus 10. caliventer 1—5, 9-10, Aeolothynnus caliventer. 6, A. westwoodi. 7—8, A. multiguttatus. 1, Ventral view of male thorax. 2, Lateral view of body. 3, Ventral view of genital capsule. 4, Lateral view of genital capsule. 5— 7, Apex of hypopygium. 8, Ventral view of male head. 9, Posterior view of apex of female abdomen. 10, Male apical gastral tergum. veloped; mesepisternum and mesopleuron separated by shallow scrobal groove; pro- podeum without discrete dorsal surface, gently sloping posteriorly; forefemur often indented basoventrally; forecoxae globular in most species; hindcoxa with short dorsal carina. Abdomen: basal segment slightly convex ventrally, foreshortened, dorsally without discrete dorsal surface; apical ter- gum hood-like, with narrow apical lip (Fig. 10); hypopygium ventrally longitudinally grooved or trough-like, apically variable, tridentate, ligulate or unidentate (Figs. 5—7); abdominal segments smoothly tapering one to the next (Fig. 2); terga II-IV with fine sublateral transverse sulcus; spiracles not apparent; terga with W-shaped transverse sulcus or line, marked by a row of setae near posterior margin. Genital capsule (Figs. 3, 4): penis valves large, dilated api- cally and often curved; volsella large flat- tened and tapering apically, forming floor of capsule; digitus not apparent; parameres slender and subtriangular; gonocoxa pro- duced into elongate, often apically notched dorsal lobe (Fig. 3); aedeagus short, origi- nating on this lobe (Fig. 3), with short api- cal loop. Color: body black with yellow, white and red markings; vertex with oblong reddish brown spot extending diagonally from dorsal eye margin to behind hindo- celli; abdomen in most species with odd VOLUME 101, NUMBER 2 comma-shaped pale markings on most seg- ments. Female.—Body length 2—5 mm; Head: broader than long or elongate and usually appearing pinched across at eyes; clypeus narrow and linear, shorter than interantennal distance; maxillary palpus with two articles, labial palpus with three; mandible edentate and sickle-shaped. Thorax: pronotum sub- quadrate; forecoxa unmodified or narrowed and separated by deep rectangular slot (multiguttatus); scutum and metanotum broadly visible dorsally. Abdomen: tergum I and III—V with broadly W-shaped trans- verse sulcus; tergum II with three transverse carinae or ridges; tergum V with thin sha- greened or roughened laterotergite marked by an arcuate bulge and sulcus; tergum VI with broadly or narrowly ovoid plate delim- ited at least laterally by carina, subtended by long tuft of setae (Fig. 9); sternum VI narrowly U-shaped. Color: pale brown. Distribution.—This genus occurs throughout southern Australia in New South Wales, South Australia, Western Aus- tralia, Tasmania, Victoria, and apparently the Northern Territory, although this record needs to be confirmed. Included species.—Thirty species are placed in Aeolothynnus including: beatrix (Turner) 1908* (Thynnus), new combina- tion; caliventer Kimsey, new species; de- coratus (Smith) 1879 (Thynnus); deductor (Turner) 1910b (Asthenothynnus), new combination; exiguus (Turner) 1910c (Thynnus); generosus (Turner) 1908* (Thynnus), new combination; incensus (Smith) 1868 (Thynnus); innocuus (Turner) 1908 (Thynnus); kurandensis (Turner) 1910d* (Asthenothynnus), new combina- tion; /actarius (Turner) 1910d* (Thynnus), new combination; /eucostictus (Turner) 1908 (Thynnus); lilliputianus (Turner) 1915a* (Asthenothynnus), new combina- tion; maritimus (Turner) 1915b* (Astheno- thynnus), new combination; minutissimus (Turner) 1910c (Asthenothynnus), new combination; minutus (Smith) 1859 (Thyn- nus); multiguttatus Ashmead 1903*; pene- 267 tratus (Smith) 1879 (Thynnus); perkinsi (Turner) 1910d* (Asthenothynnus), new combination; planiventris (Turner) 1908 (Thynnus); pleuralis (Turner) 1915a (As- thenothynnus), new combination; pulchel- lus (Klug) 1842 (Thynnus) (=Thynnus mul- tipictus Smith 1879); pulcherrimus (Turner) 1908* (Thynnus), new combination; pyg- maeus (Turner) 1908 (Thynnus); quadricar- inatus (de Saussure) 1867 (Thynnus); rub- romaculatus (Turner) 1908 (Thynnus); ten- uis (Turner) 1908 (Thynnus); vicarius (Turner) 1915a (Asthenothynnus), New combination; westwoodi (Guérin de Mene- ville) 1842 (Agriomyia) (=Thynnus intri- catus Smith* 1859); longiceps (Smith) 1859 (Thynnus); nanus (Smith) 1879 (Thynnus). Discussion.—Members of the genus Aeolothynnus are small-bodied and locally abundant Australian wasps. The vast ma- jority of species average | cm in length or less. Thousands of individuals, both males and females, may be found on a single flowering Eucalyptus tree. Members of the genus occur in most habitats throughout at least the southern part of Australia. They are for the most part unremarkably modi- fied. However, a new species, collected in South Australia has a bizarrely modified male. The male modifications in this spe- cies are unusual for the entire subfamily, so it is described as new below. Hosts are ap- parently unknown for Aeolothynnus. How- ever, given the parasitic habits of the rest of the subfamily, the hosts are undoubtedly small, locally abundant, species of larval Scarabaeidae. A number of traits are diagnostic for members of this genus. The most distinctive feature of the males is the longitudinally grooved or impressed apical abdominal sternum (hypopygium). This characteristic coupled with their smoothly tapering ab- domen, and flattened face will distinguish Aeolothynnus males from closely related genera. Aeolothynnus belongs to a group of genera characterized by the presence of a transverse carina, ridge or welt, across the 268 apical edge of the apical abdominal tergum (epipygium), the female apical abdominal tergum is smoothly convex, narrowed, with an oval medial plate margined laterally by a carina, which is in turn subtended later- ally by a long brush of setae. This group consists of Epactiothynnus Turner, Tmeso- thynnus Turner, Thynnoturneria Rohwer, [s- waroides Ashmead, Gymnothynnus Turner, Acanthothynnus Turner, Doratithynnus Turner and Aspidothynnus Turner. Females are problematic in this group of genera and too few are associated with males to distin- guish between species level and generic characteristics. Aeolothynnus caliventer Kimsey, new species (Figs. 1-5, 9, 10) Male.—Body length 8-9 mm; Head: face nearly flat from frons to upper clypeus; frons and vertex densely punctate, punc- tures small and contiguous; prementum strongly convex medially, with long medial fringe of setae; flagellum I 1.5X as long as broad; flagellum II 2.5 as long as broad; flagellum III three times as long as broad. Thorax (Figs. 1, 2): Pronotum extended ventrally; propleura convex; forecoxa ven- trally flat and expanded sharp-edged later- ally; forefemur flat, broadly expanded and rounded ventrally; mesepisternum strongly produced and flange-like, giving the thorac- ic venter a strongly cuplike appearance. Ab- domen: epipygium (Fig. 10); hypopygial apex subtruncate with rounded lateral cor- ners and short acute medial spine, ventral groove teardrop-shaped (Fig. 5). Genital capsule (Figs. 3, 4). Color: body black with pale whitish markings on apical margin of clypeus, a small spot on each antennal lobe, pronotum with small spot adjacent to tegula and on either end of transverse anterior ca- rina; small spot at posterior angle of me- sopleuron above midcoxa outlining meso- pleural lamellae, and small comma-shaped lateral spot on gastral terga I-VI; vertex with small oval red spot between hindocelli and nearest eye margin; fore- and midlegs PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with femoral apex and tibia entirely or part- ly red; hindfemur medially red; hindtibia somewhat reddish medially; wing mem- brane untinted, veins dark brown. Pubes- cence: long erect and silvery, except dense and golden along mesepisternal edge. Female.—Body length 5.5 mm; Head: head broader than long; mandible sickle- shaped and edentate. Thorax: pronotal disk with anterolateral corners acute, posterior margin with two submedial warts; propleu- ra strongly convex ventrally; scutum broad- ly visible; scutellum flattened, length sub- equal to breadth; propodeum strongly con- vex and rounded laterally and posteriorly. Abdomen: tergum I with W-shaped sulcus; II with three transverse ridges; HI-IV with large submedial U-shaped sulcus; pygidium narrowly longitudinally ovoid with lateral carina, with row of setae laterad of carina (Fig. 9); sternum VI apex narrowly U- shaped. Color: yellowish brown. Material examined.—Holotype ¢: Aus- tralia: SA, 79 km nnw Renmark, 33°31'S 140°24’E, 9 Aug.—7 Sept. 1995, K. R. Pul- len, Casuarina woodland, malaise trap. Ho- lotype deposited in the Australian National Insect Collection (ANIC), Canberra, ACT. Paratypes: two d, one 2, same data as ho- lotype except also collected in flight inter- cept trap (deposited in ANIC and Bohart Museum of Entomology, University of Cal- ifornia, Davis). These specimens derived from the Calperum Station/Bookmark Bio- sphere Reserve Invertebrate Survey. Etymology.—The name refers to the pe- culiar modifications of the male thorax; ca- lix = cup, venter = belly, Latin, noun. Discussion.—The unusually modified male thorax and forefemur will serve to dis- tinguish this species from other Aeolothyn- nus or related genera. The female is less remarkable but can be distinguished by shape of the pronotum with a small acute tooth on the anterolateral corner and pos- terior submedial swellings. In addition, the scutellum is relatively flat and the propleura are strongly bulging ventrally. VOLUME 101, NUMBER 2 ACKNOWLEDGMENTS Many thanks to Richard M. Bohart for reviewing the manuscript, and to the vari- ous collections and curators who made the study possible. LITERATURE CITED Ashmead, W. H. 1903. Classification of the fossorial, predaceous and parasitic wasps, of the superfam- ily Vespoidea. Canadian Entomologist 35: 95— 107. Given, B. B. 1959. Notes on Australian Thynninae. V. Aeolothynnus multiguttatus Ashmead and [swa- roides koebelei Ashmead. Proceedings of the Lin- naean Society of New South Wales 83: 400—402. Guérin de Meneville, EF E. 1842. Matériaux sur les Thynnides. Magasin de Zoologie 4: 1-15. International Code of Zoological Nomenclature. 1985. Third edition. University of California Press, Berkeley, xx + 338 pp. Klug, J. C. E 1842. Ueber die Insectenfamilie Heter- ogyna Latr. und die Gattung Thynnus E insbeson- dere. Physikalische und Mathematische Abhan- dlungen der Koniglichen Akademie der Wissen- schaften zu Berlin, pp. 1—44. Rohwer, S. A. 1910a. Turner’s genera of Thynnidae with notes on Ashmeadian genera. Entomological News 21: 345-351. . 1910b. A preoccupied generic name (Thyn- noturneria n. n.). Entomological News 21: 474. de Saussure, L. F 1867. Hymenoptera. /n Reise der Osterreichischen Fregatte Novara um die Erde in den Jahren, etc. Zoologischer Theil, Zweiter Band: Hymenoptera. Vol. 2, 156 pp. Smith, F 1859. Catalogue of the Hymenopterous in- 269 sects in the collection of the British Museum 7: 10-69. . 1868 (May). Descriptions of aculeate Hyme- noptera from Australia. Transactions of the Ento- mological 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. II. Proceedings of the Linnaean So- ciety of New South Wales, Sydney 33: 70—208 (June), 209-256 (August). . 1910a. Hymenoptera, Family Thynnidae. Jn Wytsman, P. ed., Genera Insectorum, 105. Brux- elles, 62 pp. . 1910b. Additions to our knowledge of the fos- sorial wasps of Australia. Proceedings of the Zoo- logical Society of London 1910: 259-306. . 1910c. New species of Thynnidae from the Australian and Austro-Malayan Regions. Annales Historico-Naturales Musei Nationalis Hungarici 8: 107-124. . 1910d. New fossorial Hymenoptera from Aus- tralia. Transactions of the Entomological Society of London 1910: 407—429. 1911. Notes on fossorial Hymenoptera. V. Further notes on the Thynnidae and Scoliidae. An- nals and Magazine of Natural History, London (8)8: 602-624. . 1912. Notes on fossorial Hymenoptera, IX. On some new species from the Australian and AustroMalayan Regions. Annals and Magazine of Natural History, London (8)10: 48-63. . 1915a. Descriptions of new fossorial wasps from Australia. Proceedings of the Zoological So- ciety of London 1915: 41-69. . 1915b. Notes on fossorial Hymenoptera. XVI. On the Thynnidae, Scoliidae and Crabronidae of Tasmania. Annals and Magazine of Natural His- tory (8)15: 537-559. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 270-273 NEW NORTH AMERICAN SPECIES AND RECORDS IN THE GENUS XENOLIMOSINA ROHACEK (DIPTERA: SPHAEROCERIDAE: LIMOSININAE) S. A. MARSHALL Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada N1G 2Wl Abstract.—Xenolimosina glabrigena, new species, is described from northern Florida, the female of Xenolimosina phoba Marshall is described, and new distributional records are given for X. phoba and X. sicula Marshall. Key Words: Xenolimosina Rohaéek is one of the most rarely collected genera of Sphaeroceridae, probably because most species seem to be active only during the colder months of the year. Xenolimosina previously included three species: X. setaria Villeneuve from Europe, X. sicula Marshall from Ontario, Quebec, Arkansas and California, and X. phoba Marshall from Quebec and Mary- land. The latter species was previously known only from males, and a description of the female is given below along with new distributional records for both de- scribed North American species. A third Nearctic species, Xenolimosina glabrigena new species, is described from northern Florida. Xenolimosina glabrigena is recog- nisable as a Xenolimosina by the long ex- serted hind tibial bristle, small eyes, tele- scoping female abdomen, lack of a mid ventral bristle on the mid tibia, two spinose lobes on the male fifth sternite (Fig. 2), and the short surstylus with a long-setose pos- terolateral surface (Fig. 3). Xenolimosina glabrigena will key out to X. sicula in Mar- shall (1985), but it is distinctly different from X. sicula in details of the male and female genitalia. Most notably, the disti- Sphaeroceridae, Diptera, Florida insects phallus of X. glabrigena is simple, broad and tubular, much like X. phoba (Marshall 1985, fig. 23), but in marked contrast to the broad, spinose and highly modified disti- phallus of X. sicula (Marshall 1985, fig. 22) and X. setaria (Rohaéek 1983, fig. 322). Some of the characteristics of X. glabri- gena, such as the dense pile of the male fore tibia and basitarsus, the shape of the postgonite (Fig. 1), and the sclerotization of the terminal sclerites of the female abdo- men, are distinctive autapomorphies not found in the other species. Rohacek (1982) suggested that Xenoli- mosina belongs to the Minilimosina genus- group, and that it is probably the sister ge- nus to Minilimosina. The distiphallus in Minilimosina species is generally simple and unadorned, suggesting that the charac- teristically adorned distiphallus of X. sicula and the European species X. setaria are synapomorphic, and these two species form a monophyletic group. The other two spe- cies in the genus, X. glabrigena and X. pho- ba, both have an elongate tubular distiphal- lus (Fig. 1) and a greatly enlarged costagial bristle, and together probably form the sis- ter group to X. setaria plus X. sicula. VOLUME 101, NUMBER 2 271 postgonite Male terminalia, left lateral. 4, Male terminalia, posterior. 5, Female terminalia, dorsal. 6, Female terminalia, ventral. 7, Spermathecae. 8-10. X. phoba. 8, Spermathecae. 9, Female terminalia, dorsal. 10, Female terminalia, ventral. Figs. 1-10. 1-7, Xenolimosina glabrigena. 1, Aedeagus and associated structures. 2, Male sternites 5—7. 3, 272 Xenolimosina glabrigena Marshall, new species (Figs. 1-7) Description.—Body length ca. 2.0 mm, dark brown to black with a heavy pruinos- ity; lower frons and gena reddish; legs brown. Interfrontal plate 1.5 as high as width at middle, bordered by 4 pairs of equal interfrontal bristles, lower two pairs weakly cruciate. Postocellar bristles absent. Eye small, height ca. 1.3 genal height, gena shining except for lower margin and a narrow vertical strip on posterior third. Palpus strongly swollen at middle, tapered at apex. Scutum with 5—6 rows of acrosti- chal bristles between anterior dorsocentral bristles; dorsocentral bristles in 2 pairs, an- terior pair 0.6 as long as posterior pair, posterior pair equal to scutellar length. Prescutellar acrostichal bristles in a single pair twice as long as acrostichal setulae. Foreleg of male with dense yellow pile ven- trally on distal half of tibia and on tarso- mere one; foreleg of female unmodified. Mid tibia with long proximal anterodorsal distal anterodorsal and distal dorsal bristles. Katepisternum pruinose, with a short pos- terodorsal bristle and a minute anterodorsal bristle. Wing length 2.5 width; second costal sector 1.0—1.4 third; costa extend- ing about 2 vein-widths beyond apex of R,,;. Costagial bristle very long, longer than alula; alula narrow. Male abdomen: Sternite 5 (Fig. 2) with two closely approximated posteromedial spinose patches, together making up a prominent posteromedial lobe; basal parts of spinose patches with continuous rows of spines, distal parts tapered and with smaller, separated spinules. Sternite 6 simple medi- ally, expanded into a broad, pale part con- nected with a broad, thin-rimmed ring scler- ite posteriorly. Epandrium (Fig. 3) uniform- ly setose; surstylus with a quadrate, ven- trally notched anteroventral lobe, a rounded posteroventral lobe with 3 stout bristles, and a densely long-setose lateral swelling. Cercus (Fig. 4) weakly differentiated from PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON epandrium, bare dorsally and medially, long setose ventrolaterally, cerci bent into broad posterior lobes at middle, not fused to form a subanal plate but narrowly contiguous ventrally. Subepandrial sclerite very broad, quadrate, articulating with inner ventral corners of cercus medially and posterior dorsal corners of surstyli laterally. Hypan- drium with a well developed anterior apo- deme; lateral arm fused with epandrium and contiguous with surstylus. Postgonite (Fig. 1) (paramere or gonostylus of authors) weakly S-shaped, distal part swollen at bend then narrowed at apex. Basiphallus simple, rounded; distiphallus simple, nar- row and tubular, ending in two dark lobes and a small, weakly spinulose membrane. Ejaculatory apodeme well developed. Female abdomen (Figs. 5—6): Tergite 8 equal in length to tergite 7, but with tripar- tite pigmentation, middle part small and pale. Tergite 10 pale anteromedially and posteromedially, with a bristle on each half. Cercus short, with 2 long, thin apical bris- tles and a long, thin dorsal bristle. Sternite 8 small, subequal in length to sternite 7 but less than half as wide, bare on anterior third, otherwise setulose and setose. Vagi- nal area weakly sclerotised with 2 ring- shaped sclerites. Sternite 10 densely setu- lose on posterior half, with a marginal row of bristles; anterior half pale and bare, with a deep, keyhole-shaped anteromedial depig- mented area. Spermathecae (Fig. 7) dark, acorn-shaped; sclerotised parts of ducts very short. Holotype—UNITED STATES. Florida: Levy Co., Archer, on rotting fungus on sand, 17.xii.1997, S.A. Marshall (¢, Uni- versity of Guelph). Paratypes—UNITED STATES. Florida: Marion Co., Ocala National Forest, 29.1.1986, flight intercept trap, R.&M. Mar- shall (22, University of Guelph). Xenolimosina phoba Marshall (Figs. 8-10) Xenolimosina phoba Marshall 1985: 764 (male only). VOLUME 101, NUMBER 2 Description of female terminalia (Figs. 8—10).—Tergite 8 completely divided; ter- gite 10 uniformly pigmented posteriorly, anteromedially pale. Cercus long, thin, with long apical, preapical and dorsal bristles. Sternite 8 large, longer than sternite 7 and over half as wide, almost entirely setulose, with 2 transverse rows of bristles. Sternite 10 with tripartite pigmentation anteriorly, middle part bare. Spermathecae (Fig. 8) acorn-shaped, sclerotised parts of ducts very short. New records since 1985.—CANADA. Quebec; Old: Chelsea, 17-x.1988. .J_R. Vockeroth (1 2, Canadian National Collec- tion, Ottawa). Ontario, St. Joseph’s Island, Hilton Beach, fish entrail baited pan traps in hardwood forest, ix—x.1987, J. Swann (8 3, University of Guelph). Comments.—The first and only female of this species was collected by Dr. Richard Vockeroth, who collected the male holotype at the same locality, and during the same month and week, twenty four years earlier. 273 Xenolimosina sicula Marshall Xenolimosina sicula Marshal] 1985: 765. New distributional records.—In addition to the original records from Ontario, Que- bec, California and Arkansas, specimens have been examined from the following lo- calities: CANADA. British Columbia: Car- manah Valley. Ontario: Wellington Co., Al- goma Co. UNITED STATES. New Hamp- shire: Strafford Co. Florida: Leon Co. All new records are from October-November. LITERATURE CITED Marshall, S. A. 1985. The genera Xenolimosina and Terrilimosina (Diptera: Sphaeroceridae: Limosi- ninae) in North America. Proceedings of the En- tomological Society of Washington 87: 759-769. Rohacek, J. 1982—3. A monograph and reclassification of the previous genus Limosina Macquart (Dip- tera, Sphaeroceridae) of Europe. Part 1, Beitraige zur Entomologie 32: 195—282 and Part 2 Beitrige zur Entomologie 33: 3-195. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 274-284 LIST OF SPECIES OF NEOTROPICAL MEGALOPTERA (NEUROPTERIDA) ATILANO CONTRERAS-RAMOS Instituto de Biologia, U.N.A.M., Departamento de Zoologia, Apdo. Postal 70-153, 04510 México, D. EF, Mexico (e-mail: atilano@mail.ibiologia.unam.mx) Abstract.—The 63 known species and subspecies of Neotropical dobsonflies and al- derflies (Neuropterida: Megaloptera) are listed. Synonymy, authors, bibliographic refer- ences, and distribution of species by country and generally by province or state are in- cluded. Six localities, each of specimens thought to belong to new species, are also pro- vided. Resumen.—Se proporciona una lista de las 63 especies y subespecies conocidas de megalopteros (Neuropterida: Megaloptera) neotropicales. Se incluye sinonimia, autores, referencias bibliograficas y la distribuci6n de las especies por pais y generalmente también por provincia o estado. Se enumeran seis localidades de ejemplares que se cree pertenecen a especies nuevas para la ciencia. Key Words: Corydalidae, Sialidae, dobsonflies, fishflies, alderflies, Megaloptera, Neo- tropics, taxonomy, species list Megaloptera is a relatively primitive en- dopterygote (holometabolous) group close- ly related to Raphidioptera and Planipennia (Neuroptera s. s.). Despite being a small group, of approximately 200 (Evans 1972) to 300 species (New and Theischinger 1993) worldwide, the alpha taxonomy of the Neotropical megalopteran fauna has only recently reached a satisfactory level of knowledge. Since the most recent species list for the Neotropics (Penny 1977) does not include results of subsequent revisions, it seems useful to provide a current list of the species with references to relevant lit- erature (1.e., original species descriptions, taxonomic revisions, significant distribution data). Species synonymy according to the most recent revision is included, as are spe- cies distributions by country and province or state if the latter were published or could be obtained. Ordering of taxa is alphabeti- cal. Several records (Corydalus and Platy- neuromus) are given as “‘sp. 1, sp. 2’’, etc., and for these locality data are given. This is in the hope that new material may be collected and the identity of these possibly new species might be established. Follow- ing Penny (1977), species of Sialidae are included in Protosialis. However, their ex- act generic position is rather uncertain at this time, as the group needs revision (e.g., Ross [1937] synonymized Protosialis under Sialis). In accordance to Cabrera and Wil- link’s (1980) limits of the Neotropical Re- gion, the list includes all species and sub- species recorded south of the United States—Mexico border. The corydalid spe- cies Corydalus cornutus (L.) and Dysmi- cohermes sp. have been collected in Texas, close to or at the Mexican border (Contrer- as-Ramos 1995b, 1998), so there is the pos- sibility they occur in Mexico (Tamaulipas). Davis (1903) included Mexico as part of the distribution range of Nigronia fasciatus VOLUME 101, NUMBER 2 (Walker). However, the southernmost dis- tribution otherwise recorded for this species is in Florida (Hazard 1960). The 63 valid species and subspecies in this list, contrast to the 46 species recorded for the Megaloptera of America north of Mexico (Evans and Neunzig 1996; 43 spe- cies [3 Corydalus spp. missing] in Penny et al. 1997). The highest diversity in the Neo- tropics lies within Corydalinae (46 spp., compared to 9 species and subspecies of Chauliodinae and 8 species of Sialidae), whereas Sialidae is the most diverse group in the Nearctic (24 spp., compared to 18 species of Chauliodinae and 4 species of Corydalinae). Four species (1 Chauliodinae and 3 Corydalinae) are shared between both regions. Family Corydalidae Subfamily Chauliodinae (fishflies) Genus Archichauliodes Weele 1909 1. Archichauliodes chilensis Kimmins. Archichauliodes chilensis Kimmins 1954: Flint 1973; Penny 1977. Distribution CHILE: Arauco, Coquim- bo, Curic6, Malleco, Santiago, Valdivia, Valparaiso. 2. Archicahuliodes pinares Flint. Archicahuliodes pinares Flint 1973. Distribution. CHILE: Concepci6n. Genus Neohermes Banks 1908 3. Neohermes filicornis (Banks). Chauliodes filicornis Banks 1903b. Neohermes filicornis (Banks): Weele 1910; Flint 1965; Evans 1972; Contreras-Ra- mos 199 Tay 9977; Distribution UNITED STATES: Ari- zona, California, New Mexico; MEXICO: Baja California, Sonora. Genus Nothochauliodes Flint 1983 4. Nothochauliodes penai Flint. Nothochauliodes penai Flint 1983. Distribution CHILE: Maule. Genus Protochauliodes Weele 1909 5. Protochauliodes bullocki Flint Protochauliodes bullocki Flint \973. Distribution.—CHILE: Bio-Bio, Linares, Malleco, Nuble. 6. Protochauliodes cinerascens cinerascens (Blanchard). Chauliodes cinerascens Blanchard 1851. Neohermes cinerascens (Blanchard): Banks 1908. Protochauliodes cinerascens (Blanchard): Weele 1910; Kimmins 1954. Protochauliodes cinerascens cinerascens (Blanchard): Flint 1973. Distribution CHILE: Curic6, Linares, Nuble, O’ Higgins, Santiago, Talca. 7. Protochauliodes cinerascens fumipennis Flint. Protochauliodes cinerascens fumipennis Flint 1973. Distribution CHILE: Concepcion. 8. Protochauliodes cinerascens reedi Kim- mins. Protochauliodes reedi Kimmins 1954. Protochauliodes cinerascens reedi mins: Flint 1973. Distribution. CHILE: Valparaiso. Kim- 9. Protochauliodes humeralis (Banks). Neohermes humeralis Banks 1908. Protochauliodes humeralis (Banks): Weele 1910; Flint 1973. Distribution CHILE: Arauco, Bio-Bio, Cautin, Concepcion, Malleco, Maule. Subfamily Corydalinae (dobsonflies) Genus Chloronia Banks 1908 10. Chloronia absona Flint. Chloronia absona Flint 1992: Contreras- Ramos 1995a. Distribution COSTA RICA: Alajuela, Guanacaste, Limon, San José. 276 11. Chloronia antilliensis Flint. Chloronia antilliensis Flint 1970: Penny and Flint 1982; Contreras-Ramos 1995a. Distribution DOMINICA. 12. Chloronia banksiana Penny and Flint. Chloronia banksiana Penny and Flint 1982: Contreras-Ramos 1995a. Chloronia bogotana (not Weele, misidenti- fication): Banks 1943. Distribution. VENEZUELA: Aragua, Carabobo. 13. Chloronia bogotana Weele. Chloronia bogotana Weele 1909: Penny and Flint 1982; Flint 1991; Contreras-Ra- mos 1995a. Distribution. BOLIVIA: La Paz; CO- LOMBIA: Cundinamarca; ECUADOR: Za- mora-Chinchipe; PERU: Cuzco, Huanuco. 14. Chloronia convergens Contreras-Ra- mos. Chloronia convergens Contreras-Ramos 1995a. Distribution. ECUADOR: Pichincha. 15. Chloronia corripiens (Walker). Hermes corripiens Walker 1858. Neuromus corripiens (Walker): Lachlan 1869. Neuromus winthemi Davis 1903. Chloronia meridionalis Weele 1909. Chloronia winthemi (Davis): Weele 1910. Chloronia ocellaris Navas 1934a. Chloronia corripiens (Walker): Penny and Flint 1982; Contreras-Ramos 1995a. Mac- Distribution BRAZIL: Espirito Santo, Minas Gerais, Rio de Janeiro, Santa Catar- ina, Sao Paulo. 16. Chloronia gloriosoi Penny and Flint. Chloronia gloriosoi Penny and Flint 1982: Flint 1992; Contreras-Ramos 1995a. Distribution COSTA RICA: San José; PANAMA: Chiriquf. 17. Chloronia hieroglyphica (Rambur). Neuromus hieroglyphica Rambur 1842. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hermes hieroglyphicus (Rambur): Walker 1853. Corydalis hieroglyphicus (Rambur): Hagen 1861. Chloronia hieroglyphica (Rambur): Banks 1908; Penny and Flint 1982; Contreras- Ramos 1995a. Distribution.— BRAZIL: Amazonas, Para; FRENCH GUIANA: GUYANA; PERU: Loreto, Madre de Dios. 18. Chloronia mexicana Stitz. Chloronia mexicana Stitz 1914: Penny and Flint 1982; Flint 1992; Contreras-Ramos 1995a, 1997. Distribution COSTA RICA: Alajuela, Guanacaste, Heredia; GUATEMALA: Alta Verapaz, Suchitepequez; MEXICO: Chia- pas, Morelos, San Luis Potosi, Tamaulipas, Veracruz. 19. Chloronia mirifica Navas. Chloronia mirifica Navas 1925a: Penny and Flint 1982; Flint 1992; Contreras-Ramos 1995a, 1997. Chloronia hieratica Navas 1928a. Distribution.—COLOMBIA: Meta; COSTA RICA: Alajuela, Cartago, Guana- caste, Heredia, Limon, Puntarenas, San José; ECUADOR: Napo, Pichincha; GUA- TEMALA: Alta Verapaz, Izabal, Solola; MEXICO: Oaxaca, Veracruz; PANAMA: Chiriqui, Col6n; PERU: Hudnuco. 20. Chloronia osae Flint. Chloronia osae Flint 1992: Contreras-Ra- mos 1995a. Distribution.—COSTA RICA: Puntaren- as. 21. Chloronia pallida (Davis). Neuromus pallidus Davis 1903. Chloronia pallidus (Davis): Penny 1977. Chloronia pallida (Davis): Penny and Flint 1982; Contreras-Ramos 1995a, 1997. Distribution. —MEXICO: Chihuahua, Guerrero, Jalisco, Michoacan, Morelos, Nayarit. VOLUME 101, NUMBER 2 22. Chloronia plaumanni Penny and Flint. Chloronia plaumanni Penny and Flint 1982: Contreras-Ramos 1995a. Distribution. — BRAZIL: Rio Grande do Sul, Santa Catarina. 23. Chloronia zacapa Contreras-Ramos. Chloronia zacapa Contreras-Ramos 1995a. Distribution. GUATEMALA: Izabal, Zacapa. Genus Corydalus Latreille 1802 24. Corydalus affinis Burmeister. Corydalus affinis Burmeister 1839: Weele 1910 (in part); Penny 1977; Contreras- Ramos 1998. Corydalus nubilus (not Erichson, misiden- tification): Weele 1910; Banks 1943; Penny 1982. Corydalus sp.: Glorioso 1981. Corydalus spec. nov.: Geijskes 1984. Distribution. —ARGENTINA: Chaco, Misiones; BOLIVIA: Beni; BRAZIL: Acre, Amapa, Amazonas, Mato Grosso, Para, Rondonia, Roraima, Sao Paulo; COLOM- BIA: Antioquia, Boyaca, Choco, Cundina- marca, Tolima; ECUADOR: Napo, Pichin- cha, Sucumbifos; FRENCH GUIANA; GUYANA; PARAGUAY; PERU: Cuzco, Loreto, Madre de Dios; VENEZUELA: Guarico. 25. Corydalus amazonas Contreras-Ramos. Corydalus amazonas Contreras-Ramos 1998. Distribution.—BRAZIL: RondOonia. Amazonas, 26. Corydalus armatus Hagen. Corydalis armata Hagen 1861. Corydalis armatus Hagen 1861. Corydalus armatus Hagen 1861: Weele 1910 (in part); Stitz 1914 (in part); Navas 1920a, 1934a, 1935; Penny 1977 (in part), 1982 (in part); Glorioso 1981 (in part); Contreras-Ramos 1998. Corydalis armata n. sp.: Davis 1903 (in part). Corydalus quadrispinosus Stitz 1914. Corydalus peruvianus (not Davis, misiden- tification): Banks 1943 (in part). Distribution ARGENTINA: Catamar- ca, Jujuy, Salta, Tucuman; BOLIVIA: Chu- quisaca, Cochabamba, La Paz, Santa Cruz; COLOMBIA: Boyaca, Cundinamarca, Val- le del Cauca; ECUADOR: Bolivar, Chim- borazo, El Oro, Esmeraldas, Imbabura, Loja, Los Rios, Napo, Pichincha, Tungura- hua, Zamora Chinchipe; PERU: Ayacucho, Cuzco, Hudnuco, Junin, Lima, Pasco; VENEZUELA: Aragua, Mérida, Tachira. 27. Corydalus arpi Navas. Corydalus arpi Navas 1936: Penny 1977, 1982; Contreras-Ramos 1993, 1998. Distribution.—BRAZIL: Amazonas, Rondonia; VENEZUELA: Territorio Fed- eral Amazonas. 28. Corydalus australis Contreras-Ramos. Corydalus australis Contreras-Ramos 1998. Corydalus affinis (not Burmeister, misiden- tification): Weele 1910 (in part); Penny 1977 (in part); Glorioso 1981. Distribution ARGENTINA: Misiones; BRAZIL: Minas Gerais, Rio Grande do Sul, Santa Catarina; URUGUAY: Artigas. 29. Corydalus batesii MacLachlan. Corydalus batesii MacLachlan 1868: Davis 1903; Geijskes 1984; Contreras-Ramos 1998. Corydalus batesi MacLachlan: Weele 1910; Stitz 1914 (in part); Kimmins 1970; Pen- ny 1977, 1982; Glorioso 1981. Distribution BOLIVIA: Cochabamba, Santa Cruz; BRAZIL: Amazonas, Para; COLOMBIA: Antioquia; ECUADOR: Napo; FRENCH GUIANA; GUYANA; SURINAME; PERU: Madre de Dios; VENEZUELA: Territorio Federal Amazon- as. 30. Corydalus bidenticulatus Contreras-Ra- mos. Corydalus bidenticulatus Contreras-Ramos 1998. 278 Corydalus lutea (not Hagen, misidentifica- tion): Glorioso 1981 (in part). Corydalus sp. B: Contreras-Ramos 1997. Distribution UNITED STATES: Ani- zona; MEXICO: Colima, Guerrero, Jalisco, Michoacan, Morelos, Nayarit, Oaxaca, Sin- aloa, Sonora. 31. Corydalus cephalotes Rambur. Corydalus cephalotes Rambur 1842: Weele 1910 (in part); Stitz 1914 (in part); Penny 1977 (in part), 1982 (in part); Contreras- Ramos 1998. Corydalis affinis (not Burmeister, misiden- tification): Walker 1853; Hagen 1861. Neuromus cephalotes (Rambur): Davis 1903 (in part). Corydalus intricatus Navas 1921: Penny 1977. Distribution.—BRAZIL: Rio de Janeiro. 32. Corydalus clauseni Contreras-Ramos. Corydalus clauseni Contreras-Ramos 1998. Distribution.—COLOMBIA: Valle del Cauca; COSTA RICA: Heredia; ECUA- DOR: Canar, Loja, Napo, Pichincha, Tun- gurahua. 33. Corydalus colombianus Contreras-Ra- mos. Corydalus colombianus Contreras-Ramos 1998. Corydalus ecuadorianus (not Banks, mis- identification): Glorioso 1981 (in part). Distribution. COLOMBIA: Valle del Cauca. 34. Corydalus diasi Navas. Corydalus diasi Navas 1915: Penny 1977; Contreras-Ramos 1998. Corydalus finoti Navas 1921: Penny 1977. Distribution. ARGENTINA: Misiones, BRAZIL: Bahia, Ceara, Goias, Minas Ger- ais, Rio Grande do Sul, Sao Paulo; PAR- AGUAY. 35. Corydalus ecuadorianus Banks. Corydalus ecuadorianus Banks 1948: Pen- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ny 1977; Glorioso 1981 (in part); Con- treras-Ramos 1998. Distribution ECUADOR: Napo, Tun- gurahua. 36. Corydalus flavicornis Stitz. Corydalus armatus flavicornis Stitz 1914: Penny 1977. Corydalus nevermanni Navas 1934b: Penny 1977; Banks 1943. Corydalus camposi Navas 1935: Penny OTe Corydalus armatus (not Hagen, misidenti- fication): Banks 1943 (in part). Corydalus peruvianus (not Davis, misiden- tification): Banks 1943 (in part). Corydalus flavicornis Stitz: Glorioso 1981; Contreras-Ramos 1998. Distribution COLOMBIA: Antioquia, Meta, Santander; COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia, Limon, Pun- tarenas, San José; ECUADOR: Guayas, Es- meraldas, Loja, Los Rios, Napo, Pastaza, Pichincha, Tungurahua; EL SALVADOR; GUATEMALA; HONDURAS; PANAMA: Chiriqui; PERU: Hudnuco, Junin, Pasco; VENEZUELA: Aragua, Bolivar, Lara, Mé- rida, Portuguesa, Zulia. 37. Corydalus flinti Contreras-Ramos. Corydalus flinti Contreras-Ramos 1998. Distribution VENEZUELA: Territorio Federal Amazonas. 38. Corydalus hecate MacLachlan. Corydalis hecate MacLachlan 1866: Kim- mins 1970. Neuromus cephalotes (not Rambur, mis- identification): Davis 1903 (in part). Corydalus cephalotes (not Rambur, mis- identification): Weele 1910 (in part); Stitz 1914 (in part); Penny 1977 (in part), 1982 (in part); Glorioso 1981 (in part). Corydalus raymundoi Navas 1920b. Corydalus sallei Navas 1920b. Corydalus hecate MacLachlan: Contreras- Ramos 1998. VOLUME 101, NUMBER 2 Distribution.—BRAZIL: Distrito Feder- al, Espirito Santo, Minas Gerais, Sao Paulo; PERU: VENEZUELA: Distrito Federal. 39. Corydalus holzenthali Contreras-Ra- mos. Corydalus holzenthali Contreras-Ramos 1998. Distribution.—BOLIVIA: Cochabamba, layPaz PERU: Pasco: 40. Corydalus ignotus Contreras-Ramos. Corydalus ignotus Contreras-Ramos 1998. Distribution. FRENCH GUIANA. 41. Corydalus imperiosus Contreras-Ra- mos. Corydalus imperiosus Contreras-Ramos 1998. Corydalus tridentatus (not Stitz, misidenti- fication): Glorioso 1981 (in part). Distribution ARGENTINA: Misiones. 42. Corydalus longicornis Contreras-Ra- mos. Corydalus longicornis Contreras-Ramos 1998. Distribution. ARGENTINA: Catamar- ca, Salta; BOLIVIA: Chuquisaca, Cocha- bamba, Santa Cruz; ECUADOR: Zamora Chinchipe. 43. Corydalus luteus Hagen. Corydalis lutea Hagen 1861: Davis 1903 (in part); Penny 1977 (as nomen nudum). Corydalus luteus Hagen: Weele 1910 (as ju- nior synonym of C. cornutus [L.]); Con- treras-Ramos 1997, 1998. Corydalus lutea Hagen: Glorioso 1981 (in part). Corydalis crassicornis MacLachlan 1868: Davis 1903; Banks 1907; Kimmins 1970. Corydalus crassicornis (MacLachlan): Weele 1910 (as junior synonym of C. cornutus [L.]). Corydalis inamabilis MacLachlan 1868: Davis 1903; Banks 1907; Kimmins 1970. Corydalus inamabilis (MacLachlan): Weele 1910 (as junior synonym of C. cornutus [L.]). 279 Corydalus armatus laevicornis Stitz 1914: Penny 1977. Distribution BELIZE; COSTA RICA: Alajuela, Guanacaste, Heredia, Lim6n, Puntarenas, San José; EL SALVADOR; UNITED STATES: Texas; GUATEMALA: Alta Verapaz, Chiquimula, Escuintla, Gua- temala, Sacatepéquez, San Marcos, Santa Rosa, Suchitepequez, Zacapa; HONDU- RAS; MEXICO: Chiapas, Coahuila, Hidal- go, Nuevo Leon, Oaxaca, Querétaro, San Luis Potosi, Tabasco, Tamaulipas, Vera- cruz; NICARAGUA; PANAMA: Zona del Canal, Chiriqui, Colon. 44. Corydalus magnus Contreras-Ramos. Corydalus magnus Contreras-Ramos 1998. Corydalus lutea (not Hagen, misidentifica- tion): Glorioso 1981 (in part). Corydalus sp. M: Contreras-Ramos 1997. Distribution COSTA RICA: Alajuela, Guanacaste, Puntarenas; EL SALVADOR; GUATEMALA: Alta Verapaz, Baja Vera- paz, Suchitepequez; MEXICO: Chiapas, Puebla, San Luis Potosi, Veracruz. 45. Corydalus neblinensis Contreras-Ra- mos. Corydalus neblinensis Contreras-Ramos 1998. Distribution. VENEZUELA: Territorio Federal Amazonas. 46. Corydalus nubilus Erichson. Corydalis nubila Erichson 1848: Hagen 1861 (in part); Davis 1903 (in part). Corydalus nubilus Erichson: Stitz 1914; Penny 1977 (in part); Glorioso 1981; Ge- ijskes 1984; Contreras-Ramos 1998. Corydalus nevermanni (not Navas, mis- identification): Penny 1982. Corydalus titschacki Navas 1928b: Penny L977 Distribution.—BRAZIL: Amazonas, Para, Roraima; FRENCH GUIANA; GUY- ANA; VENEZUELA: Territorio Federal Amazonas. 47. Corydalus parvus Stitz. 280 Corydalus parvus Stitz 1914: Penny 1977; Contreras-Ramos 1998. Corydalus armatus (not Hagen, misidenti- fication): Glorioso 1981 (in part). Distribution ECUADOR: Pastaza, Za- mora Chinchipe; PERU: Cuzco, Huanuco, Pasco. 48. Corydalus peruvianus Davis. Corydalis peruviana Davis 1903. Corydalus armatus (not Hagen, misidenti- fication): Weele 1910 (in part); Stitz 1914 (in part); Penny 1977 (in part), 1982 (in part); Glorioso 1981 (in part). Corydalis crassicornis (not MacLachlan, misidentification): Banks 1914. Corydalus primitivus fera Navas 1927: Pen- ny 1977. Corydalus peruvianus Davis: Banks 1943 (in part); Contreras-Ramos 1997, 1998. Distribution.—ARGENTINA: | Jujuy, Salta, Tucuman; BOLIVIA: Chuquisaca, Cochabamba, La Paz, Tarija, COLOMBIA: Antioquia, Cundinamarca, Meta, Putumayo, Valle del Cauca; COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia, Lim6n, Pun- tarenas, San José; ECUADOR: Cotopaxi, Guayas, Los Rios, Morona Santiago, Napo, Pastaza; Pichincha; Sucumbios; GUATE- MALA: Escuintla, Izabal, Quiché, San Marcos; MEXICO: Chiapas, Oaxaca, Pueb- la, Tabasco, Veracruz; PANAMA: Bocas del Toro, Chiriqui, Col6n, Darién; PERU: Amazonas, Huanuco, La Libertad, Pasco; VENEZUELA: Aragua, Barinas, Distrito Federal, Mérida. 49. Corydalus primitivus Weele. Corydalus primitivus Weele 1909: Weele 1910 (in part); Stitz 1914; Navas 1929; Penny 1977, 1982 (in part); Glorioso 1981 (in part), Contreras-Ramos 1998. Distribution ARGENTINA: Catamar- ca, Jujuy, Salta, Tucuman; BOLIVIA: Santa Cruz. 50. Corydalus tesselatus Stitz. Corydalus batesi tesselatus Stitz 1914. Corydalis nubila (not Erichson, misidenti- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fication): Hagen 1861 (in part); Davis 1903 (in part); Banks 1943. Corydalus bolivari Banks 1943: Glorioso 1981; Contreras-Ramos 1993. Corydalus tesselatus Stitz: Penny 1977 (in- correct locality); Contreras-Ramos 1998. Distribution. —COLOMBIA; VENE- ZUELA: Aragua, Distrito Federal, Mérida, Tachira. 51. Corydalus texanus Banks. Corydalis texana Banks 1903a: Penny 1977 (as junior synonym of C. cornutus [L.]). Corydalis cognata (not Hagen, misidentifi- cation): Banks 1892, 1903b, 1907; Davis 1903; Chandler 1956. Neuromus pallidus (not Davis): Davis 1903 (mislabeled photograph). Corydalus pallidus (not Davis): Weele 1910 (invalid combination based on mislabeled photograph; as junior synonym of C. cor- nutus [L.]). Corydalus cognatus (not Hagen, misiden- tification): Weele 1910 (in part; as junior synonym of C. cornutus [L.]); Evans 1972 (in part). Corydalus texanus Banks: Weele 1910 (as junior synonym of C. cornutus [L.]); Contreras-Ramos 1997, 1998. Corydalus cornutus (not Linnaeus, mis- identification): Stitz 1914 (in part); Glo- rioso 1981 (in part); Hermann and Davis 1991: Corydalus similis Stitz 1914: Banks 1943 (misidentification); Penny 1977 (in part). Corydalus constellatus Navas 1934b: Pen- ny 1977: Distribution —UNITED STATES: Ari- zona, California, Colorado, Nevada, New Mexico, Texas, Utah; GUATEMALA: Baja Verapaz, Chiquimula; MEXICO: Baja Cal- ifornia, Baja California Sur, Chiapas, Chi- huahua, Colima, Distrito Federal, Guerrero, Jalisco, Michoacan, Morelos, Nayarit, Oa- xaca, Puebla, Querétaro, Sinaloa, Sonora, Veracruz. 52. Corydalus tridentatus Stitz. Corydalus tridentatus Stitz 1914: Penny VOLUME 101, NUMBER 2 1977; Glorioso 1981 (in part); Contreras- Ramos 1998. Corydalus tridentatus nigripes Stitz 1914. Distribution.—BRAZIL: Espirito Santo. Parana, Rio Grande do Sul. Corydalus sp. 1. Corydalus sp. 1 (near C. affinis Burmeis- ter): Contreras-Ramos 1998. Distribution.—BRAZIL: Sao Paulo: Sao José des Campos. Corydalus sp. 2 Corydalus tesselatus (not Stitz, misidenti- fication): Banks 1943. Corydalus ecuadorianus (not Banks, mis- identification): Glorioso 1981 (in part). Corydalus sp. 2 (near C. ecuadorianus Banks): Contreras-Ramos 1998. Distribution COLOMBIA: [Cundina- marca?]: St. [San] Antonio, 2000 m. Corydalus sp. 3. Corydalus sp. 3 (near C. nubilus Erichson): Contreras-Ramos 1998. Distribution.—VENEZUELA: Bolivar: Ciudad Bolivar. Corydalus sp. 4. Corydalus sp. 4 (near C. tesselatus Stitz): Contreras-Ramos 1998. Distribution. ECUADOR: Sucumbios: El Reventador, [aprox. 900 m]. Corydalus sp. 5. Corydalus sp. 5 (near C. colombianus Con- treras-Ramos): Contreras-Ramos 1998. Distribution. ECUADOR: Pichincha: Palmeras. Genus Platyneuromus Weele 1909 53. Platyneuromus honduranus Navas. Platyneuromus soror hondurana Navas 1928b. Platyneuromus auritus Kimmins 1928. Platyneuromus honduranus Navas: Glorio- so and Flint 1984; Contreras-Ramos Sone 281 Distribution GUATEMALA: Alta Ver- apaz, El Petén, Iz4bal; HONDURAS: At- lantida, Cortés; MEXICO: Chiapas. 54. Platyneuromus reflexus Glorioso and Flint. Platyneuromus reflexus Glorioso and Flint 1984: Contreras-Ramos 1997. Distribution —GUATEMALA: Alta Ver- apaz; MEXICO: Chiapas. 55. Platyneuromus soror (Hagen). Corydalis soror Hagen 1861. Neuromus soror (Hagen): Davis 1903. Neuromus (Chloronia) soror (Hagen): Banks 1908. Platyneuromus soror (Hagen): Weele 1909, 1910; Stitz 1914; Penny 1977; Glorioso 1981; Glorioso and Flint 1984; Contrer- as-Ramos 1991b, 1997; Contreras-Ramos and Harris 1998. Doeringia christel Navas 1925b. Distribution COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia, Puntarenas, San José; MEXICO: Chiapas, Distrito Fed- eral, Hidalgo, Estado de México, Nuevo Le6n, Oaxaca, Puebla, Querétaro, San Luis Potosi, Tamaulipas, Veracruz; PANAMA: Chiriqui. Platyneuromus sp. Platyneuromus larval form A: Contreras- Ramos and Harris 1998. Distribution — MEXICO: Guerrero: 56 km NE Atoyac on road to Puerto del Gallo, 17.417°N, 100.217°W, 1372 m; Sinaloa: 4.83 km W Palmito. Family Sialidae (alderflies) Genus Protosialis Weele 1909 56. Protosialis bifasciata (Hagen). Sialis bifasciata Hagen 1861: Davis 1903. Protosialis bifasciata (Hagen): Weele 1910; Penny 1977. Distribution CUBA. 57. Protosialis bimaculata Banks. 282 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Protosialis bimaculata Banks 1920: Penny 1977S AIST 9821: Distribution.—BOLIVIA: La Paz. 58. Protosialis brasiliensis Navas. Protosialis brasiliensis Navas 1936: Penny 1997, WSS a 1982]: Distribution.—BRAZIL: Sao Paulo. 59. Protosialis chilensis (MacLachlan). Sialis chilensis MacLachlan 1870: Davis 1903; Flint 1973. Protosialis chilensis (MacLachlan): Weele 1910: Penny 1977. Distribution.—CHILE: Arauco, Concep- cion, Llanquihue, Malleco, O’ Higgins, Tal- ca, Valdivia. 60. Protosialis flammata Penny. Protosialis flamatta Penny 1981 [1982]. Distribution.—BRAZIL: Amazonas. 61. Protosialis flavicollis (Enderlein). Sialis flavicollis Enderlein 1910. Protosialis flavicollis (Enderlein): Penny 1977, 1981 [1982]. Distribution.—COLOMBIA: Tolima. 62. Protosialis mexicana (Banks). Sialis mexicana Banks 1901: Henry et al. 1992. Protosialis mexicana (Banks): Weele 1910; Penny 1977, E. D. Evans (in litt.); Con- treras-Ramos 199 1a. Distribution MEXICO: Chiapas, Vera- cruz; PANAMA. 63. Protosialis nubila Navas. Protosialis nubila Navas 1933: Penny 1977, 1981 [1982]: Distribution BRAZIL: Matto Grosso? ACKNOWLEDGMENTS Many of the references cited in this paper were obtained during my Ph.D. research at the Department of Entomology, University of Minnesota, St. Paul (UMSP). Thanks to Drs. Ralph W. Holzenthal and Philip J. Clausen for all their support while I was a student at UMSP. Thanks as well to the Di- reccion General de Asuntos del Personal Académico of UNAM for giving me a doc- toral fellowship, to Fernando Mufioz-Que- sada (UMSP) who kindly helped obtain several missing references, and to two anonymous reviewers for improving the quality of the manuscript. Several new rec- ords from Mexico were obtained through project KO22 funded by CONABIO. LITERATURE CITED Banks, N. 1892. A synopsis, catalogue, and bibliog- raphy of the neuropteroid insects of temperate North America. Transactions of the American En- tomological Society 19: 327-373. 1901. A list of neuropteroid insects from Mexico. Transactions of the American Entomo- logical Society 27: 361-371. . 1903a. Some new neuropteroid insects. Jour- nal of the New York Entomological Society 11: 236-243. 1903b. Neuropteroid insects from Arizona. Proceedings of the Entomological Society of Washington 5: 237-245. . 1907. Catalogue of the neuropteroid insects (except Odonata) of the United States. American Entomological Society, Philadelphia, Pennsylva- nia. 53 pp. . 1908. On the classification of the Corydalinae, with description of a new species. Proceedings of the Entomological Society of Washington 10: 27— 30. . 1914. Neuroptera and Trichoptera from Costa Rica. Entomological News 25: 149-150. . 1920. New neuropteroid insects. Bulletin of the Museum of Comparative Zoology, Harvard University 64: 299-362. . 1943. Neuroptera of northern South America. Boletin de Entomologia Venezolana 2: 59-66. . 1948. A new species of Corydalus (Neurop- tera). Psyche 55: 82-83. Blanchard, E. 1851. Neurdopteros, pp. 85-142 Jn Gay, C., Historia Fisica y Politica de Chile, Zoologia, Volumen 6. Burmeister, H. 1839. Handbuch der entomologie (Ber- lin), Vol. 2, parts 2—3: 397-1050. Cabrera, A. and A. Willink. 1980. Biogeografia de América Latina. Organizacién de los Estados Americanos, Monografias Cientificas, Ser. Biolo- gia 13: 1-122. Chandler, H. P. 1956. Megaloptera, pp. 229-233. In Usinger, R. L., ed., Aquatic Insects of California. University of California Press, Berkeley, Califor- nia. VOLUME 101, NUMBER 2 Contreras-Ramos, A. 1991a. Distribution of the Me- galoptera in Mexico. Bulletin of the North Amer- ican Benthological Society 8(1): 70. . 1991b. The first-instar larvae and pupae of Platyneuromus and Corydalus (Megaloptera: Cor- ydalidae: Corydalinae). Bulletin of the North American Benthological Society 8(1): 163. . 1993. Four unique dobsonflies (Megaloptera: Corydalidae: Corydalinae) from South America. Bulletin of the North American Benthological So- ciety 10(1): 130-131. . 1995a. New species of Chloronia from Ec- uador and Guatemala, with a key to the species in the genus (Megaloptera: Corydalidae). Journal of the North American Benthological Society 14: 108-114. . 1995b. A remarkable range extension for the fishfly genus Dysmicohermes (Megaloptera: Cor- ydalidae). Entomological News 106: 123-126. . 1997. Clave para la identificacio6n de los Me- galoptera (Neuropterida) de México. Dugesiana 4(2): 51-61. 1998. Systematics of the dobsonfly genus Corydalus (Megaloptera: Corydalidae). Thomas Say Publications, Entomological Society of Amer- ica. Contreras-Ramos, A. and S. C. Harris. 1998. The im- mature stages of Platyneuromus (Corydalidae), with a key to the genera of larval Megaloptera of Mexico. Journal of the North American Bentho- logical Society 17: 489-517. Davis, K. C. 1903. Aquatic Insects in New York State. Part 7. Sialididae [sic] of North and South Amer- ica. New York State Museum, Bulletin 68: 442— 486, 499, 2 pls. Enderlein, G. 1910. Eine neue Sialis aus Columbien. Stettiner entomologische Zeitung 1910: 380-381. Erichson, W. FE. 1848. Insekten, pp. 583-586 In M. R. Schomburgk’s Reisen in British-Guiana in den Jahren 1840-1884, Vol. 3. J. J. Weber, Leipzig. Evans, E. D. 1972. A study of the Megaloptera of the Pacific coastal region of the United States. Ph.D. dissertation, Oregon State University, Corvallis. Evans, E. D. and H. H. Neunzig. 1996. Megaloptera and aquatic Neuroptera, pp. 298-308. Jn Merritt, R. W. and K. W. Cummins, eds., An Introduction to the Aquatic Insects of North America, 3rd edi- tion. Kendall/Hunt Publishing Company, Du- buque, Iowa. Flint, O. S., Jr. 1965. The genus Neohermes (Mega- loptera: Corydalidae). Psyche 72: 255-263. 1970. The Megaloptera of Dominica. Pro- ceedings of the Entomological Society of Wash- ington 72: 240-242. . 1973. The Megaloptera of Chile (Neuroptera). Revista Chilena de Entomologia 7: 31—45. . 1983. Nothochauliodes penai, a new genus and species of Megaloptera from Chile (Neurop- 283 tera: Corydalidae). Entomological News 94: 15— G72 . 1991. On the identity of Chloronia bogatana [sic] Weele (Neuropterida: Megaloptera: Corydal- idae). Proceedings of the Entomological Society of Washington 93: 489-494, . 1992. A review of the genus Chloronia in Costa Rica, with the description of two new spe- cies (Neuropterida: Megaloptera: Corydalidae). Proceedings of the Biological Society of Wash- ington 105: 801-809. Geijskes, D. C. 1984. Notes on Megaloptera from the Guyanas, S. Am., pp. 79-84. In Gepp, J., H. As- pock, and H. Ho6lzel, eds., Progress in World’s Neuropterology; Proceedings of the Ist Interna- tional Symposium on Neuropterology. Graz, Aus- tria. Glorioso, M. J. 1981. Systematics of the dobsonfly subfamily Corydalinae (Megaloptera: Corydali- dae). Systematic Entomology 6: 253-290. Glorioso, M. J. and O. S. Flint, Jr. 1984. A review of the genus Platyneuromus (Insecta: Neuroptera: Corydalidae). Proceedings of the Biological So- ciety of Washington 97: 601-614. Hagen, H. 1861. Synopsis of the Neuroptera of North America with a list of the South American spe- cies. Smithsonian Miscellaneous Collections 4(1): xx + 1-347. Hazard, E. I. 1960. A revision of the genera Chauli- odes and Nigronia (Megaloptera: Corydalidae). M.S. thesis. The Ohio State University. Henry, C. S., N. D. Penny, and P. A. Adams. 1992. The neuropteroid orders of Central America (Neu- roptera and Megaloptera), pp. 432—458. /n Quin- tero, D., and A. Aiello, eds., Insects of Panama and Mesoamerica. Oxford University Press, Ox- ford. Herrmann, S. J. and H. L. Davis. 1991. Distribution records of Corydalus cornutus (Megaloptera: Cor- ydalidae) in Colorado. Entomological News 102: 25-30. Kimmins, D. E. 1928. New and little known Neurop- tera of Central America. Eos 4: 363-370. . 1954. A new genus and some new species of the Chauliodini (Megaloptera). Bulletin of the British Museum (Natural History), Entomology 3: 417-444. . 1970. A list of the type-specimens of Plecop- tera and Megaloptera in the British Museum (Nat- ural History). Bulletin of the British Museum (Natural History), Entomology 24(8): 337-361. Latreille, P, A. 1802. Histoire Naturelle, Genérale et Particuliere, des Crustacés et des Insectes. Tome troisiéme (Vol. 3). FE Dufart, Paris. MacLachlan, R. 1866. Description of a new neurop- terous insect belonging to the genus Corydalis, Latreille. Journal of Entomology (London) 2: 499-500, 1 pl. . 1868. New genera and species, &c., of neu- ropterous insects; and a revision of Mr. E Walker’s British Museum catalogue of Neuroptera, part 1 (1853), as far as the end of the genus Myrmeleon. Journal of the Linnean Society (Zoology) 9: 230— Zolealeple . 1869. Considerations on the neuropterous ge- nus Chauliodes and its allies; with notes and de- scriptions. The Annals and Magazine of Natural History 4(4): 35—46. . 1870. On the occurrence of the neuropterous genus Sialis in Chili. Entomologist’s Monthly Magazine 7: 145-146. Navas, L. 1915. Neurdpteros sudamericanos. Segunda serie. Brotéria (Série Zoologica) 13: 5—13. . 1920a. Insectos sudamericanos (2a serie). An- ales de la Sociedad Cientifica Argentina 90: 44— Sil; . 1920b. Insectos de América. Boletin de la So- ciedad Entomologica de Espana 3: 90—99. . 1921. Insectos americanos nuevos 0 criticos. 1921. Brotéria (Série Zoologica) 19: 113-124. . 1925a. Insectos exd6ticos nuevos 0 poco con- ocidos, Segunda Serie. Memorias de la Real Ac- ademia de Ciencias y Artes de Barcelona, serie 3, 19: 181-200. 1925b. Neuropteren, Megalopteren, Plecop- teren und Trichopteren aus dem Deutschen Ento- mologischen Institiit (Berlin-Dahlem). I. serie.— Entomologische Mitteilungen 14: 205-212. . 1927. Insectos del Museo de Paris (4a serie). Brotéria (Série Zoold6gica) 24: 5-33. . 1928a. Insectos del Museo de Estocolmo. Re- vista de la Real Academia de Ciencias Exactas, Fisicas y Naturales de Madrid 24: 1-12. . 1928b. Insectos del Museo de Hamburgo (pri- mera serie). Boletin de la Sociedad Entomolégica de Espana 11: 59-67. . 1929. Insectos de la Argentina (quinta serie). Revista de la Sociedad Entomologica Argentina 2: 219-225. . 1933. Décadas de insectos nuevos. Brotéria 2: 34-44, 101-110. . 1934a. Insectos suramericanos (octava serie). Revista de la Academia de Ciencias de Madrid 31: 9-28. . 1934b. Insectos del Museo de Hamburgo (2a PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON serie). Memorias de la Academia de Ciencias y Artes de Barcelona 23: 499-508. . 1935. Insectos suramericanos (décima serie). Revista de la Academia de Ciencias Exactas, Fis- ico-Quimicas y Naturales de Madrid 32: 360-375. . 1936. Insectos del Brasil (Sa. serie). Revista do Museu Paulista 20: 722-734. New, T. R. and G. Theischinger. 1993. Megaloptera (Alderflies, Dobsonflies). Handbuch der Zoologie, Vol. 4 (Part 33). Walter de Gruyter, Berlin. Penny, N. D. 1977. Lista de Megaloptera, Neuroptera e Raphidioptera do México, América Central, il- has Caraibas e América do Sul. Acta Amazonica 7(4): Suplemento, 61 pp. . 1981[1982]. Neuroptera of the Amazon Basin. Part 4, Sialidae. Acta Amazonica 11: 843-846. . 1982. Neuroptera of the Amazon Basin. Part 7, Corydalidae. Acta Amazonica 12: 825-837. Penny, N. D., P. A. Adams, and L. A. Stange. 1997. Species catalog of the Neuroptera, Megaloptera, and Raphidioptera of America north of Mexico. Proceedings of the California Academy of Scienc- es 50: 39-114. Penny, N. D. and O. S. Flint, Jr. 1982. A revision of the genus Chloronia (Neuroptera: Corydalidae). Smithsonian Contributions to Zoology 348: 1—27. Rambur, J. P. 1842. Histoire naturelle des insectes. Névropteres. Librairie Encyclopédique de Roret, Paris. Ross, H. H. 1937. Studies of Nearctic aquatic insects, I. Nearctic alder flies of the genus Sialis. Ilinois Natural History Survey Bulletin 21: 57-78. Stutz, H. 1914. Sialiden der Sammlung des Berliner Museums. Sitzungsberich der Gesellschaft natur- forschender Freunde zu Berlin 5: 191—205, 2 pls. Walker, FE 1853. List of the specimens of neuropterous insects in the collection of the British Museum. Part II (Sialidae-Nemopterides). Edward Newman, London, pp. 193-476. . 1858. Characters of undescribed Neuroptera in the collection of W. W. Saunders. Transactions of the Royal Entomological Society of London, new series 5: 176-199. Weele, H. W., van der. 1909. New genera and species of Megaloptera Latr. Notes from the Leyden Mu- seum 30: 249-253. . 1910. Megaloptera (Latreille), monographic revision, pp. 1-93 + 4 pls. /n Collections Zoolo- giques du Baron Edm. de Selys Longchamps Fasc. V (Premiere partie), Bruxelles. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 285-289 IDENTITY OF SYRISTA SPECIOSA MOCSARY AND NOTES ON THE GENUS UROSYRISTA MAA (HYMENOPTERA: CEPHIDAEF) DAVID R. SMITH Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, % National Museum of Natural History, MRC 168, Washington, DC 20560-0168, U.S.A. (e-mail: dsmith@sel.barc.usda.gov) Abstract.—The holotype of Syrista speciosa Mocsary, described from Vietnam, is re- described and illustrated and confirmed as belonging to the genus Urosyrista Maa. Pos- sible characters are presented to separate it from other species of this southeastern Asian genus. Key Words: In the treatments of world Cephidae (Benson 1946, Muche 1981) and Asian Ce- phidae (Maa 1944, 1949), the species de- scribed as Syrista speciosa by Mocsary (1904), an unusually large and colorful ce- phid from ‘“‘Tonkin,”’ has never been stud- ied and confidently placed. Benson (1946) placed S. speciosa in his new genus Ce- phalocephus, qualifying it with the state- ment that “‘it seems almost certain’’ that it belongs to this genus. Maa (1949) synon- ymized Cephalocephus with Urosyrista which he had described earlier (Maa 1944). Although Maa (1949) did not see Cephal- ocephus xanthus Benson, the type species of Cephalocephus, both generic descrip- tions are almost identical and their synon- ymy cannot be disputed. Maa (1949) did not mention S. speciosa or put it in his key to species of Urosyrista. Muche (1981), who based much of his work on the liter- ature, placed S. speciosa in Urosyrista but did not include it in his key to Urosyrista species, which was taken directly from Maa (1949). Here, I redescribe, illustrate, and confirm the generic placement of Syrista speciosa based on examination of the holotype. Cephidae, Urosyrista, stem sawflies Urosyrista speciosa (Mocsary) (Figs. 1-9) Syrista speciosa Mocsary 1904: 496. Fe- male. “Tonkin: Montes Mauson, in alti- tudine 2—3,000 pedum a H. Fruhstorfer detecta. (Mus. Hung.).”’ Cephalocephus speciosa: Benson 1946: 100. Urosyrista speciosa: Muche 1981: 265. Description.—Length, 16 mm. Head and body smooth, shining, without punctures. Color: Yellowish with black markings (as shown in Figs. 1—9); antenna pale yellowish ventrally, black dorsally, with basal 3 seg- ments black. Wings hyaline, apices slightly blackish; most of stigma black, and veins dark brown, costa and margins of stigma yellowish. Head: In dorsal view, enlarged and swollen behind eyes, elongated, 1.2 broader than long, distance behind eyes 1.3X eye length; distance from front ocellus to hind margin of head 3X distance from front ocellus to antennal sockets. Antennal- tentorial distance 2 distance between an- tennae. Antenna 30-segmented. Left man- dible bidentate, subapical tooth slightly lon- ger and stouter than apical tooth, without Figs. 1-3. Urosyrista PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON speciosa. 1, Lateral view. 2, Dorsal view. 3, Dorsal view of thorax. VOLUME 101, NUMBER 2 287 eis Figs. 4-6. Urosyrista speciosa. 4, Head, front view. 5, Head, lateral view. 6, Head, dorsal view. 288 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-9. Urosyrista speciosa. 7, Thorax, lateral view. 8, Apex of abdomen and sheath, lateral view. 9, Apex of abdomen and sheath, dorsal view. intermediary tooth (Fig. 4). Sixth segment cell. Hindtibia with 2 preapical spines. Mid- of maxillary palpus originating from near tibia with one preapical spine. Hindbasitar- apex of Sth segment; segment 4 about 1.5X sus subequal in length to remaining tarsal length of segment 6. Thorax: Forewing segments combined. Tarsal claws bifid, in- with anal crossvein. Hindwing with cubital ner tooth about as long as outer tooth, with- VOLUME 101, NUMBER 2 out basal lobe. Abdomen: Cercus less than half length of sawsheath (Figs. 8, 9). Sheath as in Figs. 8, 9; in dorsal view, thick and parallel sided; length about .6x length of basal plate and slightly less than half as long as hind basitarsus. Holotype.—Female, labeled ‘*Tonkin, Montes Manson, April, Mai, 2—3,000’, H. Fruhstorfer,”’ ““Typus 1904 Syrista speciosa Mocs.” (red label). The third label on the pin is a blank red label. In the Hungarian Natural History Museum, Budapest. Discussion.—Almost all characters of Syrista speciosa are consistent with Uro- syrista, and placement in this genus is cor- rect according to generic definitions and keys by Maa (1944, 1949), Benson (1946), and Muche (1981). Some minor differences are the lack of a basal swelling on the tarsal claws (according to Benson 1946 the claws have a small basal swelling) and the sub- apical tooth of the left mandible which, al- though longer than the apical tooth, is not as large in relation to the apical tooth as figured by Benson (1946, fig. 13). Unique characters for Urosyrista in the Cephidae are the enlarged head behind the eyes (Figs. 5, 6), the lack of an intermediate tooth and large innter tooth of the left mandible (Fig. 4), the apical segment of the maxillary pal- pus emerging from near the apex of the fifth segment, and the bifid tarsal claws, lacking a basal lobe. Urosyrista is known only from south- eastern Asia, and three species are currently included: U. speciosa (Mocsary) from Vi- etnam; U. montana (Maa) (= Cephaloce- phus xanthus Benson according to Maa 1949) from China and Burma; and U. men- cioyana Maa from China. Maa (1949) con- sidered three color forms of U. mencioyana, the typical form, var. unicolor Maa, and var. xanthobalteata Maa. All three are separated by the amount of black on the pronotum. The host plant is known only for U. men- cloyana; specimens were reared from Acan- thopanax trifoliatus (Lour.) Merr. (Arali- aceae) (Maa 1944). Urosyrista speciosa appears to be most 289 similar to U. montana because they share the thick, parallel sided sheath in dorsal view. The sheath of U. mencioyana is slen- der and gradually tapering toward its apex. I hesitate to use coloration. In species as these with numerous black markings, vari- ation in the amount of black can be exten- sive. Maa (1949) has alluded to this by sep- arating several color forms of U. mencioy- ana. However, as it may be useful, U. spe- ciosa 18 more extensively yellow than the other two species and the antennae are black dorsally, pale whitish ventrally, with the basal three segment entirely black. Uro- syrista montana has the antennae apically dull brown to black, basally distinctly paler, and the mesepisternum mostly black with a median yellow band. Urosyrista mencioy- ana has the antennae apically yellow, ba- sally distinctly darker, and the head and tho- rax mostly black. ACKNOWLEDGMENTS I thank L. Zombori, Hungarian Museum of Natural History, Budapest, for loan of the holotype. Terry Nuhn and Cathy An- derson, Museum Technicians, Systematic Entomology Laboratory, USDA, took the photographs of the holotype and arranged the plates. I thank the following for review of the manuscript: N. M. Schiff, USDA, Forest Service, Stoneville, MS, and M.E. Schauff and S.W. Lingafelter, Systematic Entomology Laboratory, USDA, Washing- ton, DC. LITERATURE CITED Benson, R. B. 1946. Classification of the Cephidae (Hymenoptera Symphyta). Transactions of the Royal Entomological Society of London 96: 89— 108. Maa, T. C. 1944. Novelties of Chinese Hymenoptera, Chalastogastra. Biological Bulletin of the Fukien Christian University 4: 33-60. Maa, T. C. 1949. A synopsis of Chinese sawflies of the superfamily Cephoidea (Hymenoptera). Chi- nese Journal of Zoology 3: 17-42. Mocsary, A. 1904. Siricidarum species quinque novae. Annales Musei Nationalis Hungarici 2: 496—498. Muche, H. 1981. Die Cephidae der Erde (Hym., Ce- phidae). Deutsche Entomologische Zeitschrift, N.E 28: 239-295. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 290-294 FRANKLINIELLA ZUCCHINI (THYSANOPTERA: THRIPIDAE), A NEW SPECIES AND VECTOR OF TOSPOVIRUS IN BRAZIL SUEO NAKAHARA AND RENATA C. MONTEIRO (SN) Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, 10300 Baltimore Avenue, Beltsville, Maryland 20705-2350, U.S.A. (e-mail: snakahar@ sel.barc.usda.gov); (RCM) Departamento de Entomologia, ESALQ, Universidade de Sao Paulo, 13418—900 Piracicaba, Sao Paulo, Brazil Abstract.—Frankliniella zucchini, new species, is described. It is a vector of a tos- povirus that causes zucchini lethal chlorotic disease of Cucurbita pepo L. cv. Caserta in Sao Paulo State, Brazil. Key Words: ZLCV, Brazil A serious disease of zucchini squash, Cu- ¢urbita pepo i. .cv. Caserta, currently known as zucchini lethal chlorotic disease (ZLC), was discovered during 1995 in ex- perimental fields at Piracicaba, Sao Paulo State, Brazil (Rezende et al. 1997). This disease is caused by a species of Tospovi- rus, Zucchini lethal chlorotic virus (ZLCV) (Pozzer et al. 1996). Tospoviruses can be transmitted only by thrips adults and larvae. The disease apparently was present sporad- ically in the state prior to 1991 when many zucchini plants were observed with symp- toms of ZLC in Campinas county. Since then, symptoms of ZLC were observed more frequently on zucchini squash and watermelon. Frankliniella zucchini, new species, described here was the predomi- nant thrips species collected from foliage and flowers of infected plants in Piracicaba. In preliminary transmission tests, this thrips was found to be a vector of ZLCV to zuc- chini seedlings (Rezende 1998, personal communication). Two polyphagous Frankliniella species are vectors of “‘tomato spotted wilt virus” (TSWV) and “‘tomato chlorotic spot virus”’ Frankliniella zucchini n. sp., Thysanoptera, Thripidae, zucchini, vector, (TCSV) in Brazil (Wijkamp et al. 1995). Frankliniella schultzei (Trybom) dark form is an efficient vector of TSWV and TCSV that damages tomato crops. Although the yellow form of this species was previously not considered to be a vector, it is an inef- ficient vector of TSWV according to Wijk- amp et al. (1995). The recently established F. occidentalis (Pergande) is a pest of var- ious agricultural crops and also a vector. Frankliniella zucchini is known to vector only ZLCV and is the fifth Frankliniella species to be implicated in the transmission of tospoviruses. The other two are F. fusca (Hinds) and F. intonsa (Trybom). For measurements and counts, the values for the holotype are given first and are fol- lowed by values for the paratypes in paren- theses. If parentheses are absent the values are either for the holotype or for all mea- sured specimens. Frankliniella zucchini Nakahara and Monteiro, new species (Figs. 1-6) Female (macropterous).—Body generally yellow; forewing pale yellow, legs paler VOLUME 101, NUMBER 2 Figs. 1-6. Frankliniella zucchini. Female. 1, Right antenna. 2, Head (a, ocellar seta Il; b, POiv seta). 3, Pronotum (a, anteromarginal seta; b, anteroangular setae; c, outer posteroangular seta; d, inner posteroangular seta, e. posteromarginal seta II, f, submarginal seta). 4, Metanotum. 5, Abdominal tergite VIII. 6, Abdominal tergite IX (a, Blseta; b, B2 seta; c, B3 seta). Scale = 0.1 mm. 292 yellow than body; body and forewing setae brown. Antennal segment I as pale as head; segment II pale yellow in proximal 2, shad- ed light brown distally; segment III pale yellow in proximal % including pedicel, distally brown; segment IV pale yellow in proximal 4%, brown distally and in pedicel; segment V pale yellow in proximal %, dis- tally brown; segments VI to VIII brown. Antenna (Fig. 1): More than twice as long as head; pedicel of segment III with slight angulation, segments III and IV each with v-shaped forked sense cones, 22—27 w long; segment III distad of subapical setae slightly converging to apex, about 4 length of segment, segment IV constricted into a neck in distal part; segment VI slightly ped- icellate at base, inner sense cone 32-35 w long, extending distally at least to apex of segment VII. Head (Fig. 2): Slightly shorter than pro- notum, about 1.5 times wider than long, cheeks rather straight, compound eyes al- most twice as long as occiput; interocellar area without sculpturing, caudad of com- pound eyes with transverse striae, those more posterior spaced farther apart. Com- pound eyes each with 4 pigmented facets in 1—2,3,5 pattern (see Nakahara 1988). Di- ameter of fore ocellus 15—17 wp. Ocellar se- tae I short, about 12 w long, just anterior of fore ocellus; ocellar setae II short, laterad of fore ocellus and just mesad of inner mar- gin of compound eye, about 12 w long; ocellar setae III well developed (Fig. 2a), between anterior part of posterior ocelli, separated by about diameter of fore ocellus, 45(37—48) w long. Postocular (PO) setae 5 pairs; POi seta absent; POii seta 12-15 wp long, caudad of posterior ocellus; next lat- erad POii seta 10-12 ww long; POijv seta longest (Fig. 2b), 32(24—33) wp long; POv and POvi setae laterad of and shorter than seta iv but longer than setae ii and iii. Mouthcone conical, longer than head, distal 7, less convergent than basal ¥, extending to posterior margin of prosternum; mandi- ble 124—136 wp long. Pronotum (Fig. 3): Rectangular, broader PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON than long, sculptured with irregularly spaced transverse striae and sparsely anas- tomosing. Median discal area with 3-5 se- tae 10—12 p long; 2 irregular rows of discal setae in posterior %, 7—9 setae in anterior row including a pair of longer submarginal setae (Fig. 3f), 20=24 i, anterior of posteroangular setae; posterior row normal- ly with 4 setae; anteromarginal setae 37— 42(42—48) pw long (Fig. 3a), 2 short setae, 10-12 pw long, between anteromarginal se- tae; anteroangular setae 48(48—-59) wp long (Fig. 3b); posteroangular seta inner pair 59 (57-67) w long (Fig. 3d), outer pair 57— 59(59—64) w long (Fig. 3c), about % as long as notum; posteromarginal seta II 32(32-— 40) pw long (Fig. 3e). Mesonotum: Subtra- pezoidal, anterior angulate area smooth with pair of campaniform sensilla, median and posterior parts transversely sculptured, laterally with striae oriented longitudinally; 2 pairs of short setae on or near posterior margin, inner pair 17—20 p long, outer pair slightly stouter, 20—24 wp long. Metanotum (Fig. 4): Reticulated with most reticles lon- ger than wide and oriented longitudinally, reticles in anterior medial area more polyg- onal and wider than other reticles; median setae 55—59(54—-59) w long, thicker than lateral setae 35—37(35—37) pw long; 2 cam- paniform sensilla in posterior % of notum. Forewing: Rather straight, apex pointed; fringe cilia wavy; costa with 20—24(19—20) cilia, 23—24(22—24) setae, setae at mid- length 40(37—42) p long, shorter than width of forewing; forevein with 19—21(18—20) setae, hindvein with 16—17(14—15) setae; scale with 4 marginal and | discal setae. Abdomen: Tergites sculptured anteriorly, and laterally of median setae and campan- iform sensilla; median setae short, on VII 17-20 pw long; short ctenidia on tergite IV; posterior margins of intermediate tergites with series of low, truncate lobes, with a few small teeth laterally; tergite VIII with posteromarginal comb complete with 14(13-17) microtrichia, longest 17-20 pw long (Fig. 5); tergite [IX with microtrichia on anastomosing striae, posterior pair of VOLUME 101, NUMBER 2 campaniform sensilla near BI setae (Fig. 6); tergite X subequal to IX, dorsal split almost complete on X. Ovipositor well de- veloped, 183(222—235) w long. Measurements: Female holotype and (paratypes) in p. Body length from anterior of eye 1176(1161-1221), distended 1423(1408—-1568). Antenna: Total length 270(265—280); length and width of segment I 24(24), 27(27-—30); I 37(35-37), 24(27); III 50(48-50), 22(22); IV 45(42-48), 20(20-21); V 37(37-40), 17(17(20); VI §2(52-54), 18(17-—20); VII 10(10-11), 7(7); VU 15(15), 5(5—6). Head length from anterior of compound eye 100(96—106), width at compound eyes 151(148—156), width at cheeks 156(143—158); length of compound eye 62(62—69), width 45; length of occiput posterior of compound eye 37(35-—37). Pronotal length 124(126—133), width 190(178—190). Forewing length 729(679-729), width at midlength 57(52-— 57). Length of abdominal tergite [IX 62(64— 67), length of Bl setae 104—109(96—109) (Fig. 6a), B2 setae 101-106 (104-109) (Fig. 6b), B3 seta 100(100) (Fig. 6c); length of tergite X 64(64—67), length of B1 seta 109(96—-111), B2 seta 98(94). Male (macropterous).—Smaller than fe- male, otherwise similar in color and most anatomical structures. Body length 1,000— 1,050 wp. Antennal length 220—246 wp. Ab- dominal tergite VIII with complete postero- marginal comb with 13-14 long, slender microtrichia; sternites III-VII each with transversve glandular area with anterior and posterior margins concave, on III 52—62 wp wide, 15—17 pw long, on VII 40—54 pw wide, 12-15 pw long, 0.30—0.38 as wide as ster- nite; sternite VIII with posteromarginal mi- crotrichia. Type material—Holotype 2 and 18 @ and 2 ¢ paratypes: Brasil, Sao Paulo, Pir- acicaba, Cucurbita pepo L, cv. Caserta, 14- VII-95. R.C. Monteiro. Holotype and 10 paratypes deposited in Departamento de Entomologia, ESALQ, Universidade de Sao Paulo, Piracicaba, Brazil, 10 paratypes in the National Museum of Natural History, 298 Smithsonian Institution, Washington D.C, and 3 paratypes in The Natural History Mu- seum, London, United Kingdom. Etymology.—The species is named after the common name of the host, ‘‘zucchini,”’ and is a noun in apposition. Distribution.—Known only from Sao Paulo State, Brazil. Collected from.—Cucurbita pepo L. cv. Caserta (zucchini). Comments.—Frankliniella gemina Bag- nall and F. rodeos Moulton in Brazil are similar to F. zucchini in color and most an- atomical characters. Frankliniella zucchini lacks POi seta and has ocellar setae III po- sitioned between the anterior part of pos- terior ocelli and separated by about the width of the fore ocellus; whereas POi setae are present in the other two species and ocellar setae III are aligned with the ante- rior margin of posterior ocelli or slightly anterior and are farther apart. Most Frankliniella species have three pairs of short postocular (PO) setae mesad or anteromesad of the fourth or longest pair of PO setae which is positioned posterior of the compound eye (Fig. 2). When only two pairs of short PO setae are present, the PO1 seta is normally absent. The normal posi- tion of POi seta is caudad of and usually slightly mesad of the inner margin of the posterior ocelli. ACKNOWLEDGMENTS We thank the following colleagues for their contributions to this article: Laurence Mound, CSIRO, Canberra, Australia, rec- ognized that this thrips was new to science and recommended that it be described; and J. A. M. Rezende, ESALQ/USP, Piracicaba, Sao Paulo, Brazil, provided information about ZLCV and the vector. For reviewing the manuscript and useful suggestions, we thank Robert Foottit, Agriculture and Agri- Food Canada, Ottawa, and Stuart Mc- Kamey and Douglass Miller, Systematic Entomology Laboratory, USDA, Washing- ton D.C. and Beltsville, Maryland respec- tively. 294 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Resende, A. C. De Avila, and S. M. Scagliusi. 1997. Incid , biological and logical char- Nakahara, S. 1988. Preliminary study of facetal pig- Fs NGS COS aan emanates ee fond eat mentation in the compound eyes of Terebrantia (Thysanoptera). Acta Phytopathologica et Ento- mologica Hungarica 23(3—4): 321-329. 7 ; Pozzer, L., R. O. Resende, M. I. Bezerra, T. Nagata, Wijkamp, I., N. Almarza and D. Peters. 1995. Median acteristics of a tospovirus infecting experimental fields of zucchini in Sao Paulo State, Brazil. Fi- topatologia brasileira 22: 92-95. M. I. Lima, E W. Kitajima, and A. C. De Avila. latent period and transmission of tospoviruses 1996. Zucchini lethal chlorotic virus (ZLCV), a vectored by thrips. pp. 153-156. Jn Parker, B. L., proposed new species in Tospovirus genus. Fito- M. Skinner and T. Lewis, eds., Thrips Biology and patologia Brasileira 21 (Suplemento): 432. Mangement. Plenum Press, New York and Lon- Rezende, J-A.M., S. R. Galleti, L. Pozzer, R. de O. don. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 295-299 THE LARVA OF CHALCIDOMORPHINA AURATA ENDERLEIN 1914 (DIPTERA: STRATIOMYIDAE) FROM “ILHA DE MARAMBATA,” RIO DE JANEIRO, BRAZIL JOSE R. PUJOL-LUZ AND ROBERTO DE XEREZ Departamento de Biologia Animal, Instituto de Biologia da Universidade Federal do Rio de Janeiro, Seropédica, Rio de Janeiro, 23890-000, Brazil (e-mail: pujol-luz@uol. com.br; rdexerez @ uol.com.br) Abstract.—The larva of Chalcidomorphina aurata is described for the first time, based on six larvae and the puparium. Larvae were collected under the bark of fallen trees in a tropical rain forest at [ha da Marambaia, State of Rio de Janeiro, Brazil (23°04'S, 43°53'W, approximately 42 km’). Some biological notes are also presented. Key Words: vae, tropical rain forest The pachygastrine genus Chalcidomor- phina Enderlein, 1914, with four species, is widespread in the Neotropics, from Mexico to Brazil: Chalcidomorphina aurata Ender- lein, 1914 (Mexico, Panama, Colombia, Peru and Brazil); C. plana James, 1967 (Dominica); C. terataspis James, 1974 (Peru); and C. argentea McFadden, 1980 in James et al., 1980 (Mexico) (James 1973, James et al. 1980). James (1974) recognized Chalcidomor- phina based on the following characters: (1) antenna elongate, with long scape and fla- gellum, and (2) scutellum projected into an elongate, spur-like process (Fig. 1). James (1974) segregated Chalcidomorphina and Dactylacantha Lindner, 1964, from the re- lated genus Dactylodeictes Kertész, 1914, based on wing venation. Pachygastrinae larvae from the Neotrop- ics have never been described. McFadden (1967) and James (1981) furnished the last records of the known North American lar- vae of this subfamily. They studied Nearc- tic representatives of some genera that are Stratiomyidae, Pachygastrinae, Chalcidomorphina aurata, soldier-flies, lar- widespread over much of the Americas (e.g., Eidalimus Kertész, 1914 [=Eucyni- pimorpha Malloch, 1915; =Eupachygaster authors, part, not Kertész, 1911]; Gowdey- ana Curran, 1928 [=Eupachygaster au- thors, part, not Kertész, 1911; =Paraeidal- imus Lindner, 1964] and Zabrachia Co- quillett, 1901) (James et al. 1980). Here we describe the larva of Chalcido- morphina aurata for the first time, based on six larvae and the puparium. Some of the larval features employed in this work are the same used by McFadden (1967) and James (1981) to describe other genera. Chalcidomorphina larvae were collected under the bark of fallen trees in a tropical rain forest at Ilha da Marambaia, State of Rio de Janeiro, Brazil, (23°04’'S, 43°53'W, approximately 42 km’) outside and inside the forest behind a dam (a restricted area of the Brazilian Navy). The junior author collected approximate- ly 35 to 50 larvae of different instars at each site belonging to two genera, Chalci- domorphina and Cyphomyia Wiedemann, 1819 (Clitellariinae). Field and laboratory 296 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-2. view; left in ventral view). Scale line = 2.0 mm. observations suggest that larvae of C. au- rata feed on microorganisms occurring in the moist areas beneath the bark of trees. The trees where the larvae were found were not identified. We also confirm McFadden’s (1967) statement concerning the gregarious behavior of larvae of pachygastrine soldier- flies. The larvae of C. aurata were reared in semi-natural conditions, segregated by size classes, and put on individual petri dishes having as substrate the same moist sub- stance present where the larvae were living. A single female was collected flying over a Chalcidomorphina aurata. 1, Female habitus (all pilosity is omitted). 2, Larva (right in dorsal tree. We believe that oviposition occurs in the moist substrate, through crevices in the bark, because a large number of the small- est larvae of C. aurata were found there. The females that emerged are extremely similar to those described by Lindner (1951), James (1974) and James et al. (1980), but showed a chromatic pattern of the eyes not described by these authors. The eyes in living insects are brownish with a greenish ‘‘9-shaped”’ pattern in lateral view Rigel): The terminology adopted in the descrip- tions follows James (1981) and Rozkosny VOLUME 101, NUMBER 2 297 Figs. 3-9. Larva of Chalcidomorphina aurata. 3, Head, dorsal view. 4, Head, ventral view. 5, Right spiracle, lateral view. 6, Abdominal segment 6, dorsal view. 7, Abdominal segment 6, ventral view. 8, Abdominal segment 8, dorsal view. 9, Abdominal segment 8, ventral view. Scale lines = 0.5 mm (Figs. 3—5), = 1.0 mm (Figs. 6— 9). Abbreviations: ant = antenna; cyb = cylindrical brushes; Ibr = labrum. 298 and Kovac (1994). The specimens upon which this study is based are in the Cole¢ao Entomologica Costa Lima (CECL), Insti- tuto de Biologia, Universidade Federal Ru- ral do Rio de Janeiro (UFRRJ), Seropédica, Rio de Janeiro, RJ, Brasil. Chalcidomorphina aurata Enderlein, 1914 (Figs. 1—9) Chalcidomorphina aurata: Enderlein 1914 (original description); Lindner 1951 (suppl. descr., figs.); James 1973 (cata- logue); James 1974 (revision, figs.); James, McFadden, and Woodley 1980 (suppl. descr., key to females, notes on the males, figs.). Distribution.—Neotropical: Mexico, Panama, Colombia, Peru, Brazil. Larva (and puparium).—Length 5.0 to 5.3 mm, flattened dorsoventrally, lateral margins of body segments strongly arched. Cuticle with usual mosaic appearance, some cells forming characteristic patches and plaques on abdominal segments 6 and 8. Chromatic pattern yellowish brown, with some dark punctuations (Fig. 2). Head: Subconical, moderately flat; mandibular-maxillary complex with well developed, cylindrical brushes almost as long as labrum, in dorsal view (Fig. 3); la- brum triangular. Antenna short, rising at an- terior part of head. Eyes prominent, round- ed, arising at the posterior part of the head. Two pairs of lateral setae, one pair of cly- peofrontal setae and one pair of dorsolateral setae inserted above eyes; three pairs of ventrolateral setae and three pairs of spine- like ventral setae (Figs. 3—4). Thorax: First segment shorter than oth- ers. Spiracle prominent and V-shaped, with a small anterolateral spiracular seta, in lat- eral view with two dorsal setae and one ventral seta (Fig. 5). First segment with two rows of setae in dorsal view: two pairs of anterodorsal setae and three pairs of dorsal setae. In ventral view, two pairs of ventral setae and one pair of ventrolateral setae near spiracle. Second and third segments PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with one row of setae with four pairs of dorsal setae and three pairs of ventral setae (Fig. 2). Abdomen: Segments 1—7 similar in shape to thoracic segments (Fig. 2), with a row of five pairs of dorsal setae and four pair of ventral setae (Fig. 6—7); ventrome- dial line of segment 6 with an elliptical ster- nal patch (Fig. 7); segment 8 rounded, with three pairs of conspicuous plaques along dorsomedial line between a pair of pennate dorsocentral setae (Fig. 8), four pairs of lat- eral setae, apical pair shorter than others; opening of spiracular chamber with a fringe of small setae, anal slit on ventral side emarginate with a long fringe of setae on each side (Fig. 9). Material examined.—Brazil, Rio de Ja- neiro, Ilha da Marambaia, 17.11.1998 (R. Xerez col.), 4 females (emerged: 04.11.1998; 2.1V.1998 and 9.IV.1998) and 2 larvae (last instar). Comments.—Chalcidomorphina larvae share some features with the genera keyed by James (1981). The larva keys to the sec- ond half of couplet 21. Then, the features are distributed in several couplets. Couplet 21 segregates two groups: first half: [Berk- shiria + Neopachygaster] and second half: [Gowdeyana + Eidalimus + Pachygaster + Zabrachia|. However, Chalcidomorphina also shares a feature with Neopachygaster (second half of couplet 22, three pairs of conspicuous plaques along dorsomedial line of abdominal segment 8) and differs in the same couplet by the shape of sternal patch of abdominal segment 6 (oval in Neopach- ygaster). ACKNOWLEDGMENTS We thank to Dr. Nelson Papavero, Dr. Francisco Racca Filho, Dr. Hélio Ricardo da Silva, Mary S. Linn, and Christopher Sheil for critical reviews of the manuscript. This research was financed by the Conselho Nacional de Desenvolvimento Cientifico e Tecnol6gico—CNPq (Grant 300265/96-4). VOLUME 101, NUMBER 2 299 LITERATURE CITED Alpine, J. F, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds. James, M. T. 1973. A catalogue of the Diptera of the Manual of Nearctic Diptera. Vol. 1. Agriculture Americas south of the United States. 26. Family Canada, Ottawa. vi + 674 pp. Stratiomyidae. Museu de Zoologia da Universi- James, M. T., M. W. McFadden, and N. E. Woodley. dade de Sao Paulo. 95 pp. 1980. The Pachygastrinae (Diptera, Stratiomyi- ——. 1974. The pachygastrine genera Dactylodeic- dae) of Middle America. Melanderia 34: 1—36. tes, Chalcidomorphina, and Thopomyia in South — Lindner, E. 1951. Vierter Beitrag zur Kenntnis der sue- America (Diptera, Stratiomyidae). Melanderia 14: damerikanischen Stratiomyiidenfauna (Dipt.). Re- 23-32. vista de Entomologia 22 (1-3). —.. 1965. Stratiomyidae, pp. 299-319. In Stone, | McFadden, M. W. 1967. Soldier Fly larvae in America A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, North of Mexico. Proceedings of the United States and J. R. Coulson, eds. A Catalog of Diptera of National Museum 121 (3569): 1-72. America North of Mexico. United States Depart- RozkoSny, R. and D. Kovac. 1994. Adults and larvae ment of Agriculture, Agriculture Handbook No. of two Ptecticus Loew from Peninsular Malaysia 276. Washington, D.C. iv + 1696 pp. (Diptera, Stratiomyidae). Tijdschrift voor Ento- 1981. Stratiomyidae, pp. 497-511. In Mce- mologie 137: 75-86. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 300-311 DESCRIPTION OF THE IMMATURE STAGES OF THREE SPECIES OF EULEPIDOTIS GUENEE (LEPIDOPTERA: NOCTUIDAE) WITH NOTES ON THEIR NATURAL HISTORY MICHAEL G. POGUE AND ANNETTE AIELLO (MGP) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, “% National Museum of Natural History, Washington, DC 20560-0168, U.S.A. (e-mail: mpogue @sel.barc.usda.gov); (AA) Smithsonian Tropical Re- search Institute, PO. Box 2072, Balboa, Ancon, Republic of Panama (e-mail: aielloa@tivoli.si.edu) Abstract.—Larvae and pupae of the genus Eulepidotis Hiibner are described for the first time. The species are E. merricki (Holland), E. juncida (Guenée), and E. superior (Guenée), all of Neotropical distribution. The larval host of E. merricki is Spanish-lime, Melicoccus bijugatus Jacq. (Sapindaceae), which is cultivated as an ornamental and fruit tree throughout the Caribbean. Larvae of EF. juncida were reared from Inga fagifolia (L.) Willd. ex Benth. (= Inga laurina (Sw.) Willd.) (Mimosaceae). Larvae of E. superior were defoliating Quararibea asterolepis Pitt. (Bombacaceae). Key Words: Melicoccus, Sapindaceae, Inga, Mimosaceae, Quararibea, Bombacaceae, host plants, Neotropical, Panama, Puerto Rico Larvae of tropical Lepidoptera are poorly known, and unless reared to adults, they are virtually impossible to identify specifically. We describe for the first time the immature stages of Eulepidotis Hubner, a large genus belonging to the subfamily Catocalinae, that is mainly of Neotropical affinity and which comprises 105 species, 3 of which occur in North America (Poole 1989, Poole and Gentili 1996). The immature stages of E. merricki (Holland), E. juncida (Guenée), and E. superior (Guenée) were studied. There have been two reports of Eulepi- dotis larvae defoliating tropical trees. Wong et al. (1990) reported larvae of E. superior defoliating Quararibea asterolepis Pitt. (Bombacaceae) on Barro Colorado Island, Panama. Nascimento and Proctor (1994) re- ported that larvae of E. phrygionia Hamp- son were defoliating a monodominant rain- forest of Peltogyne gracilipes Ducke (Cae- salpiniaceae) on Maraca Island, Roraima, Brazil. Unfortunately no larvae from the latter study were preserved. Larvae, pupae, and adults of E. merricki were sent to one of us (MGP) by Lionel- Pagan, U.S.D.A., Animal and Plant Health Inspection Service, Plant Protection and Quarantine, San Juan, Puerto Rico, for identification. The larvae were defoliating Spanish-lime, Melicoccus bijugatus Jacq. (Sapindaceae), a tree used for fruit and as ornamental purposes in the Caribbean. Spanish-lime is native to continental tropi- cal America from Nicaragua to Surinam and is planted widely and becoming natu- ralized in the Caribbean (Adams 1972, Proctor 1984). One of us (AA) reared the immature stages of E. juncida and E. superior. The host of E. juncida was Inga fagifolia (L.) Willd. ex Benth. (= IJIngalaurina (Sw.) VOLUME 101, NUMBER 2 “eo aie Nerina A “ey naga Wea att i Larval habitus. Willd.) (Mimosaceae). The host of Eulepi- dotis superior 1s Quararibea asterolepis Pitt (Bombacaceae). METHODS AND MATERIALS Twenty-two larvae of E. juncida were collected and designated as Aiello Lot 80- 003. Fifteen of the larvae were preserved in 80% ethanol. Seven were placed in a cy- lindrical rearing cage along with both old and young foliage. The cage was construct- ed from petri dishes and aluminum window screen with a circle of paper towel on the floor and measured 10 cm tall by 9 cm in diameter. To maintain and regulate humid- ity, a damp, folded strip of paper towel was placed on the cage cover, and the entire as- sembly was kept inside a clear plastic ZipLoc® bag. After pupation the pupae were placed into separately numbered cages to await eclosion. Fifteen larvae, one pupa, and four adults with associated pupal skins are in the collection of the National Muse- um of Natural History, Smithsonian Insti- tution, Washington, D.C. 1, Eulepidotis merricki. 2, E. juncida. 3, “~ censtee | Ye ementas E. superior. Three larvae and 10 pupae of E. superior were obtained and designated as Aiello Lot 85-19. The larvae were preserved in 80% ethanol, and seven adults with associated pupal skins were reared and are in the col- lection of the Smithsonian Tropical Re- search Institute, Republic of Panama. KEY TO KNOWN EULEPIDOTIS LARVAE 1. Head dark brown to black, reticulate pattern reduced; mid-dorsal stripe black, solid (Fig. 1); setae Al—A2—A3 form an acute angle (Fig. 5) ee eegeto a oo ere oes ase E. merricki (Holland) — Head brown, reticulate pattern covers most of head; mid-dorsal stripe broken (Figs. 2—3); se- tae Al—A2—A3 form an obtuse angle cane POON ieee Gs yaa sions, cscs oy eee ee ea eee 2 2. Head with reticulate pattern restricted to lateral area; dorsolateral markings on abdomen in pairs (Fig. 2); labrum with one pair of setae on lateral margin (Fig. 14) .... E. juncida (Guenée) — Head uniformly covered with a reticulate pat- tern; abdominal segments with single large dorsal markings (Fig. 3); labrum with two pairs of setae on lateral margin (Fig. 21) E. superior (Guenée) 302 KEY TO KNOWN EULEPIDOTIS PUPAE 1. Length less than 12 mm; labial palpus disjunct (ioe O) rs Se ese ee Core E. juncida (Guenée) — Length greater than 13 mm; labial palpus con- timuous (Figss 25sand) 34) eras eee 2 . Profemur absent (Fig. 25) tN Eulepidotis merricki (Holland) (Figs. 1, 4-10, 25-29) Larva Diagnosis.—Dorsal black stripe from pronotum to transverse stripe on segment 8 and extending beyond band to segment 9. Head black to dark brown, without reticu- late pattern. Description.—Head (Figs. 5—10): Width of head capsule 1.8 + .06 mm (range, 1.8— 1.9 mm) (n = 5). Black to dark brown; ec- dysial line, epicranial suture, and posterior margin of head to stemmatal area cream. Prothoracic shield dark brown; medial patch dark brown withthin cream stripe; dorsolateral band cream; lateral edges dark brown. Labrum medially cleft; 3 pairs of dorsal setae forming an oblique line medi- ally (Fig. 7); 3 pairs of ventral epipharyn- geal setae (Fig. 8). Mandible with 3 distinct broad teeth; oral surface with broad molar- bearing process (Fig. 10). Thorax: Dorsal stripe black; dorsolateral band cream; lateral band broad, black, di- vided by a series of cream spots, extending to just below setae L1 and L3. Legs dark brown. Underside dark brown to metatho- racic legs, caudal half of mesothorax cream; V1 setae surrounded by dark brown spot. Abdomen (Fig. 1): Dorsal stripe black to broad transverse band on segment 8 and ex- tends between segments 8 and 9; dorsolat- eral band cream; lateral stripe black; lower lateral stripe with irregular margins, cream; spiracular stripe black with irregular mar- gins and not enclosing spiracle; below spi- racles and venter cream. Prolegs cream; plantae black; crochets in an uniordinal me- soseries. Segments | and 2 with 3 SV setae. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Pupa Diagnosis.—Labial palpi present, contin- uous; profemur absent. Description.— Male (Figs. 25-27): Length 15.8 + 1.04 mm (range, 15.0—17.0 mm) (n = 3). Labial palpi present, contin- uous. Profemur absent. Mesothoracic leg- reaching eye. Wings do not extend beyond caudal margin of segment 4. Segments 1—3 and 8 with shallow circular pits on dorsum; segments 4—7 with shallow circular pits in a dense band extending about %4 width of segment caudally and completely encircling segments 5—7; segment 9 smooth. Genital opening on a circular plate. Anal opening below genital opening. Cremaster consist- ing of a large median pair of slightly curved hooks and 3 pairs of small hooks, 1 pair located mediodorsally, 2 pairs located lat- erally (Fig. 27). Female (Figs. 28-29): Similar to male except: length 15.2 = 0.35 mm (cange, 15.0—15.5 mm) (n = 2). Genital opening at caudal border of segment 8, dividing seg- ment 8 ventrally. Anal opening well caudad of genital opening. Host.—Melicoccus bijugatus Jacq.; Fam- ily Sapindaceae; common names: Spanish- lime, genip, honeyberry, mamoncillo, que- nette, Quenepa, Hongibeere. Natural history.—The only information known is that thousands of larvae were de- foliating the host plant (Lionel Pagan, per- sonal communication). Discussion.—The adult of E. merricki most closely resembles Eulepidotis carcis- tola Hampson and E. fumata (Felder and Rogenhofer). In E. merricki the prothorax and tegula have a black stripe; this is absent in E. carcistola and E. fumata. An elongate white spot at the base of median line of the forewing is larger and more distinct in E. carcistola than in E. merricki and E. fu- mata. The forewing lines in E. fumata are very faint compared with the bold distinct lines in E. merricki. Eulepidotis merricki was described from a specimen collected flying around low VOLUME 101, NUMBER 2 303 8 0% Figs. 4-10. Eulepidotis merricki larva. 4, Setal map. 5, Head. 6, Head, lateral view (scale = 0.5 mm). 7, Labrum, dorsal view. 8, Labrum, ventral view. 9, Mandible, dorsal view. 10, Mandible, ventral view (scale = 0.1 mm). 304 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON T 2-3 14 | 16 15 line Figs. 11-17. Eulepidotis juncida larva. 11, Setal map. 12, Head. 13, Head, lateral view (scale = 0.5 mm). 14, Labrum, dorsal view. 15, Labrum, ventral view. 16, Mandible, dorsal view. 17, Mandible, ventral view (scale = 0.1 mm). herbage outside New Brighton, Pennsylva- onym of Palindia fumata Felder and Ro- nia, on August 5, 1900. Holland (1902) in genhofer (= E. fumata). The point is that the original description stated that itis very E. merricki is not a Nearctic species but a close to Palindia mabis Guenée (= E. jun- Neotropical one, and that H. D. Merrick cida (Guenée)) which he confused as asyn- just happened to collect a specimen that VOLUME 101, NUMBER 2 was somehow imported from the Neotrop- ics, possibly on tropical fruits. There are no other specimens of FE. merricki from the Nearctic in the National Museum, Washing- ton, D.C., or The Natural History Museum, London, collections. In Franclemont and Todd (1983) E. merricki is noted as being of questionable occurrence in America north of Mexico. Eulepidotis merricki has a Caribbean dis- tribution, having been collected from Ja- maica, Cuba, and Puerto Rico. Eulepidotis juncida (Guenée) (Figs. 2, 11-17, 30-33) Larva Diagnosis.—Head brown with reticulate pattern restricted to lateral area. Dorsum of abdomen with pair of dorsolateral brown patches encompassing setae D1 and D2. Description.—Head (Figs. 12-17): Width of head capsule for three instars as follows: 1.2 mm (n = 1); 1.8 + .03 mm (range, 1.7—1.8 mm) (n = 7); 2.0 + .05 mm (range, 2.0—2.1 mm) (n = 7). Brown with lateral reticulate pattern; adfrontal area from ecdysial line to just beyond setae AF1 cream; setae Pl and P2 with cream pina- cula. Labrum medially cleft; 3 pairs of dor- sal setae with second pair below other pairs; 3 pairs of ventral epipharyngeal setae (Fig. 15). Mandible with 2 distinct broad teeth, | smaller tooth mostly obscured from dorsal view; oral surface with a broad molar-bear- ing process with small pointed processes on either side (Fig. 17). Thorax: Prothoracic shield brown; dorso- lateral band cream; small cream patch be- tween D2 and XD2. Dorsal stripe or patch on segment 2 brown, on segment 3 cream; dorsolateral band cream; lateral band broad, brown, interrupted by several cream spots, extending to just below setae LI and L3. Legs brown. Underside mostly cream, brown shading between coxae and encom- passing V1 setae. Abdomen (Fig. 2): Dorsum with pair of dorsolateral brown patches encompassing setae D1 and D2; lower lateral stripe cream, 305 consisting of irregular spots and stripes that are more or less contiguous to segment 6, line not contiguous to absen! on segments 7 to 9; spiracular band brown, broad, ex- tending below L1 seta; below L1 seta and venter cream. Prolegs cream to brown lat- erally; plantae cream; crochets in an unior- dinal mesoseries. Segments | and 2 with 3 SV setae. Pupa Diagnosis.—Small, less than 12 mm long. Labial palpi present, but disjunct. Pro- femur present. Description.—Male (Figs. 30-33): Length 9.3 + 0.40 (range, 8.9—-9.7 mm) (n = 3). Labial palpi present, disjunct. Profe- mur present. Mesothoracic leg reaching eye. Wings do not extend beyond caudal margin of segment 4. Segments 1—3 and 8 dorsum with shallow circular pits; segments 4-7 with shallow circular pits in a dense band extending about %4 width of segment caudally and completely encircling seg- ments 5—7; segment 9 smooth. Genital opening on a circular plate. Anal opening caudad of genital opening. Cremaster con- sisting of a large median pair of curved hooks and 3 pairs of small hooks, | pair located mediodorsally, 2 pairs located lat- erally (Figs. 32—33). Female (Fig. 33): Similar to male except: length 9.4 + 0.99 mm (range, 15.0—17.0 mm) (n = 2). Genital opening at caudal border of segment 8, dividing segment 8 ventrally. Anal opening well caudad of gen- ital opening. Host.—Inga fagifolia (L.) Willd. ex Benth. (= Inga laurina (Sw.) Willd.); fam- ily Mimosaceae. Natural history.—On March 5, 1980, an outbreak of larvae occurred on Barro Col- orado Island, Republic of Panama, near the 900 meter mark on Thomas Barbour Trail on the central plateau of the island. Larvae were so abundant that the rainlike sound of their fecula landing on the leaf litter could be heard many meters away. Although the majority were in the cano- 306 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 22 24 Figs. 18-24. Eulepidotis superior larva. 18, Setal map. 19, Head. 20, Head, lateral view (scale = 0.5 mm). 21, Labrum, dorsal view. 22, Labrum, ventral view. 23, Mandible, dorsal view. 24, Mandible, ventral view (scale = 0.1 mm). VOLUME 101, NUMBER 2 py, larvae fed at various levels in the tree eating only young leaves. They dropped on silk lines to the forest floor and surrounding vegetation to molt. Molting took place un- der a sheet of silk across a leaf. Larvae walked by “‘looping,”’ as do geometrids, but they have the full complement of five pairs of prolegs. Some larvae dropped, presumably in re- sponse to some perceived danger, and be- gan reascending almost immediately. They progressed slowly by gathering the silk line into a ball using their thoracic legs. Upon completion of their journey, they aban- doned the ball of silk. Seven larvae were brought into the lab- oratory on March 5; one died on March 7. The remaining six molted on March 9, and late on March 10 began preparing pupation chambers of fecula and intact leaves on the cage floor. Five pupated on March 11, and the sixth larva failed to pupate and died on March 12. Two adults eclosed during the night of March 19-20, and two more eclosed during the night of March 20-21. The fifth pupa died and was preserved. Pu- pal duration was 9 days for females and 10 days for males. Discussion.—In the adult stage, E. junci- da is similar to E. juliata (Stoll). In E. jun- cida the median brownish-yellow stripe ex- tends from the costa to the inner margin of forewing, in contrast to E. juncida, which has a more whitish-yellow median stripe that does not extend to the inner margin. Eulepidotis juncida has a wide distribu- tion from Mexico to Panama in Central America and from Guyana, Venezuela, and Colombia to Bolivia in South America. Eu- lepidotis juliata is known, so far, only from Guyana, Venezuela, and Bolivia. Eulepidotis superior (Guenée) (Figs. 3, 18-24, 34-37) Larva Diagnosis.—Reticulated pattern covering entire head. A single large mid-dorsal light brown (in alcohol specimens) mark on all abdominal segments. 307 Description.—Head (Figs. 19-24): Width of head capsule for two instars as follows: 1.9 mm (n = 1); 2.6 mm (n = 2). Brown with cream reticulated pattern. La- brum cream, medially cleft; 3 pairs of dor- sal setae forming a straight line medially; 2 pairs of small setae along lateral edge; 3 pairs of ventral epipharyngeal setae (Fig. 22). Mandible with 2 outer setae; cutting surface with 3 indistinct teeth; oral surface with broad molar-bearing process (Fig. 24). Thorax: Prothoracic shield brown; mid- dorsal stripe faint, cream; lateral stripe wide, cream; marked with various spots and patches of cream. Cream dorsally with a few faint mid-dorsal patches of brown. Pro- thorax with SD and L setal groups on brown pinacula. Thoracic segments 2 and 3 with seta SD1 and setae LI and L3 on brown pinacula. Legs cream. Underside cream. Abdomen (Fig. 3): Mid-dorsal quadrate marks light brown, less distinct on seg- ments 4—6 and 9; dorsolateral band cream; lateral band broad, brown, broken by series of cream spots along dorsal margin; distinct cream spots slightly dorsal and posterior to spiracles. D1 seta inside, D2 seta outside dorsal patch. Segments | and 2 with SD1, L1, and L3 on brown pinacula. Segments 3—6 and 8 with SD1 and L group setae on brown pinacula. Prolegs cream; crochets in an uniordinal mesoseries. Segments | and 2 with 2 SV setae, SV1 represented by a sclerotized ring near SV3. Pupa Diagnosis.—Labial palpus present, con- tinuous; profemur present. Description.—Male (Figs. 34-36): Length 15.8 + 1.04 mm (range, 15.6—19.6 mm) (n = 4). Labial palpi present, elongate, continuous. Profemur present. Mesothorac- ic leg reaching eye. Wings do not extend beyond caudal margin of segment 4. Seg- ments 1—3 and 8 with shallow circular pits on dorsum; segments 4—7 with shallow cir- cular pits in a dense band extending about % width of segment caudally and complete- 308 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ant Da oe" ee 3 Pr ee 00 0 oro ° snes oe =i oo OOS coo ° b 0, Os * 2005400 © 900 oetO2 Oped fo 400s “FORE “op8%° Cts. ® OFM foo ** 000 #09" O, .9% Poea0 * 2° con x ro coe ee O82 2005 22900024208 D008 Be ESOL? se OMe Oe 4 of o: 000.9% 0005 224900%%%08 Soe 2 084 80 oo 09388 0 O'de? Do ° * c@08 o ° eo oe : rela srocamoeapaccaire d oeoF 08S +2 oo. . eo ©6825 “eact oa Ralston ccleei@>oeet ee cine Sees BY . fee oe - 8 9 *20 ° 2 oe 0000 om % 2%20 oO, > oss > © 9 0.g 2-22 05200008" % 25 Bk BSG Birete 4O SB Me ® 000 440° Cn eeQ00 °F P25 fo fe ° 2° Poo cary 28 29 26. ¢ dorsal view (scale = 1.0 mm). Df es 27 Eulepidotis merricki pupa. 25, 4 ventral view. 28. 2 ventral view of terminal segments. 29, Dorsal view of terminal segments labial palpi; 11 Figs. 25-29. ventral view of terminal segments. (scale = 0.5 mm). Abbreviations: = mesothoracic leg; mx ant —antenna; ao = anal opening; go = genital opening; Ip = = prothoracic leg; 12 = maxilla. VOLUME 101, NUMBER 2 00 0, ,0° 00 Siete 'o* 32 fl = femur of prothoracic leg; lp = labial palpi. 20s = = 02.2 oOo 2,0 2 0 Oe," 00 2°° ° ° © PPO 2.2,9,°, 209,22 og2 © 2 232°) ° ° Oo Pee OSt OS Poe Oe 8 Oe 290 ° ° oe %& ©2060 = elo ° oo 0% oO 2 go? ° 0000 °2S0 ,0o,4 oo Snel e ©) (les:8]a\e'o= 33 Figs. 30-33. Eulepidotis juncida pupa. 30, d ventral view. 31, ¢d dorsal view (scale 309 kOSmm) 32, ventral view of terminal segments. 33, 2 ventral view of terminal segments (scale = 0.5 mm). Abbreviations: 6) 310 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Taine too tte Barre Le, Wb Ce DOT ae 34, 3 ventral view. 35, 3 dorsal view (scale = 1.0 mm). 36, 6 al segments (scale = 0.5 mm). Abbreviations: Figs. 34-37. Eulepidotis superior pupa. ventral view of terminal segments. 37, ° ventral view of termin fl = femur of prothoracic leg; Ip = labial palpi. VOLUME 101, NUMBER 2 ly encircling segments 5—7; segment 9 smooth. Genital opening on an oval plate. Anal opening caudad of genital opening. Cremaster consisting of a large median pair of curved hooks and 3 pairs of small hooks, 1 pair located mediodorsally, 2 pairs locat- ed laterally (Fig. 36). Female (Fig. 37): Similar to male except: length 16.1 + 1.03 mm (range, 15.0—17.0 mm) (n=3). Genital opening at caudal border of segment 8, dividing segment 8 ventrally. Anal opening well caudad of genital opening. Host.—Quararibea asterolepis Pitt.; family Bombacaceae. Natural History.—During late May through early June 1985, a massive, highly synchronized outbreak of larvae was ob- served on the central plateau of Barro Col- orado Island, Republic of Panama (Wong et al. 1990). The larval host plant was under- going leaf flush and the larvae were feeding on young leaves and often defoliating the trees. Once feeding was completed, the lar- vae descended on silk lines and pupated be- neath dried leaves on the forest floor. Lar- vae were so abundant that it was difficult to avoid walking into their silk lines or stepping on larvae and pupae. A total of 10 pupae and 3 larvae were brought to one of us (AA) by Maria Wong, Seiji Tanaka, and Peter Becker. Three pupae died, and seven were reared to adults. Adult no. 1 (2) eclosed 21 June from a pupa col- lected 7 June. Adult nos. 4 (¢) and 5 (9) eclosed 23 and 24 June, respectively, from pupae collected 11 June. Adult nos. 6—9 eclosed 18 June (2), 20 June (d), and 21 June (2d d), respectively, from pupae col- lected 14 June. All eclosions took place in early evening, some as early as 5:30 PM, and others as late as 9 PM. Discussion.—Adults of E. superior are the largest (forewing length 13—9.5 mm) of the brown species of Eulepidotis and cannot be confused with any other species in the genus. It is distributed from Mexico to Pan- ama in Central America, northwestern South Americain Venezuela, Colombia, and Ecua- 311 dor, and in the Caribbean on the islands of Puerto Rico, Grenada, and St. Lucia. ACKNOWLEDGMENTS We thank Robin B. Foster of the Smith- sonian Tropical Research Institute, Balboa, Republic of Panama, for identifying the host plant of E. juncida; Robert W. Poole of Entomological Information Services, Rockville, Maryland, for identifying E. sw- perior, and Lionel Pagan of U.S.D.A., AVPELLES:, BPO. San: Juan, Puerto. Rico, for supplying the immature stages and reared adults of E. merricki. We thank Wil- liam E. Miller, University of Minnesota, St. Paul, Minnesota, and Douglass R. Miller and David R. Smith of the Systematic En- tomology Laboratory, U.S.D.A., Beltsville, Maryland, and Washington, D.C., for criti- cally reviewing and offering suggestions that greatly improved the manuscript. Linda Lawrence, Systematic Entomology Labo- ratory, U.S.D.A, Washington, D.C., pre- pared the habitus illustrations. LITERATURE CITED Adams, C. D. 1972. Flowering plants of Jamaica. Uni- versity of the West Indies, Mona, Jamaica, 848 pp. Franclemont, J. G. and E. L. Todd. 1983. Noctuidae, pp. 120-159. In Hodges, R. W. et al., eds., Check list of the Lepidoptera of America North of Mex- ico. The Wedge Entomological Research Foun- dation, Washington, D.C., 284 pp. Holland, W. J. 1902. Description of an apparently new species of Palindia. Entomological News 13: 172-173. Nascimento, M. T. and J. Proctor. 1994. Insect defoli- ation of a monodominant Amazonian rainforest. Journal of Tropical Ecology 10: 633-636. Poole, R. W. 1989. Lepidopterorum Catalogus (New Series), Fascicle 118, Noctuidae. E. J. Brilland Flora and Fauna Publications, Leiden, 1,313 pp. Poole, R. W. and P. Gentili (eds.). 1996. Nomina In- secta Nearctica. A check list of the insects of North America, Vol. 3: Diptera, Lepidoptera, Si- phonaptera. Entomological Information Services, Rockville, Maryland, 1,143 pp. Proctor, G. R. 1984. Flora of the Cayman Islands. Roy- al Botanical Gardens, Kew, London, 834 pp. Wong, M., J. Wright, S. P. Hubbell, and R. B. Foster. 1990. The spatial pattern and reproductive con- sequences of outbreak defoliation in Quararibea asterolepis, a tropical tree. Journal of Ecology 78: 579-588. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 312-315 OBSERVATIONS ON THE PREY AND NEST CLUSTERS OF PODALONIA VALIDA (CRESSON) (HYMENOPTERA: SPHECIDAE) KEVIN M. O’ NEILL AND HOWARD E. EVANS (KMO) Department of Entomology, Montana State University, Bozeman, MT 59717, U.S.A. (e-mail: koneill@montana.edu); (HEE) Department of Entomology, Colorado State University, Fort Collins, CO 80523, U.S.A. Abstract.—In northern Colorado, Podalonia valida preyed upon mature larvae of the saltmarsh caterpillar, Estigmene acrea (Drury), the same prey it uses in Arizona. Prey were deposited in shallow, single-celled nests that occurred within clusters probably pro- visioned by single females. Because females forage for abundant nearby prey, stock each nest cell with a single caterpillar, and dig simple nests in a localized area, they have the potential to provision multiple nests over short time periods. Several observations are consistent with this hypothesis. First, within clusters of nest cells, there was a high degree of synchrony in developmental stages of the wasps, suggesting that eggs were laid in quick succession. Second, dissections showed that female P. valida carry more mature eggs in their ovaries than is typical for non-parasitoid aculeate wasps. Key Words: marsh caterpillar, Arctiidae Wasps of the genus Podalonia dig short burrows terminating in a cell provisioned with a single prey (Bohart and Menke 1976). Podalonia luctuosa_ (Smith) (O’Brien and Kurczewski 1982), Podalonia robusta (Cresson) (Kurczewski et al. 1992), and Podalonia argentifrons (Cresson) (O’Brien 1983) restrict themselves to cut- worms (Noctuidae), but may take a variety of species at any one site. Similarly, Po- dalonia occidentalis Murray apparently prey solely on tent caterpillars (Lasiocam- pidae: Malacosoma) (Evans 1987) and Po- dalonia valida (Cresson) limit themselves to arctiid moth caterpillars. Steiner (1974, 1975) found P. valida specializing on salt- marsh caterpillars (Estigmene acrea (Dru- ry)) in southern Arizona, whereas Rust et al. (1985) found two female P. valida tak- ing Apantesis proxima (Guérin-Méneville) on San Clemente Island, California. Here, Digger wasp, Sphecidae, nesting behavior, egg size, Estigmene acrea, salt- we present results of our observations of the nesting behavior and prey of P. valida in northern Colorado and compare our find- ings to those of Steiner, who concentrated on the hunting and territorial behavior of females. METHODS We studied P. valida females at the Paw- nee National Grasslands in northern Weld Co., Colorado, from 12 July to 13 August 1984. The site was situated along a sandy, little-used road that passed through prairie consisting of grasses mixed with several common forbs, notably white sweetclover (Melilotus alba Desr.) and sunflower (He- lianthus spp.). RESULTS AND DISCUSSION Our 64 prey records indicate that P. val- ida preyed exclusively on mature larvae of VOLUME 101, NUMBER 2 the saltmarsh caterpillar Estigmene acrea (Drury) (Arctiidae), the same species used by this wasp in Arizona (Steiner 1974). The prey of P. valida, which were probably tak- en on their host plants (white sweetclover), were carried in the mandibles as females walked forward straddling the prey. One fe- male carried a caterpillar at least 10 m be- fore reaching her nest. Prey-carrying fe- males occasionally stopped and cached their prey ~5 cm above the soil surface on small plants, where they were left for sev- eral minutes while females searched for their nests. Unlike other species of Podalonia, P. valida dig nests before hunting (Steiner 1974, 1975). Each nest at our site consisted of a short, oblique burrow about | cm in diameter, terminating in a single | X 3 cm cell situated 3—5 cm beneath the soil surface (N = 7). Typically, when digging the nest, a female backed out 5—10 cm from the bur- row entrance and scraped soil backwards with her forelegs, while elevating her ab- domen, flicking her wings rapidly, and buzzing loudly. Upon returning to her nest with prey, a female dropped it at the en- trance, cleared the burrow, and pulled the prey inside while moving backwards. After several minutes inside the nest, during which time she laid an egg on the prey, the female emerged and permanently closed the burrow by scraping in soil from the edge of the burrow and placing lumps of soil or pebbles in the hole. She then tamped the loose soil with her head (mandibles wide open) or with a lump of soil which was held in the mandibles and which broke up due to the impact. The entire sequence between entering the hole with prey and completing closure typically took about 4 minutes. These observations are in accord with those of Steiner (1974, 1975), who found that individual females provisioned series of single-celled nests within small patches of bare soil sometimes no more than 60 cm across. Although we did not observe indi- vidually marked females over prolonged periods, we also found that cells tended to B18 be clustered. In 2 of our 7 excavations of recently completed nests, we unearthed just a single cell, but in the others, we found clusters of 8, 15, 16, 18, and 27 cells within areas no more than 0.25 m’*. Some cells were separated by as little as 5 cm, but were definitely parts of different nests. Clusters of nest cells were well-separated from one another and each was apparently used ex- clusively by a single female, who vigorous- ly attacking conspecific females intruding upon their nest cluster in interactions that included bouts of grappling between the combatants. Similar interactions between P. valida females have been described in de- tail by Steiner (1975). The contents of unparasitized and non- moldy P. valida cells within clusters sug- gest that, if a single female was responsible for all of the cells, some were provisioned during short time intervals. For example, in the cluster of 15 cells, 5 had prey with wasp eggs, 3 had prey with small wasp larvae, and | had a large wasp larva. In the cluster of 7 cells, all had prey with unhatched eggs. Similarly, the cluster of 18 cells included 7 prey with wasp eggs and 7 with small wasp larvae, and the cluster of 16 cells contained 7 with eggs or small larvae. The potential ability of P. valida fe- males to provision multiple nests in rapid succession may be possible because they 1) forage for abundant prey nearby. 2) stock each cell with a single prey, and 3) dig simple, shallow nests without search- ing widely for successive nesting sites. This strategy may allow them to exploit a single developmental stage of a single prey species that is available for just a brief pe- riod during the summer. However, the rap- id stocking of multiple nests would also require that a female produce the requisite numbers of eggs. Typically, female sphe- cids carry no more than two mature eggs at a time, although they have three ovari- oles in each ovary (Iwata 1964, O’ Neill 1985). Nevertheless, the three P. valida fe- males that we captured within minutes of their laying eggs had 3, 5, and 6 mature 314 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (or nearly mature) oocytes still in their ovaries. Thus, the latter female had been carrying seven well-developed oocytes just prior to capture and had one ovariole with two mature oocytes. In Iwata’s extensive survey of the ovaries of solitary aculeate wasps, only parasitoid sphecids of the gen- era Larra (with up to 21 mature oocytes) and Chlorion (with up to 10) carried more mature eggs than the maximum for P. val- ida. Such high levels of short-term fecun- dity (for a digger wasp), however, may be achieved at the cost of producing smaller eggs. The mature oocytes of P. valida ranged in length from 2.6—3.1 mm in length and one egg found on prey was 2.9 mm long. In contrast, in the similarly-sized but less fecund wasp Philanthus bicinctus (Mickel), mature oocytes and newly laid eggs ranged as high as 5.4 mm in length, and had perhaps 3—4 times the volume of P. valida eggs (O’ Neill 1985). Bembecinus quinquespinosus (Say) and Philanthus pulcher Dalla Torre carry 1—2 mature o0o- cytes within the same size range as P. val- ida, although they are both much smaller wasps. Of the 86 cells we excavated, four had fly puparia 0.5—1.0 cm below the cell, which in each case-contained the remains of a single saltmarsh caterpillar. One pu- parium gave rise to a tachinid of the genus Exorista, at least one species of which is a known parasitoid of FE. acrea (Arnaud 1978). The wasps had probably captured previously parasitized caterpillars and suf- fered incidental cleptoparasitism. Another 23% of the cells contained molded or rot- ting caterpillars. Our observations complement those of Steiner (1974, 1975), indicating that two widely separated populations of P. valida have identical prey preferences and simi- lar nesting behaviors that are unique for this genus. Podalonia valida seems to have adopted a strategy intermediate be- tween those of the relatively high fecun- dity parasitoid sphecids (Larra and Chlo- rion) and the more typical low fecundity nest provisioning species that spend con- siderable time and energy on each off- spring. This strategy is facilitated by the greater number of mature eggs they carry and, perhaps, by shorter periods of search- ing for potential nest sites once nesting has begun. ACKNOWLEDGMENTS We thank Patricia Denke, Wendell Mor- rill, Mark O’Brien, and Catherine Seibert for commenting on the manuscript and N. Woodley (Systematic Entomology Labora- tory, USDA, Washington, DC) for deter- mination of the tachinid fly. The work was supported by the National Science Foun- dation, the Colorado Agricultural Experi- ment Station, and and the Montana Agri- cultural Experiment Station. LITERATURE CITED Arnaud, P. H. 1978. A host-parasite catalog of North American Tachinidae (Diptera). Miscellaneous Publication No. 1319, Science and Education Ad- ministration, United States Department of Agri- culture, 860 pp. Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps of the World: A Generic Revision. Berkeley: Uni- versity of California Press, ix + 965 pp. Evans, H. E. 1987. Observations on the prey and nests of Podalonia occidentalis Murray (Hymenoptera: Sphecidae). Pan-Pacific Entomologist 63: 130— 134. Iwata, K. 1964. Egg gigantism in subsocial Hymenop- tera, with ethological discussion on tropical bam- boo carpenter bees. Nature and Life in Southeast Asia 3: 399—434. Kurezewski, EF E., M. EF O’Brien, and M. G. Spofford. 1992. Nesting behavior of (Cresson) (Hymenoptera: Sphecidae). Journal of Hymenoptera Research |: 235-239. O’Brien, M. F. 1983. Observations on the nesting be- havior of Podalonia argentifrons. Southwestern Entomologist 8: 194—197. O’Brien, M. E and FE E. Kurcezewski. 1982. Ethology and overwintering of Podalonia luctuosa (Hy- menoptera: Sphecidae). Great Lakes Entomologist 15: 261-275. O'Neill, K. M. 1985. Egg size, prey size, and sexual Podalonia robusta size dimorphism in digger wasps (Hymenoptera: Sphecidae). Canadian Journal of Zoology 63: 2187-2193. Rust, R., A. S. Menke, and D. Miller. 1985. A biogeo- graphic comparison of the bees, sphecid wasps, VOLUME 101, NUMBER 2 and mealybugs of the California Channel Islands (Hymenoptera, Homoptera). pp. 29-59 in Menke, A. S. and D. R. Miller, eds., Entomology of the Channel Islands. Santa Barbara Museum of Nat- ural History, Santa Barbara, California. Steiner, A. L. 1974. Unusual caterpillar-prey records and hunting behavior for a Podalonia digger 315 wasp: Podalonia valida (Cresson). Pan-Pacific Entomologist 50: 73-77. Steiner, A. L. 1975. Description of territorial behavior of Podalonia valida (Hymenoptera: Sphecidae) females in southeast Arizona, with remarks on digger wasp territorial behavior. Quaestiones En- tomologica 11: 113-137. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 316-324 IDENTIFICATION OF LATE-INSTAR NYMPHS OF COCKROACHES (BLATTODEA) KEVIN T. HATHORNE AND PATRICIA A. ZUNGOLI Department of Entomology, Clemson University, Box 0365, Clemson, SC 29634-0365, U.S.A. (e-mail: kevinh@clemson.edu) Abstract.—A dichotomous key to late-instar nymphs of 12 cockroach species common in the United States is presented. Notes on biology, distribution, and taxonomy are given. Key Words: More than 3600 species of cockroaches are known worldwide, 69 of which are found in North America. Fifteen to twenty of these species are of economic importance as nuisance pests. Classification of cock- roaches has undergone many changes. Rehn (1951) classified adult cockroaches based on wing characteristics. Princis (1960) pro- vided a comprehensive history of cock- roach taxonomy and reported previously unknown information on cockroach evolu- tion. McKittrick (1964) grouped the cock- roaches with mantids based on evolutionary studies, morphology of genitalia and pro- ventriculus, and oviposition behavior. Corn- well (1968) provided a thorough review of the history of cockroach classification. He took the reader from 1758 when all cock- roaches were placed in the genus Blatta, in the order Coleoptera, through the revision of Imms’ Textbook of Entomology (Rich- ards and Davies 1957) in which cockroach- es are placed in their own order, Dictyop- tera. Cockroaches are currently placed in the order Blattodea as recognized by the Entomological Society of America (Bosik 19977)): Dichotomous keys to the cockroaches are primarily restricted to the adult stage (Blatchley 1920, Rehn 1950, Pratt and Sto- janovich 1962, Dakin and Hays 1970) ex- Insecta, Blattodea, late-instar nymphs, descriptions, dichotomous key cept for those by Powell and Robinson (1980), Fisk (1987), and Gordon (1992). Powell and Robinson (1980) included only first-instar nymphs of five Periplaneta spe- cies and Gordon (1992) distinguished the mid-instar nymphs of Periplaneta ameri- cana and P. fuliginosa. Fisk (1987) includ- ed nymphs of 16 cockroach species with identification based on comparative char- acters. Because behavior and habitat preference differ substantially among species, and de- velopment of effective control strategies de- pends on correct identification, a dichoto- mous key to late-instar cockroach nymphs is needed. The present work provides a means of identifying 12 of the pest species found in North America. MATERIALS AND METHODS Based on their pest status, late-instar nymphs of 12 species of cockroaches rep- resenting 3 families (Blaberidae, Blattelli- dae, and Blattidae) are included in this key. A cockroach may go through 5 to 12 molts before reaching the adult stage. The indi- vidual specimens were selected for size (ap- proximately the same size as the adult of the same species). Exact instar was not known for each individual. Specimens of the following eight species were obtained VOLUME 101, NUMBER 2 from colonies maintained by the Clemson University Urban Entomology Laboratory: Blattella germanica (L.), Supella longipal- pa (FE), Periplaneta americana (L.), Peri- planeta australasiae (FE), Periplaneta brun- nea Burmeister, Periplaneta fuliginosa (Serville), Blatta orientalis (L.), and Par- coblatta lata (Brunner). Three species (Blattella asahinai Mizukubo, Panchlora nivea [L.], and Eurycotis floridana [Walk- er]) were obtained from the USDA-ARS laboratory in Gainesville, Florida. One spe- cies, Blattella vaga Hebard, was obtained from Virginia Polytechnic Institute and State University, Blacksburg, Virginia. Ten to twenty specimens of each species were examined for morphological differ- ences, using a dissecting microscope (WILD Heerbrugg Switzerland M5-3984). Three species of cockroaches in this key are wingless as adults. These wingless spe- cies, on close examination, have truncated wings (B. orientalis (female), E. floridana), or wing pads (female P. lata). This char- acter may confuse the identification of the adult form with a late-instar nymph of an- other species. All characters used in this key can be seen when the specimen is viewed dorsally. KEY TO LATE-INSTAR COCKROACH NYMPHS 1. Body tan. Thorax with 2 dark brown to black ParallelMlinesss vey sete tees ees eee 2 — Body color variable. Thorax without parallel INBOVESS G38 n't OM LONR te een oie DEES Gi Reet - N Abdomen with 2 tan dots on some or all ter- gites in center of dark longitudinal area. Ab- domen with tan markings on lateral edges separated by dark brown markings between tergites (note: All above characters may vary slightly or may not be present). Blattella germanica (Fig. 2) Abdomen with markings different from above 3 3. Abdomen with tan, horizontal bar on some or all tergites in center of dark longitudinal area. Body with black, vertical line through center. Cercus dark brown to black on terminal ends. Abdomen with tan markings on lateral edges separated by dark brown markings between tergites (note: All above characters may vary slightly or may not be present)........ Blattella vaga (Fig. 3) n 317 Abdomen with tan, horizontal bar bearing an- terior emargination on some or all tergites in center of dark longitudinal area. Body without black, vetical line. Cercus tan with dark brown to black markings on terminal ends. Abdomen with tan markings on lateral! edges not separated by dark brown markings be- tween tergites (note: All above characters may vary slightly or may not be present) Blattella asahinai (Fig. 4) Cercus longer than distance between their ba- GES Toute blogs “edit eeoieticne eae ce ek tran A ial 5 Cercus shorter than distance between their ba- SCSe- eh: RA eye et ee LISP EME ee Psy) foe Se 10 Body variably black and tan. Pronotum with black horizontal bar bearing anterior and pos- terior median emarginations 5 arc eee Periplaneta australasiae (Fig. 5) Body uniform reddish-brown or with black on lateral edges of abdominal tergites or tan with dark brown markings. Pronotum with mark- ings different from above. Pronotum dark brown with tan on lateral edg- es. 2nd and 3rd thoracic sclerites with hori- zontal dark brown markings. Length less than INO 0000 ale fhe listat pe rots Supella longipalpa (Fig. 1) Pronotum with markings different than above. 2nd and 3rd thoracic sclerites without hori- zontal dark brown markings. Length greater than 15 mm Pronotum dark with litthe color variation. Male with styli shorter than 10th tergite SOW Ss eee Periplaneta fuliginosa (Fig. 6) Pronotum with some color variation, dark- ened areas. Styli variable Abdomen black, with lighter areas in center of tergites. 10th tergite truncate Saige weapon tes Ee ake Parcoblatta lata (Fig. 7) Abdomen with color pattern different from aboves lOthtergite notched™ 25>. 52.05 9 Male with styli slightly longer than distance between their bases. Cercus with last segment twice as long as wide (Fig. 8a) Dre erent Ea at Periplaneta americana (Fig. 8) Male with styli shorter than distance between their bases. Cercus with last segment less than twice as long as wide (Fig. 9a) Body black. Cercus longer than 10th tergite Blatta orientalis (Fig. 10) Body light brown or dark red and black. Cercus as long as, or shorter than, 10th tergite .... 1] Periplaneta brunnea (Fig. 9) . Body dark red in center, with black laterally. 9th abdominal tergite with posterior corners prolonged backward into sharp points. Tho- racic and abdominal lateral margins forming a smooth line in dorsal view. 10th tergite 318 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Lik eh) 4 ee Eurycotis floridana (Fig. 11) — Body uniformly reddish brown. 9th abdomi- nal tergite without points. Thoracic and ab- dominal lateral margins not forming a smooth line in dorsal view. 10th tergite longer than CEICUSHRa hia cent Panchlora nivea (Fig. 12) Notes on Species Supella longipalpa (Fig. 1).—The brown-banded cockroach nymph has a broad black stripe on the pronotum and oth- er black markings on the thorax. It is typi- cally less than 15 mm. This species is found throughout the United States in relatively sanitary conditions such as office buildings. It prefers temperatures over 27°C and is of- ten found above floor level. It will infest furniture, and deposit egg cases behind pic- ture frames and appliances. Blattella germanica, B. asahinai, and B. vaga are similar species. The German, Asian, and field cockroach nymphs are tan with two broad, dark brown to black lon- gitudinal stripes on the thorax. Because these species are so similar to one another, multiple characters must all be used for identification of species. As late-instar nymphs there are differences in body size between these three species. However, these size variations may overlap. Blattella germanica (Fig. 2).—When body length is compared to the other two Blattella species represented in this key, it usually is the median. The German cock- roach is perhaps the most economically im- portant of all cockroach pests. It is found throughout the United States in human hab- itations, and rarely outside. Blattella vaga (Fig. 3).—The field cock- roach nymph is typically smaller and lighter in color than B. germanica and B. asahinai. It is found in the southwestern United States in irrigated fields and yards. Blattella asahinai (Fig. 4).—The Asian cockroach nymph is typically 2 to 3 mm longer than B. germanica. Most individuals have three small black dots arranged in a triangle, a character not always observed in B. germanica or B. vaga. Found primarily outdoors in Florida, it occasionally enters structures. Parcoblatta lata (Fig. 7).—The broad wood cockroach nymph has a uniformly reddish brown thorax. The abdomen is mostly black with lighter markings in the center of the first three to four tergites. The 10th tergite is pointed and unnotched. It is found throughout the United States in wooded habitats. Adult females are wing- less. Adult males will occasionally enter buildings. Periplaneta australasiae (Fig. 5).—The Australian cockroach nymph is easily dis- tinguished by its coloration, it is black and tan and has a dark horizontal bar with an- terior and posterior emarginations on the pronotum. The adults share this character. This species is most commonly found in both indoor and outdoor situations in the coastal states of the southern United States and portions of California. Periplaneta fuliginosa (Fig. 6).—The smokybrown cockroach nymph, very simi- lar to P. americana and P. brunnea, can be distinguished by its uniform reddish brown color. It is found in the southeastern United States in hot, humid areas in and around structures. Periplaneta americana (Fig. 8).—Like P. brunnea, the American cockroach nymph is reddish brown with variable dark- ened areas. The cerci are long and slender and the last segment is twice as long as it is wide. If styli are present, they are longer than the distance between their bases. This species is found throughout the United States in association with buildings and warm, humid areas. Periplaneta brunnea (Fig. 9).—The cerci and styli on the brown cockroach are the key characters used to distinguish this spe- cies from P. americana. The cerci are more flattened and broader and the last segment is not twice as long as it is wide. If styli are present, they are shorter than the dis- tance between their bases. The brown cock- roach is found around buildings in the southeastern United States. VOLUME 101, NUMBER 2 319 2 Figs. 1-2. Dorsal view of late-instar nymphs. 1, Supella longipalpa, the brown-banded cockroach. 2, Blatella germanica, the German cockroach. 320 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 3-4. Dorsal view of late-instar nymphs. 3, Blatella vaga, field cockroach. 4, B. asahinai, the Asian cockroach. VOLUME 101, NUMBER 2 32 an 7 Kh Figs. 5-7. Dorsal view of late-instar nymphs. 5, Periplaneta australasiae, the Australian cockroach, 6, P. fuliginosa, the smokybrown cockroach. 7, Parcoblatta lata, the broad wood cockroach. 322 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON KTH Figs. 8-9. Dorsal view of late-instar nymphs. 8, Periplaneta americana, the American cockroach; 8a = dorsal view of left cercus. 9, P. brunnea, the brown cockroach; 9a = dorsal view of left cercus, note size and shape of last segment. VOLUME 101, NUMBER 2 KH Figs. 10-12. Dorsal view of late-instar nymphs. 10, Blatta orientalis, the Oriental cockroach. 11, Eurycotis floridana, the Florida woods cockroach. 12, Panchlora nivea, the Cuban cockroach. Blatta orientalis (Fig. 10).—The Oriental cockroach, easily identified by its uniform dark, reddish-black color, can also have a shiny appearance. Adult females have trun- cated wings. It is found in and around struc- tures over most of the temperate United States and Canada. Eurycotis floridana (Fig. 11).—The Flor- ida woods cockroach nymph is compara- tively large (35 mm) and dark in color. The center of the body is dark red and the outer edges are black. The posterior corner of the 9th abdominal tergite is prolonged back- ward to sharp points. Adults have truncated wings. An important pest in Florida, it also is found in southeastern Georgia and along the lower Gulf and Atlantic coasts of the United States. The adult emits an oily liquid with a repellent odor. Panchlora nivea (Fig. 12).—The Cuban cockroach nymph is reddish brown, and the lateral margins of the thoracic and abdom- 324 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON inal segments appear jagged when viewed dorsally. The 10th tergite is rectangular and longer than the cerci. It is found around houses and wooded areas throughout Flor- ida and along the Gulf Coast of the United States. ACKNOWLEDGMENTS We thank Peter H. Adler (Clemson Uni- versity, Clemson, South Carolina) for his advice with this project. We also thank Mary Ross (Virginia Polytechnic Institute and State University, Blacksburg, Virginia), Richard Brenner (USDA-ARS MAVERL Laboratory, Gainesville, Florida), and Phil Koehler (University of Florida, Gainesville, Florida) for generously providing us with cockroaches. Al Wheeler and Mike Hood, both of Clemson University (Clemson, South Carolina), reviewed and improved this manuscript. This manuscript is techni- cal contribution number 4399 of the South Carolina Agricultural Experiment Station. LITERATURE CITED Blatchley, W. S. 1920. Orthoptera of Northeastern America. Nature Publishing Company, Indianap- olis. 784 pp. Bosik, J. B. 1997. Common Names of Insects & Re- lated Organisms 1997. The Entomological Society of America. Lanham, Maryland. 232 pp. Cornwell, P. B. 1968. The Cockroach Vol. 1. A Lab- oratory Insect and an Industrial Pest. Hutchinson & Co. London. 391 pp. Dakin, M. E., Jr. and K. L. Hays. 1970. A Synopsis of Orthoptera (sensu lato) of Alabama. Auburn Uni- versity Agricultural Experiment Station Bulletin 404, 118 pp. Fisk, EF W. 1987. Order Blattodea, pp. 120-131, In Stehr, EF W., ed., Immature Insects. Kendall/Hunt Publishing, Dubuque, Iowa. Gordon, J. M. 1992. Comparison of mid-instar nymphs of Periplaneta fuliginosa (Serville) and Peripla- neta americana (L.) (Blattodea: Blattidae) Pro- ceedings of the Entomological Society of Wash- ington 94: 151-156. McKittrick, EF A. 1964. Evolutionary Studies of Cock- roaches. Cornell University Agricultural Experi- ment Station. Memoir # 389, 197 pp. Powell, P. K. and W. H. Robinson. 1980. Descriptions and keys to the first-instar nymphs of five Peri- planeta species (Dictyoptera: Blattidae). Proceed- ings of the Entomological Society of Washington 82: 212-228. Pratt, H. D. and C. J. Stojanovich. 1962. Key to the Common Cockroaches found in the United States of Public Health Importance. United States De- partment of Health and Environmental Welfare. Public Health Service. Centers for Disease Con- trol. Princis, K. 1960. Zur Systematik der Blattarien. Eos 36: 429-449. (Authors have not read original work.) Rehn, J. W. H. 1950. A key to the genera of North American Blattaria, including established adven- tives. Entomological News 61(3): 64—67. . 1951. Classification of the Blatteria as indi- cated by their wings (Orthoptera). Memoirs of the American Entomological Society 14: 1-134. Richards, O. W. and R. G. Davies. 1957. A. D. Imms. A General Textbook of Entomology. Ninth Edi- tion. Methuen & Co. Ltd. London. 886 pp. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 325-331 STUDIES ON STONEFLIES OF NORTH DAKOTA WIT THE DESCRIPTION OF A NEW PERLESTA SPECIES (PLECOPTERA: PER LIDAE) Boris C. KONDRATIEFF AND RICHARD W. BAUMANN (BCK) Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, CO 80523, U.S.A. (e-mail: bkondrat @ceres.agsci.colostate.edu); (RWB) Department of Zoology and Monte L. Bean Life Science Museum, Brigham Young University, Provo, UT 84602, U.S.A. (e-mail: richard-baumann @ byu.edu) Abstract.—Records for fifteen stonefly taxa, including eleven confirmed species are given for North Dakota. Literature records for an additional three species could not be confirmed. Perlesta dakota, n. sp., is described from the adult male, female and egg. Diagnostic characters are presented using illustrations and a SEM photomicrograph. Key Words: Only two species of stoneflies were listed for North Dakota by Stark et al. (1986), a state covering 18,299,503 hectares that in- cludes two major river systems, the Mis- souri and the Red River of the North. The landscapes of North Dakota, as common with most Northern Great Plains states, ex- hibit relatively little topography, and the state is better known for its numerous prai- rie marshes (Van der Valk 1989). Many counties of the state have 80—90% of their total area in agricultural production. The Red River forms the eastern border of the state with Minnesota, traversing the former bed of the glacial Lake Agassiz. The area west of the Red River Valley, a region often referred to as the Drift Prairie is poor- ly drained, with few streams of any gradi- ent. West of the Drift Prairie is the Missouri Plateau, also an area of poor drainage, and further west is the Missouri River and its associated tributaries. These streams are of- ten located in broad valleys of native grass- es or hay and small grain production areas. The Little Missouri River is a prominent feature of western North Dakota and drains through the spectacular landscape known as stoneflies, Plecoptera, North Dakota, Perlesta, new species the ‘“‘Badlands.”’ Streams of the Badlands are mostly silty and relatively unproductive (Gordon and Post 1965). A southern exten- sion of the Northern Coniferous Forest can be found in the northwestern portion of the state. Despite the presence of suitable lotic habitats throughout the state, only nine spe- cies of stoneflies have been recorded from North Dakota. Stark et al. (1986) in their compilation of North American stonefly records, listed only Strophopteryx fasciata (Burmeister) and Haploperla orpha (Fri- son). However, we have found seven addi- tional records in the literature. Harden and Mickel (1952) noted a record for Jsoperla longiseta (Banks), and Neel (1985) in his ecological study of the Turtle River in east- ern North Dakota discussed a species of Ac- roneuria, Perlesta placida (Hagen), and Taeniopteryx nivalis (Fitch). Finally, Stoaks (1975) listed nymphal records of Pteronar- cys dorsata (Say), P. placida, A. arenosa (Pictet) and J. bilineata (Say) from the For- est River. A recent collecting trip by the authors into southwestern North Dakota yielded an 326 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Perlesta dakota. 1, Adult head and pronotum. 2, Male terminalia, dorsal. 3, Paraproct, caudal. 4, Paraproct, lateral. 5, Penis. 6, Female subgenital plate, ventral. undescribed species of Perlesta. The genus Perlesta in North America presently in- cludes sixteen species (Stark 1989, Poulton and Stewart 1991, Stark and Rhodes 1997, Kirchner and Kondratieff 1997, DeWalt et al. 1998). The terminology in the descrip- tion follows that proposed by Stark (1989). Perlesta dakota Kondratieff and Baumann, new species (Figs. 1-7). Male.—Forewing length 8-9 mm. Gen- eral color brown. Head yellow with dark brown pattern as Fig. 1. Pronotum dark brown (Fig. 1). Antennal scape and pedicel a VOLUME 101, NUMBER 2 brown, basal flagellular segments becoming darker brown beyond segments 10—16. Fore- femur dorsally brown. Wing membrane and veins brown except costal margin pale. Ab- dominal terga brown, sterna yellow brown usually with darker brown triangle-shaped shading. Cercus yellow brown. Tergum 10 mesal sclerite not divided, sensilla basicon- ica small and scattered (Fig. 2). Paraproct short in caudal view (Fig. 3), more slender in lateral view (Fig. 4), spine anteapical and directed mesad (Figs. 2 and 3). Penis tube + sac moderately long with raised shelf dorsobasally; caecum prominent, dorsal hair patch broad basally narrowing to base of caecum, lateral hair patch elongate (Fig. 5). Female. Forewing length 10-12 mm. Color pattern similar to male but paler. Sub- genital plate truncate with notch deep and narrowly V-shaped (Fig. 6). Egg.—Collar button-like. Chorion sur- face finely pitted with scattered coarser pit- ting. Micropylar row in posterior region (Fig. 7). Nymph.—Unknown. Types.—Holotype ¢, allotype 2: North Dakota, Hettinger Co., Cannonball River, at New England, Hwy 22, 15 July 1997, R. W. Baumann and B. C. Kondratieff. Para- types, same data as holotype: 14 6,9 9; Ransom Co., Fort Ransom State Park, 11 July 1970s S. -M. “Anders, 16, 1 ¢ (NDSU); Stark Co., Heart River, Hwy 22, Dickinson, 14 July 1997, R. W. Baumann and B.'@> "Kondratieff, 2 2° (BYUC, CSUC). Holotype and allotype deposited in the National Museum of Natural History, Smithsonian Institution, other paratypes at the Monte L. Bean Life Sciences Museum, Brigham Young University (BYUC); C. P. Gillette Museum of Arthropod Diversity, Colorado State University (CSUC); North Dakota State University (NDSU) and the B. P. Stark Collection (BPSC). Etymology.—The Sioux were the largest tribe of Native Americans of the North American Plains and prairies, and are called the Dakota in the Santee dialect. Diagnosis.—Perlesta dakota appears | sua Wee micrograph of egg. Perlesta dakota, scanning electron photo- similar to a group of dark species, including P. cinctipes (Banks), P. adena Stark, P. fus- ca Poulton and Stewart, and P. xube Stark and Rhodes. Perlesta dakota seems most similar to P. fusca, a species distributed throughout the Ozark-Ouachita Mountain region of Arkansas, Missouri and Oklahoma (Poulton and Stewart 1991). Males of P. dakota can be distinguished from P. fusca by the well developed thumb- like caecum (Fig. 5) and maculation of the head (Fig. 1). The penis structure of P. dak- ota is similar to P. decipiens (See Stark 1989; Figs. 42—43) but the former species can be distinguished by the dark brown wing and body coloration and head pattern. Females of P. dakota can be distinguished from P. fusca by the more truncate lobes of the subgenital plate and the deep and nar- row notch (Fig. 6). The egg collar of P. dakota is button-like (Fig. 7), and the cho- rion finely pitted (Fig. 8). Poulton and Stewart (1991) indicated that the egg of P. fusca lacks a collar and the surface of the 328 egg is reticulate. The egg of P. decipiens has a distinctive short collar. Remarks.—The type locality, the Can- nonball River at New England, is typical of an agriculturally-impacted stream of the re- gion, little or no riparian vegetation, heavily silted and with only few small riffles. Adults of P. dakota were collected by beat- ing tall overhanging streamside grasses. Other aquatic insects collected with P. dak- ota include the damselfly Calopteryx ae- quabilis Say, the mayfly Caenis sp., and the caddisflies, Cheumatopsyche pettiti (Banks), Ceratopsyche morosa (Hagen), Hydroptila consimilis Morton, and Limnephilius hyali- nus Hagen. North Dakota Stonefly Records Collections examined, primarily from North Dakota State University (NDSU), yielded new records or added to or substan- tiated previous records of North Dakota stoneflies. Other institutions listed include Brigham Young University (BYUC), Col- orado State University (CSUC); Dickinson State University (DSUC); Minot State Uni- versity (MSUM); Mississippi College (BPSC); Museum of Comparative Zoology, Harvard University (MCZC), University of Minnesota (UMSP), and University of Mis- sissippi (UMIC). Some of these records are only identified to the generic level, but are included for distributional purposes, since so few stonefly records are currently avail- able for North Dakota. A total of fifteen stonefly taxa are known from the state of North Dakota, representing at least eleven species. Allocapnia sp. Records.—Ransom Co., cold spring-fed stream, joining Sheyenne River, Hwy 46, Little Yellowstone Park, 1 Nov 1962, R. D. Gordon, 6 N (NDSU). Several species of this genus of winter stoneflies could occur in North Dakota, es- pecially A. pygmaea (Burmeister) (Ross and Ricker 1971). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Oemopteryx fosketti (Ricker) Records.—Billings Co., Little Missouri River, Medora, 19 March 1997, C. P. Milne, IG 35 672 (BYUCG ESUG, DSUCG)2 Little Missouri River, Sully Creek Campground, 19=March, 1997— C2 Paine. Wi Greleee (BYUC). The species originally described from Saskatchewan, Canada, can be abundant in large and often silty rivers of the Missouri, Colorado, and Saskatchewan drainages (Baumann et al. 1977). Records for this species are known from nearby Dawson Co., Montana. The males have peculiar up- turned forewing tips. Strophopteryx fasciata (Burmeister) Records.—Cass Co., Fargo, 24 April 1900, E. Cleveland, 1 6 (MCZC). Strophopteryx fasciata is a widespread species occurring throughout eastern and midwestern North America, and typically a larger stream species emerging in late win- ter and early spring (Stewart and Stark 1988). The record for 7. nivalis by Stoaks (1975) could not be confirmed due to the lack of material, but is possible, since this species is common in Minnesota (Ricker and Ross 1968). Pteronarcys pictetii Hagen Records.—North Dakota: Cavalier Co., 20 Oct 1962, 2 N (NDSU); Pembina Co., Pembina River, Walhalla City Park, | July 1970, Perkins and R. L. Post, 1 2 (NDSU); Pembina River, Walhalla, 9 July 1961, R. L. Post and H. Osborn, 4 N (NDSU); same but 20 Oct 1962, E. Saugstad, 2 N (NDSU); Richland Co., Sheyenne River, 14 mi NW Walcott, 24 June 1963, D. Aarhus, 1 2 (NDSU). Pteronarcys pictetii has been reported from adjacent Minnesota and Manitoba (Stark et al. 1986), and was expected in the state. The separation of the nymphs of P. dorsata (Say) and P. pictetii is difficult, es- pecially immature specimens. The transcon- VOLUME 101, NUMBER 2 tinental species, P. dorsata has been re- ported from all surrounding states and Ca- nadian provinces except South Dakota (Stark et al. 1986). Pteronarcys sp. Records.—Cass Co., Stearn, 25 Aug 1960, 1 N (NDSU); Cavalier Co., 20 Oct 1962, | N (NDSU); Grand Forks Co., Tur- tle River State Park, 8 Aug 1962, R. D. Gordon, 2 N (NDSU); Forest River, Hwy 18, 3 mi NE Inkster, | July 1970, R. Stoaks, 1 N (NDSU); same but 25 Aug 1970, 1 N (NDSU); same but 19 June 1971, 1 N (NDSU)s same, but 28 Aug 1971, 3. .N (NDSUW)s¢same. but. 15° Sept. 1991, 3. IN (NDSU); Ward Co., Mouse River, Nedrose #1, SE Minot, 13 July 1957, R. Nelson, 1 2 (MSUM). The nymphs collected by Ralph D. Stoaks are apparently the specimens listed by him as P. dorsata, a species that remains unconfirmed for the state. The adult female listed above could not be determined to spe- cies. Acroneuria abnormis (Newman) Records.—Richland Co., Mirror Pool, Sheyenne River, 23 June 1975, P. K. Lago, 1 d (UMIC). This widespread species was expected in North Dakota, and is known from all sur- rounding states and Canadian provinces (Stark et al. 1986, Huntsman et al. 1999). Acroneuria lycorias (Newman) Records.—Cass Co., Sheyenne River, 5 mi E Kindred, 28 June 1996, K. Mundal, 2 3, 3 2 (NDSU), same but 30 June 1996, K. Mundal, 2 ¢, 3 2 (NDSU); Grand Forks Co., Forest River, Hwy 18, 3 mi NE Inkster, 16 Oct 1971, R. Stoaks, 1 N (NDSU); Pem- bina Co., Pembina River, Walhalla, 9 Sept 1961, R. L. Post and H. Osborn, 14 N (NDSU); same but 20 Oct 1962, E. Saug- stad, 2 N (NDSU); same but 20 Oct 1996, D. G. Aarhus, 1 N (NDSU). This widespread eastern and upper mid- western Nearctic species has been recorded 329 from adjacent Minnesota and Manitoba. Harden and Mickel (1952) indicated that this species is one of the few stoneflies that occur in the western prairie regions of Min- nesota. Acroneuria sp. Records.—Pembina Co., Pembina River, Walhalla, 9 Aug 1961, H. Osborn, 1 N (NDSU); Walsh Co., Forest River, USGS Gage, 3: mi SE Fordville; 15)Sept 1971, R. D. Stoaks, 1 N (NDSU). Other possible species of Acroneuria re- ported from adjacent states and Canadian provinces include A. carolinensis (Banks) and A. internata (Walker) (Stark et al. 1986). Stoaks (1975) listed A. arenosa, however, this nymphal determination is doubtful since this species is restricted to the eastern U.S. (Stark and Gaufin 1976). Perlesta decipiens (Walsh) Records.—Cass Co., Sheyenne River, 5 mi E Kindred, 28 June 1996, K. Mundal, 3 2? (NDSU); NDAC, Fargo, USDA UV light trap, 6 July 1956, 1 2 (NDSU); Richland Cox, 13’ July 1962; 1 ¢, 4 2 (NDSU); Trail Co., Elm River, 8 Aug 1969, R. L. Post, 1 3d (NDSU). The occurrence of this geographically widespread species was expected in the state. Perlesta xube Stark and Rhodes Records.—Walsh Co., Forest River, Hwy 35, 8 July 1966,,R° LE: Post;'2°6, 2 233) N (NDSU). Perlesta xube was recently described from a small stream in Cherry County, Ne- braska (Stark and Rhodes 1997). This rec- ord from the Forest River represents a northeastern range extension, indicating that this species may occur in remnant stream systems throughout the northern Great Plains. Perlesta sp. Records.—Cass Co., Fargo, 13 July 1973, P. K. Lago, 1 2 (UMIC); Dunn Co., 330 Knife River, Manning, 29 May 1991, Mott, 1 N (DSUC); Grand Forks Co., Turtle Riv- er, 23 June 1970, D. M. Huntsinger, 1 N (NDSU); Forest River, Hwy 18, 3 mi NE Inkster, 23 June 1970, R. D. Stoaks, 4 N (NDSU); Slope Co., Burning Coal Vein, 0.25 mi W Logging Camp Ranch, Little Missouri National Grassland, 6 July 1968, R. D. Stoaks, 6 N (NDSU); Walsh Co., For- est River, 3.5 mi W Fordville, Hwy 32, 19 June 1971, R. D. Stoaks, 31 N (NDSU); same but 11 July 1971, R. D. Stoaks, 4 N (NDSU); same but at USGS gage, 19 May 1971, R. D. Stoaks, 1 N (NDSU). Nymphs of Perlesta are difficult to spe- cifically distinguish, and much of the above material is poorly preserved or represents early instars. Stoaks (1975) and Neel (1985) reported P. placida as nymphs and as nymphs and adults, respectively. Stark (1989) determined that P. placida was a complex of at least twelve (now sixteen) species. No adults from Neel (1985) were available for study. With no associated males, the single female adult from Fargo was left undetermined. Isoperla bilineata (Say) Records.—Cass Co., Fargo, NDSU Cam- pus:.7 June, 1963, R: WW. Poole, 7 io, 1°92 (NDSU); same but 16 June 1963, R. W. Poole, 7 gd, 1 2 (NDSU) same but 18 June 1993, 6 6 (NDSU); Sheyenne River, 5 mi E Kindred, 30 June 1996, K. Mundal, | ¢, 4 2 (NDSU); NDAC, Fargo, USDA UV light trap, 9 June 1956, 20 d, 3 2 (NDSU). Emmons Co., 16 mi W Linton, 20 June L975;2P. K Lago.2 6 (UMIC); Grant,Co:; Heart Butte Dam, 19 June 1975, P. K. and B. A. Lago, 2 6 (UMIC); Pembina Co., Tongue River, Kotchman Farm, near Cav- alier, 29 June 1974, P K. and B. A. Lago, 3 6 (UMIC); same but 2 July 1975, 2 ¢ (UMIC). Stoaks (1975) previously reported this species, but no specimens from his study were available for examination. Szczytko and Stewart (1978) indicated material from PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the Red River of the North, Minnesota, which forms the border with North Dakota. Isoperla longiseta (Banks) Records.—Billings Co., Little Missouri River, Elkhorn Ranch, 20 June 1965, R. L. Post, 1 2 (NDS); Cass"Cos Parco, 1M May 1939, D. G. Denning, 1 2 (UMSP); Slope Co., Little Missouri River, at Mar- marth, Hwy 12, 14 July 1997, R. W. Bau- mann and B.C: Kondratiefii, (6465.72 (BYUG, CSUCG): Isoperla longiseta is known from west- ern and midwestern North America, and oc- curs further east than any other typically western Jsoperla (Szczytko and Stewart 1979). It is often considered a typical prai- rie stonefly of larger streams and rivers (Harden and Mickel 1952, Ricker 1946, 1964). Harden and Mickel (1952) previous- ly mentioned the 1939 record of J. longiseta from Fargo. Haploperla orpha (Frison) Records.—Pembina Co., Tongue River, Kotchman Farm near Cavalier, 21 June 1974. PK anduB. -Anskago! Qedeesee, (BPSG); same. but) 23) July 1975, id. See (UMIC). Haploperla orpha is a relatively poorly known species having also been recorded from Maine, Minnesota, Quebec, New Brunswick, and Wisconsin (Stark et al. 1986). Typically, nymphs of this species are found in medium and small-sized streams with some gradient. ACKNOWLEDGMENTS We thank Dr. B. P. Stark for reviewing the manuscript, providing the SEM photo- graphs and material. Drs. P. J. Clausen, Uni- versity of Minnesota; R. D. Gordon, Wil- low City, North Dakota; P. K. Lago, Uni- versity of Mississippi; C. P Milne, Jr., Dickinson State University; D. A. Rider, North Dakota State University; R. A. Roy- er, Minot State University; and P. D. Per- kins, Museum of Comparative Zoology, VOLUME 101, NUMBER 2 Harvard University, are thanked for loaning valuable material. Special thanks are given to Charles P. Milne, Jr. for making a concerted effort to collect winter stonefly species, which re- sulted in the discovery of Oemopteryx fos- ketti. Dr. Gerald M. Fauske, NDSU, helped us locate questionable localities from across the state. LITERATURE CITED Baumann, R. W., A. R. Gaufin, and R. F Surdick. 1977. The stoneflies (Plecoptera) of the Rocky Mountains. Memoirs of the American Entomolog- ical Society 31, 208 pp. DeWalt, R. E., B. P. Stark, and M. A. Harris. 1998. Perlesta golconda (Plecoptera: Perlidae), a new stonefly species from Illinois. Entomological News 109: 315-317. Gordon, R. D. and R. L. Post. 1965. North Dakota water beetles. North Dakota Insects. Publication 5, Agricultural Experiment Station, North Dakota State University 53 pp. Harden, P. H. and C. E. Mickel. 1952. The stoneflies of Minnesota (Plecoptera). University of Minne- sota Agricultural Experiment Station, Technical Bulletin 201. 82 pp. Huntsman, B. O., R. W. Baumann, and B. C. Kondra- tieff. 1999. The stoneflies (Plecoptera) of the Black Hills of South Dakota and Wyoming, USA: Distribution and zoogeographic affinities. Great Basin Naturalist 59: 1-17. Kirchner, R. E and B. C. Kondratieff. 1997. A new species of Nearctic Perlesta (Plecoptera: Perlidae) from Virginia. Proceedings of the Entomological Society of Washington 99: 290-293. Neel, J. K., Sr. 1985. A northern prairie stream. Uni- versity of North Dakota Press, Grand Forks. 274 PPp- Poulton, B. C. and K. W. Stewart. 1991. The stoneflies of the Ozark and Ouachita Mountains (Plecop- tera). Memoirs of the American Entomological Society 38, 116 pp. Ricker, W. E. 1946. Some prairie stoneflies (Plecop- 331 tera). Transactions of the Royal Canadian Institute 26: 3-8. . 1964. Distribution of Canadian stoneflies, pp. 50-71. In Illies, J. ed., 3. Internationales Sympo- sium tiber Plecopteren. August. Gewasser und Ab- wasser, Bagel, Dusseldorf. 138 pp. Ricker, W. E. and H. H. Ross. 1968. North American species of Taeniopteryx (Plecoptera: Insecta). Journal of the Fisheries Research Board of Canada 25: 1423-1439. Ross, H. H. and W. E. Ricker. 1971. The classification, evolution, and dispersal of the winter stonefly ge- nus Allocapnia. Illinois Biological Monograph 45, 166 pp. Stark, B. P. 1989. Perlesta placida (Hagen), an eastern Nearctic species complex (Plecoptera: Perlidae). Entomologica Scandinavica 20: 263—286. Stark, B. P. and A. R. Gaufin. 1976. The Nearctic spe- cies of Acroneuria (Plecoptera: Perlidae). Journal of the Kansas Entomological Society 49: 221- DS Stark, B. P. and H. A. Rhodes. 1997. Perlesta xube, a new stonefly species from Nebraska (Plecoptera: Perlidae). Entomological News 108: 92-96. Stark, B. P, S. W. Szczytko, and R. W. Baumann. 1986. North American stoneflies (Plecoptera): sys- tematics, distribution, and taxonomic references. Great Basin Naturalist 46: 383-397. Stewart, K. W. and B. P. Stark. 1988. Nymphs of North American stonefly genera (Plecoptera). Thomas Say Foundation, Entomological Society of Amer- ica. 12, 460 pp. Stoaks, R. D. 1975. Seasonal and spatial distribution of riffle dwelling aquatic insects in the Forest Riv- er, North Dakota. Ph.D. dissertation, North Dakota State University, Fargo. 163 pp. Szcezytko, S. W. and K. W. Stewart. 1978. /soperla bilineata: designation of a neotype and allotype, and further descriptions of egg and nymph. An- nals of the Entomological Society of America 71: 212-217. 1979. The genus /soperla (Plecoptera) of western North America; holomorphology and sys- tematics, and a new stonefly genus Cascadoperla. Memoirs of the American Entomological Society. 323 120° pp: Van der Valk, A. 1989. Northern Prairie Wetlands. Iowa State University Press, Ames. 400 pp. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 332-336 A NEW NEARCTIC SPECIES OF LESTODIPLOSIS (DIPTERA: CECIDOMYHDAE) PREYING ON AN OAK LEAF TIER, PSILOCORSIS QUERCICELLA (LEPIDOPTERA: OECOPHORIDAE) RAYMOND J. GAGNE AND JOHN T. LILL (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); (JTL) Department of Biology, University of Missouri-St. Louis, St. Louis, MO 63121, U.S.A. Abstract.—A new species of cecidomyiid, Lestodiplosis venusta Gagné, is described, illustrated, and compared to its most similar congeners. Larvae of the new species were found in Missouri preying on caterpillars of Psilocorsis quercicella Clemens, a leaf skel- etonizer of oaks. An unidentified species of Ceraphronidae was reared from pupae of the Lestodiplosis. Key Words: A new species of Lestodiplosis attacking larvae of the oecophorid Psilocorsis quer- cicella Clemens was discovered by one of us (JTL) during the course of an ecological study of the natural enemies of the lepidop- teran in Missouri. Psilocorsis quercicella is a common bivoltine moth found throughout eastern United States (Covell 1984). The larvae skeletonize the surface of oak leaves within leaf shelters formed by tying togeth- er adjacent oak leaves with silk. Larvae of the gall midge predator appear to be highly mobile ectoparasites of early instar P. quer- cicella larvae and are often gregarious. When present, one to four of the preda- ceous cecidomyiids were found per cater- pillar. The predator was found feeding on P. quercicella on both white oak (Quercus alba L.) and black oak (Q. velutina Lam.). After feeding, full grown Lestodiplosis \ar- vae spin a silken cocoon within the leaf shelter, often incorporating pieces of cater- pillar frass onto the surface of the cocoons. To date, larvae of the predator have been observed only from second generation P. Lestodiplosis, predator, oak leaf tier quercicella larvae collected in late summer and fall in Missouri. An unidentified gre- garious parasitoid (Hymenoptera: Cera- phonidae) has been reared from the Lesto- diplosis pupae. The genus Lestodiplosis contains some 175 known species in the world that attack many kinds of insects and mites. Some ap- pear to be specialist predators, others are evidently generalists. Many species are known from only one or a few specimens so it is especially helpful to have several series of the same species from a particular niche, such as JTL found in leaf ties of P. quercicella. METHODS Larvae of the predator were reared to the adult stage in 16-0z. closed clear plastic containers kept at 23°C and with a photo- period of 14:10 (L:D) h. Humidity was maintained with moist filter paper. Speci- mens of immature stages and reared adults were preserved in 70% isopropyl alcohol. Specimens were mounted on microscope VOLUME 101, NUMBER 2 slides using the method outlined in Gagné (1989). Terminology for adult morphology follows usage in McAlpine et al. (1981) and for larval morphology that in Gagné (1989). Lestodiplosis venusta Gagné, new species (Figs. 1-13) Adult.—Head: Eyes connate, 11—12 fac- ets long at vertex; facets hexagonal, all closely adjacent. Occiput with dorsal pro- tuberance. Frons with 10—14 setae. Labella hemispherical but pointed apically, each with several lateral setae. Palpus 4-seg- mented. Male antennal flagellomeres (Fig. 1) binodal; basal node and the distal third of the neck dark; one circumfilum on the basal node, two on the distal, the loops of the three circumfila unequal in length; the proximal and distal circumfila with ventral- most loops greatly elongated and much lon- ger than remaining loops. Female flagel- lomeres (Fig. 2) with basal part of node and distal part of neck darker than remainder of flagellomere; circumfila with some short loops. Thorax: Wing (Fig. 3) 1.8—2.3 mm long, R, slightly curved, joining C posterior to wing apex; 7—8 irregular dark spots present on wing. Legs (Fig. 4) with alternating light and dark groups of scales. Tarsal claws (Fig. 5) untoothed, curved beyond mid- length; empodia attaining bend in claws. Male abdomen: First through seventh tergites entire, rectangular, with single pos- terior row of setae, several lateral setae, scattered scales, and 2 anterior trichoid sen- silla; eighth tergite undifferentiated, the only vestiture the anterior pair of trichoid sensilla. First through seventh sternites quadrate, with single to double posterior row of setae, scattered setae elsewhere, no scales, and 2 anterior trichoid sensilla; eighth sternite similar to preceding except weakly sclerotized anterolaterally. Genitalia (Figs. 6-8): cerci each with rounded pos- terior margin; hypoproct simple, rounded at apex, as long as but slightly wider in ventral view than cerci, with 2 pairs of apicoventral 333 setae, produced anteroventrally into large lobe lying in close juxtaposition with dorsal surface of aedeagus and covered posteriorly with short spinules; aedeagus elongate with blunt apex, sinuous in side view: gonocox- ite elongate cylindrical with short, triangu- lar mesobasal lobe bearing short spinules; gonostylus elongate cylindrical, with setu- lae near base and covered with minute ca- rinae and only several short setae beyond. Female abdomen (Figs. 9-11): First through seventh tergites entire, rectangular, with mostly single row of posterior setae, several lateral setae, extensively covered with scales, and 2 anterior trichoid sensilla. Second through seventh sternites quadrate, extensively covered with setae and scales and with anterior pair of trichoid sensilla. Eighth segment much narrowed, without differentiated tergite and sternite, the ter- gum with anterior pair of trichoid sensilla the only vestiture, the sternum with anterior pair of trichoid sensilla and covered with setae some distance beyond. Tenth segment without vestiture dorsally, with scattered se- tae ventrally; cercus large, ovoid, with ven- tral field of short, closely-set, sensory setae, and scattered setae elsewhere. Hypoproct short, wide, the convex posterior edge with 2 short setae. Apex of ninth sternite pro- truding posteriorly below hypoproct, rigid, triangular. Third larval instar (Figs. 12—14).—Integ- ument mostly smooth, 2 pseudopods pres- ent ventrally on mesothorax and metathorax and 3 each on first through seventh abdom- inal segments. Antenna much longer than wide. Cephalic apodemes slightly longer than head capsule. Spatula absent. Lateral papillae in 2 groups of 3 on each side of central line, only one setose in each group. Second and fifth dorsal papillae of each segment much shorter than remaining four. Terminal segment with 6 elongate setae, usually with expanded apices. Holotype.—<¢, reared from larva preying on larva of Psilocorsis quercicella on oak, Cuivre River State Park, Troy, Missouri, 1- IX-1997, J. Lill, deposited in the National 334 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-8. Lestodiplosis venusta. 1, Male third antennal flagellomere. 2, Female third antennal flagellomere. 3, Wing. 4, Foreleg. 5, Tarsal claw and empodium. 6, Gonopod, cerci, hypoproct, and aedeagus (dorsal). 7, Cercus, hypoproct, and aedeagus (lateral). 8, Hypoproct (ventral). VOLUME 101, NUMBER 2 335 13 14 Figs. 9-14. Lestodiplosis venusta. 9-11, Female postabdomen. 9, Seventh tergite to cerci (not all setal bases and setae indicated: dorsal). 10, Cerci, hypoproct and apex of ninth sternum, the postabdomen retracted (lateral). 11, Seventh sternite to cerci (not all setal bases and setae indicated; ventral). 12-14, Larva. 12, Head, neck, and first two thoracic segments (dorsal). 13, part of mesothorax with two groups of three lateral papillae, a ventral papilla, and two pseudopods (ventrolateral). 14, Apex of seventh and all of eighth and terminal abdominal segments (dorsal). 336 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Museum of Natural History (USNM), Washington, DC. Other material examined.—All associat- ed with leaf ties of Psilocorsis quercicella, Cuivre River State Park, Troy, Missouri: 6 3,5 ¢, 9 larvae, same data as holotype; 6 3d, 1 2, same data as holotype except col- lected 11-IX-1996, all in USNM. Etymology.—The specific name, venus- ta, is an adjective meaning beautiful, with reference to the striking black and white banded legs and antenna. Remarks.—The banded antenna and legs and the spotted wings of this species are generally similar to some other Lestodi- plosis species, but the combination of un- equal male circumfila and a blunt-tipped ae- deagus is characteristic in the Nearctic Re- gion of a narrower group of three species that differ chiefly in details of the male gen- italia. These three species are: Lestodiplosis florida Felt, Lestodiplosis cinctipes (Felt), and Lestodiplosis satiata Felt. These other species are now under investigation by one of us (RJG) for a revision in progress of Nearctic Lestodiplosis. Compared to the new species, the ventral hypoproctal lobe of L. florida is covered by fewer but more robust spinules and its ae- deagus is much wider on the basal two- thirds and tapers abruptly to a narrow neck. Lestodiplosis florida was originally taken from a leaf roll on a Crataegus in Florida. This species is represented in the USNM collection by a specimen from Florida that was reared from galls on Croton linearis Jacq. and another that was swept from Tsu- ga canadensis (L.) Carr. in Ontario. The ventral lobe of the hypoproct of both L. cinctipes and L. satiata is covered more extensively and by finer spinules than in the new species and the aedeagus in these spe- cies is narrower throughout its length than on L. venusta. In addition, the antennal cir- cumfila loops of both species are all much longer than in L. venusta. Both species were originally caught in flight in New York, but a specimen of L. satiata in the USNM was reared in association with cynipine leaf galls on Quercus sp. in Pennsylvania and another was reared from flower heads of Borrichia frutescens (L.) DC. in North Car- olina. There is one other Nearctic Lestodiplosis species, besides L. venusta, that has been recorded as feeding on caterpillars. Felt (1933) described Lestodiplosis novangliae Felt from specimens associated with a tor- tricid Epinotia ‘“‘nanana’’ on spruce in Massachusetts. He noted also that larvae similar to that species were found on an- other tortricid, Rhyacionia sp. on pine. The gonocoxites of L. novangliae, unlike those of L. venusta, have prominent and spiny mesobasal lobes. ACKNOWLEDGMENTS We thank B. Schuette and the Missouri Department of Natural Resources for logis- tical support and collection permits, P. Ma- likul for making the slide preparations, C. Anderson for computer assistance in mak- ing the plates, and, for their comments on drafts of the manuscript: P. Kolesik, Uni- versity of Adelaide, South Australia; A. L. Norrbom of the Systematic Entomology Laboratory, USDA, Washington, D.C.; M. Solinas, Universita degli Studi di Perugia, Italy; and J. Yukawa, Kyushu University, Fukuoka, Japan. The field research was funded with a National Science Foundation Dissertation Improvement Award (DEB- 9700887) to JTL. LITERATURE CITED Covell, C. V., Jr. 1984. A Field Guide to the Moths of Eastern North America. Houghton Mifflin Co., Boston. Felt, E. P. 1933. A new Lestodiplosis. Psyche 40: 113-— 114. Gagné, R. J. 1989. The Plant-Feeding Gall Midges of North America. Cornell University Press, Ithaca, New York. xiii + 355 pp. & 4 pls. 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. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 337-346 DESCRIPTION OF TWO NEW SPECIES OF AMBLYCERUS YAUNBERG (COLEOPTERA: BRUCHIDAE) WITH A PROBABL© STRIDULATORY MECHANISM CIBELE S. RIBEIRO-COSTA Departamento de Zoologia, Universidade Federal do Parana, C. P. 19020, 81.531-990, Curitiba, Parana, Brazil (e-mail: stra@bio.ufpr.br) Abstract.—Amblycerus atypicus, new species, and A. ischiodontus, new species, are described and illustrated. The former has an area with overlapping scales in part of the external elytral margins and the inner distal portion of the hind femur with fine transverse striation. Amblycerus ischiodontus has an area transversely striate on the metepisternum and an apical tooth on the ventral margin of the hind femur. The areas on the elytra and metepisternum probably act as a file and the striate area or the tooth of the hind femur as the scraper. The elytro-femoral method of stridulation is postulated for the first time in Bruchidae and is shared with Amblycerus cistelinus (Gyllenhal), A. jatayensis (Pic), A. sosia Ribeiro-Costa and Kingsolver, A. whiteheadi Kingsolver and A. guazumicola King- solver and Johnson. The other kind, involving the metepisternum and hind femur, has already been described for three other Amblycerus species: A. eustrophoides (Schaeffer), A. pollens (Sharp), and A. stridulator Kingsolver, Romero N., and Johnson. A key for the bruchid species with modified body areas probably involved in stridulation is presented. Key Words: Kingsolver (1970) first recorded the pres- ence of areas of the integument probably involved in stridulation in Bruchidae. He indicated the presence of a fusiform node with transverse striations on the metepister- num and the presence of an apical blunt tooth on the ventral margin of the hind fe- mur for Amblycerus eustrophoides (Shaef- fer), which has a distribution restricted to North America. Kingsolver et al. (1993) no- ticed the same stridulatory areas for A. stri- dulator described from Mexico, Costa Rica, and Venezuela and for A. pollens (Sharp) recorded from Belize, Costa Rica, and Bra- zl. Amblycerus atypicus, n. sp. (French Gui- ana), has an area with overlapping scales on the external margins of the elytra and fine transverse striation on the inner distal por- Amblycerus, stridulation, taxonomy, Bruchidae tion of the hind femur. The other new spe- cies, Amblycerus ischiodontus (Brazil), has the metepisternum and hind femur with ar- eas transversely striate, similar to those mentioned for A. eustrophoides, A. pollens, and A. stridulator. These areas for each spe- cies are in contact when the hind leg is moved, and, even though no sound emis- sion has been perceptible from dead speci- mens, it is possible that they form a strid- ulatory mechanism. Since males and fe- males of both species have these areas, I hypothesize that the mechanism probably is not related with courtship behavior. The type of stridulation in A. atypicus can be characterized as the elytro-femoral method, established by Dumortier (1963). The area with overlapping scales on the el- ytron would be the ‘‘pars stridens”’ (file) 338 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and the striate area of the hind femur, the ‘““plectrum’’ (scraper). Dumortier (1963) cited the presence of the elytro-femoral method for the Coleopteran families Scar- abaeidae, Lucanidae, Cerambycidae, Cicin- delidae, Tenebrionidae, and Carabidae. This method of stridulation is hypothesized for the first time in Bruchidae and is shared with the following Amblycerus Thunberg species: A. cistelinus (Gyllenhal), A. jatay- ensis (Pic), A. sosia Ribeiro-Costa and Kingsolver, A. whiteheadi Kingsolver, and A. guazumicola Kingsolver and Johnson. The methodology used for the descrip- tions follows Ribeiro-Costa (1997). Amblycerus atypicus Ribeiro-Costa, new species (Figs. 1—10) Description.—Measurements (pronotum + elytra): Length 8.17—8.83 mm; width 4.83—5.17 mm. Integument: Antenna dark red. Dorsal surface (Fig. 1) dark red with scattered black spots on elytra, two on basal one- third, four at middle one-third and two on apical one-third. Lateral margins of elytra gently darker. Undersurface and legs dark red. Pygidium with large, ovate, central, dark brown spot (Fig. 4). Vestiture: General coloration on dorsal surface golden yellow (Fig. 1). Pronotum with white setae forming six small, round spots arranged in triangle in middle lateral and scattered patches along basal lobe (Fig. 1). Scutellum usually white. Elytra (Fig. 1) with white setae in scattered patches along basal margin and around scutellum, often in a few small, irregular spots on median and external margins, in lines along interstitial margins and strial sulci, and condensed into a few, very small irregular patches on re- mainder of elytra. Pygidium (Fig. 4) with large, ovate, velvety dark brown spot, bor- dered with dense white setae and with two small patches of dark brown pillosity in an- terolateral areas. Undersurface gently mot- tled brown and white. Lateral margins of abdominal sterna with white setae gently condensed into round patches. Frons slightly flat, evenly punctate ex- cept on frontal carina. Eyes moderately fac- eted (3 ommatidia), protruding laterally; oc- ular index 3.9; ocular sinus 1/3.8 diameter of eye in lateral view; postocular lobe 1/9.5 times largest length of eye in lateral view. Antenna (Fig. 5) serrate from fourth to tenth segments, all gently longer than wide (1.1 times); last segment elliptical. Prono- tum trapezoidal with lateral margins slight- ly arched in dorsal view, cervical sulcus present in lateral third, a sulcus outlining the emargination of basal lobe; long lateral carina, nearly reaching apex of pronotum, not forked anteriorly; surface densely punc- tulate, lateral one-third on either side also coarsely punctate. Prosternal process wide, margins sulcate, apex subacute, moderately expanded exceeding fore coxae and fitting into sulcate mesosternum. Scutellum (Fig. 2) about 1.9 times as long as wide, with tridentate apex. Elytra slightly convex in cross section along elytral suture, with an area with overlapping scales in part of the external margins (Fig. 9), truncated apically and with striae strongly impressed. Mete- pisternum with some punctures moderately, sparsely coarser; transverse axis of mete- pisternal sulcus (Fig. 3) strongly divergent from metapleural suture, very short longi- tudinal axis, less than half metepisternal length, not modified in a fusiform node with transverse striations (Fig. 3). Hind coxa with punctures moderately coarse and slightly dispersed, except in basal third. Hind femur (Fig. 6) 2.8 times as long as wide; apical tooth on ventral margin absent; internal face on distal portion with trans- verse striations (Figs. 6, 10). Hind tibia (Fig. 6) with coronal teeth approximately of same size; lateral spur 2.9 times length of median, first hind tarsal segment about 1.2 times as long as lateral spur and 3.6 times median spur; ventral face slightly convex with inconspicuous lines of punctures and setae in margins. Male pygidium vertical in lateral view, female oblique; male eighth VOLUME 101, NUMBER 2 339 my] Dark red integument extsh! Golden yellow hairs a Black integument a1 i Golden yellow hairs a Dark red integument White hairs ey Dark brown integument Dark brown hairs ) Imm wif {Ty a ane a i ih air alh iW Ata) hued Ni ui bil i LN Figs. 1-6. Amblycerus atypicus. 1, Dorsal habitus. 2, Scutellum. 3, Lateral view of head and thorax. 4, Pygidium. 5, Antenna. 6, Hind leg. 340 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON awry 500-4 Figs. 7-8. Amblycerus atypicus. 7, Male terminalia, median lobe. 8, Same, tegmen and lateral lobes. tergite acute. Fifth visible abdominal ster- num of female longer than that of male at middle, margin in both sexes entire. Male terminalia (Figs. 7, 8): Eighth ter- gite acute apically. Median lobe (Fig. 7) with length 4.2 times its largest width in basal area; ventral valve moderately long, acute apically, with straight lateral margins; dorsal valve rounded. Basal area of internal sac (Fig. 7) without anterior and median sclerites; a pair of posterior tooth-like scler- ites. Median region of internal sac (Fig. 7) VOLUME 101, NUMBER 2 with two central, long laminar sclerites each with angulate apical portion and each one with one or two teeth near base, one sub- basal and two on median portion; pair of sclerites laterad to laminars, longer than wide, each with one small curved tooth di- rected upward; unpaired median sclerite, 0.3 times as long as laminars, slightly sin- uate in lateral view, with one side serrate. In apical area of internal sac (Fig. 7) one sclerite with long stems among many den- ticles. Tegmen with shallow emargination among enlarged lateral lobes (Fig. 8). Discussion.—Romero et al. (1996) estab- lished the cistelinus group including A. cis- telinus, A. sosia, and A. guazumicola main- ly considering both the integument and pu- bescence pattern on the dorsal surface, not mentioning the presence of areas on elytra or hind femur possibly involved in stridu- lation. Amblycerus atypicus (French Gui- ana), A. jatayensis (Brazil) and A. white- headi (Panama, Nicaragua, Costa Rica, Co- lombia), probably will be included in this group (revisionary study currently under- way) because they share characters cited by Romero et al. (1996) and also the areas on the external margins of the elytra with over- lapping scales (Fig. 9) and transverse stri- ation on the inner distal portion of hind fe- mur (Fig. 10), which are apparently strid- ulatory in function. Amblycerus atypicus is clearly separated from all the other Amblycerus species and especially from those in the cistelinus group by the form, number and arrangement of sclerites in the male internal sac (Fig. 7). Types.—Holotype ¢°. FRENCH GUI- ANA: Maroni River; Collection Wm Schaus, deposited in the National Museum Natural of History, Smithsonian Institution, Washington, D.C. U.S.A. One paratype ¢ with same label deposited in the Cole¢cao de Entomologia Pe. Jesus S. Moure, Curi- tiba, Brazil. Etymology.—tThe species name refers to the atypical elytro-femoral method of pos- sible stridulation in Amblycerus. 341 Amblycerus ischiodontus Ribeiro-Costa, new species (Figs. 11—18) Description.—Measurements (pronotum + elytra): Length 4.48 mm; width 2.72 mm. Integument: General coloration, dark red. Eyes, legs and undersurface of thorax darker. Vestiture: Head, appendages and ventral area of thorax and abdomen, flavous; met- episternum in part, golden yellow. Prono- tum (Fig. 11) golden yellow rarely with small brown patches. Scutellum flavous. El- ytra (Fig. 11) golden yellow; brown hairs in rare irregular patches and flavous hairs in lines along interstitial margins and strial sulci. Pygidium (Fig. 11) golden yellow with two small central, brown patches. Frons slightly flat, evenly punctuated, frontal carina absent. Eyes moderately fac- eted (7 ommatidia), protruding laterally; oc- ular index 3.8; ocular sinus 1/5.0 diameter of eye in lateral view; postocular lobe 1/6.7 times largest length of eye in lateral view. Antenna (Fig. 16) serrate from fourth to tenth segments, all perceptibly longer than wide (1.4 times), last segment subelliptical. Pronotum trapezoidal with lateral margins moderately arched in dorsal view, cervical sulcus in lateral thirds and a sulcus outlin- ing basal lobe; long lateral carina, nearly reaching apex of pronotum, not forked an- teriorly; surface densely punctulate, lateral one-third either side also coarsely punctate. Prosternal process wide, not sulcate, apex subacute, moderately expanded between an- terior coxae and fitting into sulcate meso- sternum. Scutellum (Fig. 12) about 1.8 times as long as wide, apex strongly triden- tate on rounded base. Elytra slightly convex in cross section along elytral suture, sub- truncated apically and with striae strongly impressed. Transverse axis of metepister- num (Fig. 13) moderately divergent from metapleural suture; transversely striate and strongly curved area running with longitu- dinal axis, apparently being a modification PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 342 Figs. 9-10. Amblycerus atypicus. 9, Scanning electron micrograph (SEM) of the external margin of elytron showing ‘file.’ 10, SEM of the inner distal portion of the hind femur showing “‘scraper.”’ VOLUME 101, NUMBER 2 Rs \ qd HANS yl) ) \\ \ i rt inhi Wits A NY KENT Ii' iA - Np Way ») ; \ \ Kt AU Ne EE Du Wii AA TEA ea) RAE REAR CA 4 1 i Ml i ‘i p NA Hal Pn (Fal i \ iY 1 ay TA gah { HURRAY SR iH i ‘\ RR AM at Tt eg ne Dark red integument Golden yellow hairs 343 14 —— fae) Dark red integument LAU ta! Brown hairs [el Dark red integument Flavous hairs imm Figs. 11-16. Amblycerus ischiodontus. 11, Dorsal habitus. 12, Scutellum. 13, Metepisternum. 14, Hind leg. 15, Ventral margin of hind femur showing the tooth. 16, Antenna. of it (Fig. 13). Hind coxa with coarse punc- tures slightly denser in distal half. Hind fe- mur (Fig. 14) 2.3 times as long as wide and with tooth pronounced in distal portion of ventral margin (Fig. 15); internal face on distal region without transverse striations. Hind tibia (Fig. 14) with coronal denticles approximately of same size; lateral spur 2.4 times length of median, first hind tarsal seg- ment 1.2 times as long as lateral spur and 344 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 17-18. 2.8 times median spur; ventral face mod- erately convex with inconspicuous lines of punctures and setae in margins. Male py- gidium vertical, round apically. Fifth visible abdominal sternum not emarginate in male. Female unknown. Male terminalia (Figs. 17, 18): Eighth tergite rounded apically. Median lobe (Fig. 17) with length 3.8 times its largest width Amblycerus ischiodontus. 17, Male terminalia, median lobe. 18, Same, tegmen and lateral lobes. in basal area; ventral valve moderately long, acute apically, with concave lateral margins; dorsal valve subtriangular with straight lateral margins and rounded apex. Basal area (Fig. 17) of internal sac without anterior and median sclerites; a pair of pos- terior thin sclerites, prolonged and with a tooth sub-basally on one side. Median area of internal sac (Fig. 17) with a pair of lam- VOLUME 101, NUMBER 2 inar sclerites, sinuous, serrate sub-basally, teeth approximate; unpaired sclerite in fork form 0.6 times as long as laminars, with moderately separate stems. In apical area of internal sac (Fig. 17) one sclerite with very long stems. Median and apical areas of in- ternal sac with denticles near apical sclerite. Tegmen (Fig. 18) with moderate emargi- nation in “V” between enlarged lateral lobes. Discussion.—Amblycerus ischiodontus shares with A. eustrophoides, A. pollens, and A. stridulator an area of the metepister- num with transverse striations (Fig. 13) and an apical tooth on the ventral margin of the metafemur (Fig. 15). The study of all char- acters together did not show great affinity among the mentioned species. Differences are easily recognized in the patterns of pu- bescence, form of the striate area, and the sclerites in the male internal sac. Kingsolver et al. (1993) commented that A. pollens is more similar to A. eustrophoi- des when compared with A. stridulator. Romero et al. (1996) included A. eustro- phoides in the alternatus group along with A. serieguttatus, A. alternatus, and A. schwarzi and stablished the stridulator group only for A. stridulator. Revisionary studies on Brazilian species under way indicate that A. pollens and A. ischiodontus are members of two distinct species groups, respectively. According to Arrow (1904) the body area where the stridulatory mechanism is placed in Coleoptera can be the same in non-relat- ed groups, as it can be different in different members of the same group. According to him, there are Coleoptera with the two types of stridulatory mechanism placed in different parts of the body. Amblycerus ischiodontus is easily sepa- rated from all other Amblycerus by its char- acteristic form and disposition of sclerites in the male internal sac (Fig. 17). Type.—Holotype 6. BRAZIL: Amazo- nas, Parana do Xiboreninho, 03°15’S— 06°00'W; 5 Aug. 79; mixed water; Canopy Fogging Project TRS#05, Tray 326 Adis, 345 Erwin, Montgomery et al. collectors; de- posited in the National Museum Natural of History, Smithsonian Institution, Washing- ton, D:C..U:S.A. Etymology.—The name of the species refers to the presence of a tooth on the ven- tral margin of the hind femur. KEY TO AMBLYCERUS SPECIES WITH AREAS OF INTEGUMENT PROBABLY INVOLVED IN STRIDULATION 1. External margins of elytra partly with an area with overlapping scales (Figs. 3, 9); inner distal portion of hind femur with fine transverse stri- ation (Figs. 6, 10); usually scattered black spots on elytra (Fig. 1); pygidium with large, ovate, velvety dark brown spot (Fig. 4) ......... 2 — Elytra and hind femur without integumental modifications; patterns of coloration on elytra and pygidium otherwise . Elytra with white hairs vaguely mottled; very coarse punctures on lateral third of pronotum; Nm male terminalia with four pairs of sclerites and onesunpainedysclenite: arcana oes or toe ss Amblycerus guazumicola (Kingsolver and Johnson) — Elytra usually with well marked white irregular spots on median and external margins and small irregular patches on remainder of elytra (Fig. 1); moderately coarse punctures on lateral third of pronotum; number of sclerites in in- ternal sac of male terminalia otherwise .... . 3 3. Male internal sac with three pairs of sclerites and one unpaired sclerite (Fig. 7) ....... RE Ce sat Se OIE Amblycerus atypicus, new species — Male internal sac with two pairs of sclerites andyvone unpalred\isclenitesea ah. = or lich ver 4 4. Pair of laminar sclerites in male internal sac with tooth — Pair of laminar sclerites in male internal sac smooth 5. Male internal sac with basal pair of sclerite spine-shaped — Male internal sac with basal pair of sclerite Y- shapedis, see. Amblycerus cistelinus (Gyllenhal) 6. Male internal sac with unpaired median sclerite serrate basally along one side ... Amblycerus sosia Ribeiro-Costa and Kingsolver — Male internal sac with unpaired median sclerite with serration only on base EL aeaseco ets care Amblycerus jatayensis (Pic) 7. Metepisternum with striate area transverse to BRAS Amblycerus whiteheadi Kingsolver metepisternal sulcus, apparently being a mod- ification of metepisternum; apical portion of ventral margin of metafemur finely striate, with conspicuous tooth; pygidium with median line 346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . Amblycerus stridulator Kingsolver, Romero N., and Johnson — Metepisternum with striate area running with longitudinal axis of metepisternal sulcus, ap- parently being a modification of it; apical por- tion of ventral margin of metafemur with con- spicuous or inconspicuous smooth tooth; py- gidium with or without median line 8. Apical portion of ventral margin of metafemur with one inconspicuous tooth; posterior end of metepisternum straight on striate area .... EA eg st be a GRE e Amblycerus pollens (Sharp) — Apical portion of ventral margin of metafemur with one conspicuous tooth; posterior end of metepisternum curved on striate area 9. Posterior end of metepisternum gently curved on striate area; scutellum tridentate; pygidium with narrow median line BRR ei Amblycerus eustrophoides (Schaeffer) — Posterior end of metepisternum strongly curved on striate area (Fig. 13); scutellum tri- dentate on round base (Fig. 12); pygidium oth- CEWISe™ Amblycerus ischiodontus, new species ACKNOWLEDGMENTS I thank Dra. Dilma Solange Napp and Dra. Catherine N. Duckett for reviewing the manuscript. This is Contribution No. 1049 from Departamento de Zoologia, Universi- dade Federal do Parana. LITERATURE CITED Arrow, G. H. 1904. Sound production in the lamelli- corn beetles. Transactions of the Royal Entomo- logical Society of London 4: 709-750. Dumortier, B. 1963. Morphology of sound emission apparatus in Arthropoda, pp. 277-345. /n Busnel R. G., Acoustic Behavior of Animals. Elsevier Publ. Co., New York. 933 pp. Kingsolver, J. M. 1970. A synopsis of the subfamily Amblycerinae Bridwell in the West Indies, with descriptions of new species (Coleoptera: Bruchi- dae). Transactions of the American Entomological Society 96: 469-497. Kingsolver, J. M., N. J. Romero, and C. D. Johnson. 1993. Files and scrapers: circumstantial evidence for stridulation in three species of Amblycerus, one new (Coleoptera: Bruchidae). Pan-Pacific En- tomologist 69(2): 122-132. Ribeiro-Costa, C. S. 1987. Descrigdes de oito novas espécies de Amblycerus Thunberg (Coleoptera: Bruchidae). Revista Brasileira de Zoologia 14(3): 629-648. Romero, J., C. D. Johnson, and J. M. Kingsolver. 1996. Revision of the genus Amblycerus of the United States and Mexico (Coleoptera: Bruchidae: Am- blycerinae). United States Department of Agricul- ture Technical Bulletin No. 1845, 166 pp. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 347-358 A REVIEW OF THE BEACH-FLY GENUS ISOCANACE MATHIS (DIPTERA: CANACIDAE) WAYNE N. MATHIS Department of Entomology, Smithsonian Institution, Washington, DC 20560-0169, U.S.A. (e-mail: mathisw @nmnh.:si.edu) Abstract.—The species of the beach-fly genus [socanace Mathis are reviewed, including the description of two new species: I. crosbyi (New Zealand. South Island. NN: Cable Bay (41°09.6'S, 173°24.9’E) and I. freidbergi (Kenya. Takaungu (50 km N of Mombasa)). Descriptions for the genus and two species groups are also revised. Key Words: Old World Recent field work on New Zealand has led to the discovery of the first species of the beach-fly family Canacidae from that country and has also prompted the research that resulted in this review of the genus J/s- ocanace Mathis. Macrocanace Tonnoir and Malloch, with two endemic species from New Zealand, had previously been included in the Canacidae (Harrison 1959), but is now assigned to the family Tethinidae (Mathis and Munari 1996). The New Zealand species of Canacidae is undescribed and is in the genus /socan- ace where it is the second known species of the Isocanace albiceps group. The first species of that group, J. albiceps (Malloch), is from Australia (Mathis 1996). The new species is apparently widespread on New Zealand, occurring on both North and South Islands and undoubtedly on some of the associated offshore islands. I am also taking this opportunity to de- scribe an Afrotropical species of /socanace that Amnon Freidberg and Fini Kaplan col- lected some years ago in Kenya. The latter species belongs to the J/socanace_ briani group, which previously comprised three Afrotropical species: /. australis Mathis review, Diptera, Canacidae, beach flies, Jsocanace, I. crosbyi, I. freidbergi, (Kenya, South Africa), I. briani Mathis (Al- dabra, Madagascar), and /. flava (Canzoneri and Meneghini; Zaire). The last species, /. flava, is unusual among beach flies, being one of just a few canacid species that occurs in freshwater habitats. As implied by the common name for the family, most species occur in saline habitats, especially along maritime coasts. The new species from Kenya is very similar to and is apparently the sister species of J. flava, although it is found in saline environments along the western coast of the Indian Ocean. The addition of two new species to /so- canace, one to each of the species groups, alters the generic and species-group char- acterizations, which are revised. A revised key to the species groups and species is also provided. Isocanace is a relatively recent genus in the nomenclatural history of the Canacidae (Mathis 1982), and the genus has been treated in only two subsequent papers (Mathis 1992, 1996). The first paper is a world catalog, and the second is a review of the Australian beach flies. Jsocanace is known only from the Afrotropical and Aus- tralasian Regions of the Old World (Mathis 348 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1992). This apparently disjunct distribution is probably an artifact due to the poor sam- pling of Canacidae from countries between Australia and Africa rather than the actual distribution of the genus. Nothing is known about the immature stages or most other as- pects about the biology of the included spe- cies except for brief comments on habitats where specimens have been collected. The descriptive format for the new spe- cies follows Mathis and Wirth (1979) and Mathis (1982, 1988). More details concern- ing the morphology and higher classifica- tion of the Canacidae are found in Mathis (1982, 1992) and Wirth (1987). I follow Crosby et al.’s (1976) geographic codes for New Zealand zoogeographic provinces: AK = Auckland, NC = North Canterbury, NN = Nelson. Two venational ratios are used in the de- scriptions. Costal vein ratio: The straight line distance between the apices of vein R,,;, and R,,. _— iF ( XS Ta Si oe , Ie - i IS pea Figs. 24-25. External male terminalia of /socan- ace briani. 24, Epandrium, cercus, and surstylus, lat- eral view (type locality). 25, Same (Aldabra), lateral view. 24 B. Cogan, A. Hutson (1 6, 1 2; USNM). Picard: Bassin Labine (trail to), 20 Mar 1986, W. N. Mathis (12 36, 7 2; USNM); La Gigi, 19-24 Mar 1986, W. N. Mathis (7 6, 20 2: USNM); Settlement, 15—21 Mar 1986, W. N. Mathis (3 6, 1 2; USNM). MADAGASCAR. Antsiranana: Nosy Be, Ambatoloaka (beach), 4—7 Apr 1991, A. Freidberg, EF Kaplan (15 ¢, 25 2; USNM). Nosy Be, Andoany (Hell-Ville), 5 Apr 1991, A. Freidberg, E Kaplan (4 6, 2 9; USNM). Nosy Be, Sambirano, Lokobe (6 m; rocks on beach), 9-23 Nov 1957, B. Stuckenberg (4 ¢, 13 2; USNM). Ramena, 10 Apr 1991, A. Freidberg, E Kaplan (2 6, 18 2; USNM). Nosy Domba, 6 Apr 1991, A Freidberg, EF Kaplan (1 2; USNM). Distribution.—Afrotropical: Madagascar (Antsiranana), Seychelles (Aldabra). Isocanace flava (Canzoneri and Meneghini) (Figs. 26—29) Canace flava Canzoneri and Meneghini 1969: 184 [HT 6 (MRAC); Zaire. Albert National Park: May ya Moto].—Cogan 1980: 694 [Afrotropical catalog]. Isocanace flava: Mathis 1982: 17 [generic combination; figures of head, thorax, ¢ terminalia]; 1992: 6 [world catalog]. Diagnosis.—This species is distinguished from congeners by the following combina- tion of characters: mesofrons with scattered setulae on middle (Fig. 26); lateroclinate fronto-orbital setae 4 (Figs. 26—27); arista with branching rays shorter, at most slightly longer than aristal width at base (Fig. 27); 3 dorsally curved genal setae (Fig. 27); an- terior notopleural seta distinctly smaller than posterior seta or lacking; anepisternal and katepisternal seta present, black; male terminalia as in Fig. 29. Distribution.—Afrotropical: Zaire (Haut- Zaire). Isocanace freidbergi Mathis, new species (Figs. 30-31) Description.—This species is similar to /. flava but is distinguished from it and other congeners by the following combination of characters: small to moderately-small beach flies, length 1.65—2.40 mm; mostly gray. Head: Mesofrons whitish gray to tannish gray, bearing scattered setulae on middle and 2-3 large, lateral proclinate setae; la- teroclinate fronto-orbital setae 3. Antenna mostly yellow but with varying amounts of blackish overlay, especially on flagellomere 1; arista bearing short branching hairs, these at most slightly longer than aristal width at base. Large, dorsally curved genal setae 3, 1 large anteroclinate seta, and 3-5 much smaller, anteroclinate setulae. Thorax: Acrostichal setulae in 4 some- what irregular rows of numerous setulae and with a larger, prescutellar pair; anterior no- topleural seta either lacking or pale and much smaller than posterior seta; anepisternal and 356 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 26-29. Epandrium and surstylus, lateral view. katepisternal seta present, black. Wing with costal vein ratio 0.21; M vein ratio 0.36—0.47. Femora of mid and hindlegs mostly micro- tomentose, gray, concolorous with pleural re- gion, midfemora more yellowish but with some faint investment of gray microtomen- tum; tibiae and tarsi yellow. Isocanace flava. 26, Head, anterior view. 27. Same, lateral view. 28, Thorax, dorsal view. 29, Abdomen: Male terminalia (Figs. 30— 31): Surstylus in lateral view irregularly hooklike to sickle shaped (Fig. 30), widest at basal 4%, apex pointed, curved anteriorly; in posterior view as in Fig. 31. Type Material.—The holotype ¢d is la- beled ““KENYA Takaungu, 50 km North VOLUME 101, NUMBER 2 Figs. 30-31. view. 31, Same, posterior view. Mombasa 3. XII. 1989 A. FREIDBERG & FINI KAPLAN.” The holotype is double mounted (minuten in a block of plastic), is in excellent condition, and is deposited in the USNM. Thirty-nine paratypes (14 4, 25 2; TAU, USNM) bear the same label data as the holotype. Other paratypes are as fol- lows: KENYA. Gazi (60 km S Mombasa; route Al4), 5 May 1991, A. Freidberg, EF Kaplan (3 6, 3 2; TAU, USNM). Distribution.—Afrotropical: Kenya. Etymology.—The species epithet is a S5i, External male terminalia of /socanace freidbergi. 30, Epandrium, cercus, and surstylus, lateral genitive patronym to recognize the impor- tance of the collecting efforts of Dr. Amnon Freidberg (TAU) to my research on the shore and beach flies of Africa. Remarks.—This species is apparently the sister species of /. flava, with supporting ev- idence being the hook to sicklelike apex of the surstylus in lateral view (Figs. 29, 31). Unlike its sister species, which is found in freshwater habitats in the Rift Valley (east- ern Zaire), this species occurs in coastal habitats that are saline. ACKNOWLEDGMENTS I am grateful for the assistance and co- operation of many organizations and indi- viduals who contributed to the field work and production of this paper. To Dr. Trevor K. Crosby (NZAC) and Drs. Daniel J. Bick- el and David K. McAlpine (AM) and their institutions, who expedited the arrange- ments for field work and loaned specimens, I express my sincere thanks. The line illustrations were either entirely produced (Figs. 26—28) or carefully inked by Mr. Young T. Sohn. For reviewing a draft of this paper I thank Drs. Amnon Freidberg, Volker Hollmann-Schirrmacher, Allen L. Norrbom, and Norman E. Wood- ley. I am also grateful to Dr. Anna K. Ber- nensmeyer, former Associate Director for Science, National Museum of Natural His- tory, Smithsonian Institution, for financial support to conduct research at the Natural History Museum (BMNH), London, Eng- land, through grants from the Research Op- portunity Fund. Field work on New Zea- land was facilitated through a grant from Mr. Helmut Hollmann, benefactor for re- search on shore flies. I gratefully acknowl- edge his generous assistance. Travel to and from New Zealand was largely provided by British Airways, and I am grateful to them for generously supporting this research, es- pecially the field work on New Zealand. Field work on New Zealand was greatly ex- pedited through the pleasant assistance of Dr. Volker Hollmann-Schirrmacher. LITERATURE CITED Canzoneri, S. and D. Meneghini. 1969. Sugh Ephy- dridae e Canaceidae della fauna etiopica. Bollet- tino del Museo Civico di Storia Naturale di Ve- nezia 19: 101-185. Cogan, B. H. 1980. 79, Family Canacidae, p. 694. In Crosskey, R. W., ed., Catalog of the Diptera of the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Afrotropical Region. London: British Museum (Natural History), 1437 pp. Colless, D. H. and D. K. McAlpine. 1991. 39. Diptera (Flies), pp. 717-786. Jn Naumann, I. D., ed., In- sects of Australia. A Textbook for Students and Research Workers. 2nd ed., Vol. 2. Melbourne University Press, Carlton, Victoria, 1137 pp. Crosby, T. K., J. S. Dugdale, and J. C. Watt. 1976. Recording specimens localities in New Zealand: an arbitrary system of areas and codes defined. New Zealand Journal of Zoology 3: 69. Harrison, R. A. 1959. Acalypterate Diptera of New Zealand. New Zealand Department of Scientific and Industrial Research Bulletin 128, 382 pp. Malloch, J. R. 1925. Notes on Australian Diptera, No. vi. Proceedings of the Linnean Society of New South Wales 50(2): 80-97. Mathis, W. N. 1982. Studies of Canacidae (Diptera), I: Suprageneric revision of the family, with revisions of new tribe Dynomiellini and new genus /socan- ace. Smithsonian Contributions to Zoology 347: iii + 1-29. . 1988. First record of the genus Procanace Hendel from North America, with the description of a new species (Diptera: Canacidae). Proceed- ings of the Entomological Society of Washington 90(3): 329-333. . 1989. 103. Family Canacidae, pp. 669-670. In Evenhuis, N. L., ed., Catalog of the Diptera of the Australasian and Oceanian Regions. E. J. Brill and B. P. Bishop Museum special publication 86, 1155 pp. . 1992. World catalog of the beach-fly family Canacidae (Diptera). Smithsonian Contributions to Zoology 536: 1-18. . 1996. Australian beach flies (Diptera: Cana- cidae). Proceedings of the Biological Society of Washington 109(2): 326-348. Mathis, W. N. and L. Munari. 1996. A catalog of the Tethinidae (Diptera). Smithsonian Contributions to Zoology 584: 1—27. Mathis, W. N. and W. W. Wirth. 1979. Beach flies of Madagascar (Diptera: Canacidae). Annals of the Natal Museum 23(3): 785-796. Wirth, W. W. 1951. A revision of the dipterous family Canaceidae. Occasional Papers of Bernice P. Bish- op Museum 20(14): 245-275. . 1956. New species and records of South Af- rican Canaceidae (Diptera). Journal of the Ento- mological Society of South Africa 19(1): 47-51. 1987. 102. Canacidae, pp. 1079-1083. In McAlpine, J. F, ed., Manual of Nearctic Diptera, Vol. 2. Research Branch, Agriculture Canada, Monograph 28. Ottawa, iv + 675-1332 pp. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 359-371 TYPES AND BIOLOGICAL NOTES OF THE EASTERN NORTH AMERICAN SAWFLIES OF PONTANIA COSTA AND PHYLLOCOLPA BENSON (HYMENOPTERA: TENTHREDINIDAE) DESCRIBED BY MARLATT, DYAR, AND ROHWER ALEXEY G. ZINOVJEV AND DAVID R. SMITH (AGZ) Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034, Russia (e-mail: zag@zisp.spb.su); (DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, MRC-168, Washington, DC 20560-0168, U.S.A. (e-mail: dsmith @sel.barc.usda.gov) Abstract.—The taxonomic placement of 18 sawfly species assigned to Pontania and Phyllocolpa or described under Pontania from eastern North America by Marlatt, Dyar, and Rohwer are discussed. The authorship of Pteronus carpini, Pontania consors, and P. borealis attributed to Marlatt is changed to Dyar. The following eight species from eastern North America belong to Pontania (Eupontania): P. (E.) s-desmodioides (Walsh) (= Pon- tania borealis Dyar 1898, n. syn.); P. (E.) s-pisum (Walsh); P. (E.) s-pomum (Walsh); P(E.) gracilis Marlatt; P. (E.) rugulosa Marlatt; P. (E.) petiolaridis Rohwer, P. (E.) consors Dyar; and P. (E.) lucidae Rohwer. Three species are included in the leaf-rolling crassispina group of the subgenus Pontania: P. (P.) pumila Rohwer; P. (P.) populi Mar- latt; and P. (P.) terminalis Marlatt. Five species are included in the genus Phyllocolpa: P. nigrita (Marlatt); P. pectoralis (Marlatt); P. robusta (Marlatt); P. leavitti (Rohwer); and P. crassicornis (Rohwer), n. comb. Pontania acuminata Marlatt is transferred to Nematus, n. comb. Lectotypes are designated for seven species. The Salix and Populus host plants are given and associated galls are illustrated. Key Words: Pontania, Phyllocolpa, sawflies, leaf galls, leaf folds, Marlatt, Dyar, Rohwer The Nearctic gall-making and leaf-fold- ing sawflies have received little attention and have not been revised due to few ap- parent morphological characters in the adults and lack of information on associated galls, habits, and other biological data. Smith (1979) listed the gall-forming species of Nematinae in the genera Euura, Pontan- ia, and Phyllocolpa, following the classifi- cation in place at that time. Subsequent work on the Palearctic fauna by Vikberg (1982) and Zinovjev (1993), has refined the classification, most notably by the recog- nition of various subgenera and species groups within each genus. Consequently, the taxa described from the Nearctic Region need to be reevaluated, and associated galls and host information recorded where known. In this paper, we discuss the species of Pontania and Phyllocolpa described by Marlatt, Rohwer, and Dyar from eastern North America. The authorship of three species are correctly attributed to Dyar rath- er than Marlatt. The species described by these authors are significant because many have associated galls, host, and biological information. These authors are also respon- sible for all of the species of Phyllocolpa and half of the species of Pontania known 360 from eastern North America. We use the di- agnostic characters and classification used by Zinovjev (1993), except that Phyllocol- pa is treated as a genus rather than a sub- genus of Pontania. Following is a summary of the classification and biological infor- mation for the subtribe Euurina: Subtribe Euurina—Produces galls or leaf folds on Salicaceae, mainly Salix spp., but a few on Chosenia in eastern Asia and on Populus in North America. Genus Euura Newman—Produces bud, stem, petiole, or midrib galls. Genus Phyllocolpa Benson—Larvae in rolled leaves or leaf margins, without swelling at site of egg lay- ing. At least two species groups in North America, /eucapsis and leu- costicta. Genus Pontania Costa Subgenus Pontania Costa—Larvae in closed galls or leaf rolls, with site of egg laying marked by dis- tinct swelling on upper surface of leaf. Three species groups: cras- sispina, dolichura, proxima. Subgenus Eupontania Zinovjev— Larvae produce closed leaf galls attached to the midvein or occa- sionally a larger lateral vein. At least two of the five Palearctic species groups occur in North America, polaris and viminalis. Five species treated here belong in Phyl- locolpa, two in the leucosticta group, two in the Jeucapsis group, and one is not placed. Phyllocolpa crassicornis (Rohwer) was not included in the review by Smith and Fritz (1996). Pontania acuminata Mar- latt, P. populi Marlatt, and P. terminalis Marlatt, included in Phyllocolpa by Smith and Fritz (1996), are here transferred to Pontania or Nematus. The subgenus Pontania is represented by three species groups in North America. The gall-making Pontania proxima (Lepeletier), which was described under the name Messa PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON hyalina by Norton (1864), was introduced from Europe together with the host plants Salix alba L., Salix fragilis L., and their hy- brids, and it is the only representative of the proxima group known in temperate North America. Galls of an undescribed species of the dolichura group on Salix sericea (Marsh.) were found in Otsego Co., N.Y., in July 1996 by A. G. Zinovjev and R. Fritz. Similar galls have been encountered on S. nigra Marsh. and other willows in the Harvard University Herbaria by AGZ. We recognize three species of the leaf-rolling crassispina group, P. (P.) pumila Rohwer, P. (P.) populi Marlatt, and P. (P.) termin- alis Marlatt. The following eight species from eastern North America belong to Pontania (Eupon- tania): P. (E.) s-pisum (Walsh), P. (E.) s- pomum (Walsh), P. (E.) s-desmodioides (Walsh), P.(E.) gracilis (Marlatt), P. (E.) rugulosa Marlatt, P. (E.) petiolaridis Roh- wer, P. (E.) consors Dyar, and P. (E.) lu- cidae Rohwer. Their galls (Figs. 1—9) help distinguish the species. The Walsh species are newly assigned to this group. For some of the species discussed, the remnants of the galls from which type spec- imens were reared are still preserved, and we illustrate these. These remnants made it possible to check the host plant identifica- tions by G. Argus. Acronyms used for mu- seums are: USNM = National Museum of Natural History, Smithsonian Institution, Washington, D.C.; CUIC = Cornell Uni- versity, Ithaca, New York. SPECIES DESCRIBED BY MARLATT Pontania acuminata Marlatt 1896a: 32. Type locality—Michigan. The holotype is labeled ‘“‘Ag.Coll. Mich.,’’ presumably from East Lansing. Type material.—Described from one fe- male. The holotype was redescribed by Smith and Fritz (1996). Deposited in the USNM. Host plant.—Unknown. Notes.—Examination of the ovipositor of VOLUME 101, NUMBER 2 61 of + Fe eee Figs. 1-2. Galls of Pontania (Eupontania) spp., Milford, Otsego Co., N.Y. 1, P. (E.) s-pisum on Salix discolor. 2, P. (E.) s-pomum on Salix eriocephala. 362 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. the holotype revealed that this species be- longs to the genus Nematus Panzer. The Ovipositors of Nematus are normally broad and straight (Benson 1958, figs. 712-725), and those of Pontania and Phyllocolpa are usually slender and curved (Benson 1958, figs. 636-641). The slight ventroapicale margination of the sheath (Fig. 10) led to the placement of this species in Phyllocolpa by Smith and: Fritz (1996). The correct combination is Nematus acuminatus (Mar- latt), n. comb. Pontania atra Marlatt 1896a: 37. Type locality—Michigan. The holotype is from “Ag. Coll. Mich.,”” and is presum- ably from East Lansing. Type material.—Described from one fe- male. The holotype female is labeled: “‘Ag. Coll. Mich. 4—21 90 62; Davis; Type fe- male; Type No 1916 U.S.N.M.; Pontania atra n. sp. female.’’ Deposited in the USNM. Gall of Pontania (Eupontania) gracilis on Salix sericea, Milford, Otsego Co., N.Y. Host plant.—Unknown. Notes.—This species is currently placed as a synonym of Amauronematus amento- rum (Forster) (Smith 1979). The latter, to- gether with the related species inhabiting willow catkins, are now placed in a separate genus, Pontoprista Malaise. Pteronus dubius Marlatt 1896a: 74. Type locality— Wellesley, Mass. Type material.—Described from one male. The holotype is labeled: ‘‘Wellesley, Mass., March 29, 1890; Type male; Type No. 1937 U.S.N.M.; Pteronus dubius male.’ The genitalia are on a separate pin in a tube and with the label: ““TYPE Pter- onus dubius Marl.’’ Deposited in the USNM. Host plant.—Unknown. Notes.—This species was correctly as- signed to Pontania (Smith 1979). Males are difficult to place and generic placement is the best that can be done at present. VOLUME 101, NUMBER 2 Pontania gracilis Marlatt, 1896a: 39. Type locality.—Virginia. Type material.—Described from two fe- males and an undetermined number of galls. Lectotype female, here designated: PE Nowi52x Iss. Apri 19:86) Type: female; Type No. 1919 U.S.N.M.; Pontania gracilis n. sp.”’ Paralectotype: ““No 152x, Iss. Apr. 19. SGewelype: aemale. TyperuNo.c 1919 U.S.N.M.”’ There are two dry galls with the following labels: “152x Va Sept. 29. 85”’’; one of them (Fig. 6) with the remnants of the leaf (the lower surface with sparse tri- chomes) and the label ‘“‘gall on Salix sericea det G. Argus, 1996.” The types and galls are in the USNM. Host plant.— Salix sericea Marsh. (iden- tified by G. Argus). Notes.—This is a valid species of Pon- tania (Eupontania). The description of P. gracilis larvae and biology by Dyar (1897b) refers to P. petiolaridis Rohwer (see below). A typical gall of this species is shown in Fig. 3 (from Otsego Co., N.Y., collected by AGZ and R. Fritz). Pontania nigrita Marlatt 1896a: 27. Type locality—Michigan. The holotype is labeled “‘Ag.Coll.Mich,’’ presumably from East Lansing. Type material.—Described from one fe- male. The holotype was redescribed by Smith and Fritz (1996). Deposited in the USNM. Host plants——According to Smith and Fritz (1996), the host plants are Salix seri- cea, §. discolor Muhl., and S. eriocephala Michx. Notes.—This is a valid species of Phyl- locolpa and belongs to the leucosticta group. Its larva and biology were described by Dyar (1897b) under the name Pontania pallicornis Norton. Pontania pectoralis Marlatt 1896a: 31. Type locality —Algonquin, Illinois. Type material.—Described from one fe- male. The holotype was redescribed by Smith and Fritz (1996). Deposited in CUIC. 363 Host plant—Unknown. Notes.—This is a valid species, Phyllo- colpa pectoralis (Marlatt). Pontania populi Marlatt 1896b: 253. Type locality—New York, as given by Marlatt. The specimen described was reared by Dyar; Dyar (1897a) stated it was from Fore Lee, NJ: Type material.—Described from one fe- male. The holotype was redescribed by Smith and Fritz (1996). Deposited in the USNM. Host plant.— Populus grandidentata Michx. Notes.—A valid species, Pontania (Pon- tania) populi Marlatt, belonging to the cras- sispina group. The larvae and biology were described by Dyar (1897a). It produces leaf rolls on the host creating a small swelling (procecidium) at the site of egg laying. There is more than one generation per year. The larvae eat the parenchyma and leave the upper cuticle intact, which is typical for species of the crassispina group. A very slight ventroapical indentation of the sheath (Fig. 11) and the leaf-folding habit led to the inclusion of this species in Phyllocolpa by Smith and Fritz (1996). Pontania robusta Marlatt 1896a: 32. Type locality—‘‘Michigan and District of Columbia (7?).’” The lectotype is labeled ‘*Ag.Coll.Mich.,’> presumably East Lan- sing. Type material—Described from one fe- male and one male. The lectotype, a female from Michigan, was selected and rede- scribed by Smith and Fritz (1996). Depos- ited in the USNM. Host.—Populus tremuloides Michx. Notes.—This is a valid species of Phyl- locolpa in the leucapsis group. The larva and biology were described by Dyar (1897b). It produces leaf folds on aspen, as was confirmed by Smith and Fritz (1996). There is one generation a year. Pontania rugulosa Marlatt 1896a: 41. 364 Type locality—Michigan. Type material.—Described from two males, “‘one reared (?) from willow gall.” The lectotype, here designated, is labeled: “O; 17: Gollection €:Vr Riley; Type male; Type No 1920 U.S.N.M.; Pontania rugulosa M. n.sp.” There is a gall on the same pin, but it does not belong to the lectotype. De- posited in the USNM. Host plant.—Willow (species unknown). Notes.—This species belongs to Pontan- ia (Eupontania) according to the shape of the mandibles, but we are not certain if it is a valid species because males are difficult to place. The gall lacks an exit hole, but the male has part of the pupal skin attached to the leg. It may have been reared from a similar gall, but not the gall on the pin. The gall on the pin is equally produced from both sides of the leaf, but it is practically without leaf remnants and the willow prob- ably cannot be identified. Pontania terminalis Marlatt 1896b: 253. Type locality—Near New York City. The specimens were from Dyar’s collec- tions and, according to Dyar (1897a), were from Van Cortland Park, New York City. Type material.—Described from three fe- males and two males. The female lectotype was selected and the species redescribed by Smith and Fritz (1996). Deposited in the USNM. Host plant.— Salix sericea, according to Smith and Fritz (1996). The host plant re- lationships of P. terminalis need to be con- firmed. There is possibly more than one- species under this name. Notes.—The valid name is Pontania (Pontania) terminalis Marlatt, and it be- longs to the crassispina group. The larvae and biology were described by Dyar (1897a). It produces leaf rolls on willow similar to P. populi except with a smaller swelling at the site of egg laying. The lar- vae eat parenchyma and leave the upper cu- ticle intact, which is typical for this species group. The slight ventroapical indentation of the sheath (Fig. 12) and leaf-folding hab- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON it led to the placement of this species in Phyllocolpa by Smith and Fritz (1996). SPECIES DESCRIBED BY DYAR Dyar (1898) attributed the authorship of Pontania consors, Pontania borealis, and Pteronus carpini to Marlatt (1898), but Dyar’s descriptions of these species precede those of Marlatt by about half a year. Mar- latt (1898) cited “‘Dyar. N. Y. Ent. Soc., VI., June 1898, p. 121” in his descriptions. Therefore we change the authorship of the following names to Dyar: Pontania boreal- is, Pontania consors, and Pteronus carpini. We treat the adults reared from larvae de- scribed by Dyar as holotypes or lectotypes. Pteronus carpini is a valid species in the genus Nematus (Smith 1979). Pontania borealis Dyar 1898: 121 (galls, larva).—Maarlatt 1898: 302 (female). Type locality.—Plattsburgh, N.Y., ac- cording to Dyar (1898). Type material—Dyar did not state the number of galls and larvae he reared. Mar- latt described the reared adults from two fe- males. The lectotype female, here designat- ed, is labeled: ““8S; Collection H. G. Dyar; Type No 4131 U.S.N.M.; Pontania borealis female Marl.’ Paralectotype:‘*8S; Collec- tion H. G. Dyar; Pontania borealis Marl.” There is also one similarly labeled speci- men without Marlatt’s identification label. Galls: There are two galls on a pin (Fig. 4) labeled: “‘8S; Collection H. G. Dyar; Type No 4131 U.S.N.M.; Pontania borealis gall Marl.”’ An additional label is added: “‘upper gall: P. (Eu.) s-pomum, lower gall: P. (Eu.) s-desmodioides, A. Zinovjev det, 1997”. There are three galls on another pin (Fig. 5) without Marlatt’s identification label, but some of them probably also belong to the type series of P. borealis. The two upper galls are of P. (E.) s-desmodiodes (A. Zi- novjev det. 1997, galls on Salix humilis det G. Argus, 1996; lower gall of P. (E.) s-po- mum). Deposited in USNM. Host plant.— Salix humilis Marsh. as de- VOLUME 101, NUMBER 2 ey) on} Nn Figs. 4-6. Galls of Pontania (Eupontania) spp. 4, P.(E.)? s-pomum (a) and P. (E.) s-desmodioides (type material of P. borealis) (b) on same pin. 5, P. (E.) s-desmodioides (a) and P. (E.) ? s-pomum (b) on same pin. 6, Type material of P. (E.) gracilis on Salix sericea. termined by G. Argus (not Salix sericea as reported by Dyar). Notes.—The valid name is Pontania (Eu- pontania) s-desmodioides (Walsh) (= Pon- tania borealis Dyar, n. syn.). Dyar’s (1898) description of the galls is very similar to those described by Walsh (1866) for P. s- desmodioides. The host plant was misiden- 366 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from which type material was reared. 8, s, type material on Salix petiolaris. pontania) spp. 7, P. (E.) consors, 1 was reared. 9, P. (E.) petiolaridi Figs. 7-9. Galls of Pontania (Eu P. (E.) lucidae, from which the type materia VOLUME 101, NUMBER 2 367 12 od a Figs. 10-12. 10, Apex of abdomen and sheath of Nematus acuminatus. 11, Apex of abdomen and sheath of Pontania populi. 12, Lateral view of P. terminalis. 368 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tified by Dyar. The remnants of leaves with the galls of this species collected by Dyar are shown in Figs. 4, 5. The leaf shape and pubescence of those associated with P. s- desmodioides is typical for Salix humilis, and the willow identification was confirmed by G. Argus. However, along with these galls of P. s-desmodioides (Figs. 4b, 5a), there are also two others on the same pins which appear to belong to P. s-pomum (Figs. 4a, 5b). A typical P. s-pomum gall is in Fig. 2. The remnants of the leaves at- tached to the latter two galls are glabrous and might be Salix eriocephala Michx., the host plant of P. s-pomum. However, none of the three reared females can be identified as P. (E.) s-pomum. Pontania consors Dyar 1898: 121 (galls, lar- va).—Marlatt 1898: 302 (female, male). Type locality—Plattsburgh, N.Y. Type material—Dyar did not give the number of galls and larvae. Marlatt de- scribed adults from one female and two males. The lectotype female, here designat- ed, is labeled: “‘8T; Collection H. G. Dyar; Type No 4132 U.S.N.M.; Pontania consors female Marl.’’ Paralectotypes: 2 males, ‘““8T; Collection H. G. Dyar; Type No 4132 U.S.N.M.; Pontania consors male Marl.”’ Five galls from two leaves are on a pin la- beled: “$i Collection H: G- Dyar (Fig: 7). There is also a cocoon in decayed wood with the label “‘8T; Pontania consors Marl.” Deposited in the USNM. Host plant.— Salix humilis as identified by G. Argus (not Salix sericea as reported by Dyar). Notes.—This is a valid species, Pontania (Eupontania) consors Dyar. Dyar described the species as follows: “‘Galls found with the preceding on S. sericea, but gregarious, hairy and spherical. Near the base of the leaf, three or two together, rarely but one, exceeding the margin often by half the di- ameter of the gall; not evenly divided by the leaf, about one third or a little more above, two thirds below; pale greenish, of- ten heavily marked and mottled with red above, paler below, rarely uniformly pale. Strongly silky hairy like the leaves below, less hairy or even smooth above; size 8.5— 8.5—7 mm or as small as 5 mm in diame- ter.’ The host plant was misidentified by Dyar. The remnants of the leaf were iden- tified by G. Argus, as in the preceding case (P. borealis), as Salix humilis. A. G. Zi- novjev and H. Goulet collected galls of P. consors at Lake Jean Venne, Masham Co., Quebec, Canada (about 50 km N of Ottawa) in the fall of 1995 on Salix humilis (host- plant determined by G. Argus). SPECIES DESCRIBED BY ROHWER S. A. Rohwer did not select single spec- imens as holotypes in his publications, even though he attached holotype or paratype la- bels and USNM type numbers to specimens of each species he described. Therefore, we designate lectotypes where necesary. Pontania amentivora Rohwer 1915: 209. Type locality—Falls Church, Virginia. Type material.—Described from four fe- males. The lectotype female (with a cocoon on the same pin), here designated, is la- beled: ‘10128 Hopk. U.S.; reared May 13 13; Falls Church, Va; S.A. Rohwer Coll.; Type female No. 18313 U.S.N.M.; Pontania amentivora Type Roh.’ Deposited in USNM. Host plant.— Salix sp. Notes.—According to Rohwer “‘this spe- cies lives, in the larval stage, in the pistil- late catkins of a small species of Salix and causes the destruction of the ovaries and the premature forming of ‘cotton’.”’ This spe- cies is currently placed as a synonym of Amauronematus amentorum (Forster) (Smith 1979). The catkin feeders, including this and some related species previously placed in Amauronematus Konow belong in the genus Pontoprista Malaise. Pontania crassicornis Rohwer, 1912: 241. Type locality.—Toronto, Ontario, Cana- da. Type material.—Described from one VOLUME 101, NUMBER 2 14 Figs. 13-15. 15, Anterolateral view of head of P. crassicornis, holotype. male. The holotype is labeled ‘‘ex galls on Salix humilis; Toronto Ont; A. Cosens. Coll.; Type male no. 14572 U.S.N.M; Pon- tania crassicornis TYPE male Roh.; Pontan- ia (Phyllocolpa) crassicornis Rohwer, det. A.Zinovjev 1996.” There are also many dry leaves with galls labeled: ‘‘Pontania crassicornis Roh.? Toronto, Ontario, Cana- 13 Pmcmssisiss 13, Lateral view of P. pumila. 14, Dorsal view of head of Phyllocolpa crassicornis, holotype. da. Cosens; galls of Pontania (Eupontania) consors Dyar, A. Zinovjev det, 1997.” De- posited in the USNM. Host plant.— Salix humilis (original identification confirmed by G. Argus). Notes.—The valid combination is Phyl- locolpa crassicornis (Rohwer), n. comb. The leaves with the galls are Salix humilis 370 and the galls those of P. consors. However, the holotype is a typical leaf roller of the Phyllocolpa leucapsis group. The mandi- bles are asymmetric, the left one in lateral view with a swollen base and a thin blade- like apex, distinctly carinate on the outer surface, and the antennal hollows are gla- brous, shining, and keeled laterally (Figs. 14, 15). Rohwer (1912) mentioned its close relationship to Pontania robusta Marlatt, which is known to produce leaf folds on aspen. Although we are not able to distin- guish the males of Pontania and Phyllocol- pa, we consider this species to be a valid one. Pontania leavitti Rohwer 1910: 199. Type locality.—Nerepis, New Bruns- wick. Type material.—Described from one fe- male. The holotype was redescribed by Smith and Fritz (1996). Deposited in the USNM. Host plant.— Salix sericea, according to Smith and Fritz (1996). Notes.—A valid species of Phyllocolpa in the /eucosticta group. The larvae form leaf rolls on Salix sericea and perhaps some other willows. Pontania lucidae Rohwer, 1912: 242. Type locality—Toronto, Ontario, Cana- da. Type material.—Described from ‘‘Males and females bred from galls on Salix lucida, by A. Cosens.”” The number of specimens is not stated. The lectotype female, here designated, is labeled: ‘‘ex galls on Salix lucida: Toronto Ont; A. Cosens Coll.; Type female No. 14571 U.S.N.M.; Pontania lu- cidae TYPE female Roh.” Paralectotypes: 4 females, 5 males with labels as on the lectotype, except ‘‘Paratype No. 14571 U.S.N.M.”? One male bears the label *‘Pon- tania lucidae male allotype Roh.’ One of the males does not have a red type label, but perhaps it also belongs to the type se- ries. There are remnants of six dry leaves with galls (Fig. 8) labeled ‘‘Pontania luci- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dae Roh. Salix lucidae. Cosens. Toronto, Canada.”’ Deposited in the USNM. Host plant.— Salix lucida (original iden- tification confirmed by G. Argus). Notes.—A valid species in Pontania (Eu- pontania), closely related to Pontania con- sors aS was stated by Rohwer. The exam- ined leaves of Salix lucidae bear only a few typical leaf galls associated with midribs. The rest of the galls in this sample might rather be called petiole or midrib galls (Fig. 8). We are not sure if all of these galls be- long to the species described by Rohwer or if some of them are Euura galls. Pontania petiolaridis Rohwer 1917: 19. Type locality——Toronto, Ontario, Cana- da. Type material.—**Described from a num- ber of females and males reared by A. Cos- ens from galls on Salix petiolaris.”” Num- bers of specimens were not given. The lec- totype female, here designated, is labeled: “Salix petiolaris; Toronto Ont; A. Cosens Coll.; Type female No. 20697 U.S.N.M.; Pontania petiolaridis TYPE female Roh.” Paralectotypes: 4 females, 2 males with the same labels, except “‘Paratype No. 20697 U.S.N.M.”’ One male bears the label *‘Pon- tania petiolaridis male allotype Roh.”’ There are many dry galls with remnants of leaves and a label: “Paratype No. 20697 U.S.N.M.; Pontania petiolaridis TYPE galls Roh.” Deposited in the USNM. Host plant.— Salix petiolaris Sm. (iden- tification confirmed by G. Argus). Notes.—This is a valid species in Pon- tania (Eupontania), is very closely related to P. gracilis, and has galls (Fig. 9) similar to those of P. gracilis (Figs. 3, 6). The galls and biology were described by Cosens (1917) and probably by Dyar (1897b) under the name Pontania gracilis (host-plant “*Sa- lix petiolata’) from Van Cortland Park, New York City, and Gouverneur, N.Y., on ‘Salix petiolata.”’ The name “‘petiolata’’ is obviously a misprint for S. petiolaris (G. Argus, personal communication). Also, the remnants of leaves with the galls are gla- VOLUME 101, NUMBER 2 brous and might belong to S. petiolaris. A. G. Zinovjev reared this species from galls collected by R. Vanderkam near Ottawa, Ontario, in 1995. Pontania pumila Rohwer 1910: 198. Type locality—St. John, New Bruns- wick. Type material.—Described from one fe- male and one male. The lectotype female, here designated, is labeled: ““St. John, New Brunswick”’ (female, July 14). Paralecto- type: male, ‘““Nerepis NB 22 Jul; AG Leav- itt; TYPE male No 12920 U.S.N.M.; Pon- tania pumila Roh. TYPE male.”’ Deposited in the USNM. Host plant.—Unknown. Notes.—This is a valid species, Pontania (Pontania) pumila Rohwer. It very likely belongs in the leaf-rolling crassispina group. The shape of the sheath (Fig. 13) and ovipositor are typical for this group. ACKNOWLEDGMENTS We thank G. Argus, Ottawa, Ontario, Canada, for identification of the willows. E. R. Hoebeke kindly loaned the type of Pon- tania pectoralis Marlatt. Robert Fritz, Vas- sar College, Poughkeepsie, N.Y., helped with collection of galls in Otsego Co., N.Y. Part of this work was supported by a Smith- sonian Institution Short-Term Visitors Grant to A. G. Zinovjev. We thank the fol- lowing for review of the manuscript: N. Schiff, USDA, Forest Service, Stoneville, Miss., and M. A. Solis and M. E. Schauff, Systematic Entomology Laboratory, USDA, Washington, DC. LITERATURE CITED Benson, R. B. 1958. Hymenoptera (Symphyta). Hand- books for the Identification of British Insects, Royal Entomological Society of London, Vol. 6, Part 2(c); pp. 139—252. Cosens, A. 1917. Reports on Insects of the Year. Di- vision No. 3, Toronto District. Forty-Seventh An- nual Report of the Entomological Society of On- tario 1916, pp. 18-19. Dyar, H. G. 1897a. On the larvae of certain saw-flies 371 (Tenthredinidae). Journal of the New York Ento- mological Society 5: 18-30 1897b. New sawflies (Tenthredinidae) with descriptions of larvae. Journal of the New York Entomological Society 5: 190—201. . 1898. On the larvae of certain Nematinae and Blennocampinae with description of new species. Journal of the New York Entomological Society 6: 121-138. Marlatt, C. L. 1896a. Revision of the Nematinae of North America, a subfamily of leaf feeding Hy- menoptera of the family Tenthredinidae. United States Department of Agriculture, Division of En- tomology, Technical Series Number 3, 135 pp. . 1896b. Some new nematids. Canadian Ento- mologist 28: 251-258. . 1898. Some new nematids. Canadian Ento- mologist 30: 302—304. Norton, E. 1864. Notes on Tenthredinidae, with de- scriptions of new species. Proceedings of the En- tomological Society of Philadelphia 3: 5—16. Rohwer, S. A. 1910. On a collection of Tenthredino- idea from eastern Canada. Proceedings of the United States National Museum 38: 197-209. . 1912. Notes on sawflies, with description of new species. Proceedings of the United States Na- tional Museum 43: 205-251. . 1915. Descriptions of new species of Hyme- noptera. Proceedings of the United States National Museum 49: 205-249. . 1917. Pontania petiolaridis, new species, p. 19. In Cosens, A. Reports on insects of the year, Division No. 3, Toronoto District. Forty-Seventh Annual Report of the Entomological Society of Ontario, 1916, pp. 18-19. Smith, D. R. 1979. Symphyta, pp. 3-137. In Krom- bein, K.V. et al., eds. Catalog of Hymenoptera in America North of Mexico, Vol. 1, Smithsonian Institution Press, Washington, D.C. Smith, D. R. and R. S. Fritz. 1996. Review of the eastern United States species of the leaf-folding sawflies of the genus Phyllocolpa Benson (Hy- menoptera: Tenthredinidae). Proceedings of the Entomological Society of Washington 98: 695— 707. Vikberg, V. 1982. Notes on the taxonomy and the no- menclature of some mainly Fennoscandian saw- flies (Hymenoptera, Symphyta). Notulae Ento- mologicae 62: 61—65. Walsh, B. D. 1866. On the insects, Coleopterous, Hy- menopterous and Dipterous, inhabiting the galls of certain species of willow.—Part 2d and last. Proceedings of the Entomological Society of Phil- adelphia 6: 223-288. Zinovjev, A. G. 1993. Subgenera and Palaearctic spe- cies groups of the genus Pontania, with notes on the taxonomy of some European species of the viminalis-group (Hymenoptera: Tenthredinidae). Zoosystematica Rossica 2: 145-154. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 372-378 DESCRIPTION OF IMMATURE STAGES OF PLATYVELIA BRACHIALIS (STAL) (HETEROPTERA: VELITIDAE) PETER P. KORCH, STEVEN L. KEFFER, AND ELISA WINTER (PPK) 1911 Bigler Avenue, PO. Box 367, Spangler, PA 15775, U.S.A.; (SLK, EW) Department of Biology, James Madison University, Harrisonburg, VA 22807, U.S.A. (SLK e-mail: keffers]@jmu.edu) Abstract.—The five immature instars of the broad-shouldered waterstrider, Platyvelia brachialis, are described for the first time. Of three characters considered diagnostic for adult Platyvelia only one, patches of silvery setae on the abdominal dorsum, was observed on immatures of P. brachialis. Key Words: immature insects During a collecting trip to northeastern North Carolina in August 1994, the two se- nior authors collected a complete series of immature instars for the veliid species Pla- tyvelia brachialis (Stal). We here provide the first description of the five immature in- stars of this species. Readers wishing to learn more about the adults of Platyvelia brachialis should consult Polhemus and Polhemus (1993) and the references therein. Excellent, more general treatments of ve- liids can be found in Andersen (1982), Smith (1988), and Schuh and Slater (1995). MATERIALS AND METHODS Specimens were collected along the banks of stagnant bodies of water at the fol- lowing localities in northeastern North Car- olina: Washington Co., Thirty Foot Canal, near junction of Thirty Foot Canal Rd. and Tom Pepper Rd., 12, 13 August 1994; Washington Co., ca. | km. E. of Creshell near junction of Old Cherry Rd. and Springhill Rd., 12 August 1994; Tyrell Co., Batavia Canal, ca. 2 km N. or Lake Phelps, 13 August 1994. They were collected by disturbing the vegetation along the banks. Broad-shouldered waterstriders, Platyvelia brachialis, Veliidae, Heteroptera, The veliids would then walk out onto the water, away from the banks, at which time they were easily collected from the water surface with an aspirator. In addition to the immatures described below, 27 adult males and 24 adult females were collected. All the adults were macropterous. Descriptions were based on 3 first, 4 sec- ond, 5 third, 6 fourth, and 4 fifth instar specimens. Dorsal habitus illustrations were produced using a camera lucida mounted on a Zeiss Stemi SV6 stereomicroscope. Voucher specimens have been deposited in the first author’s private collection. DESCRIPTIONS First instar (Fig. 1).—Length, 1.25 mm = 10:09 mm;_.width;0.6lamnie2=, 0:03 amm: Body elongate and elliptical, greatest width at mesothorax; general ground color ochre- ous; covered with dense, short setae. Head declivent, dorsally brown, ventrally ochreous, vertex with dark median stripe; anteclypeus protruding, rounded anteriorly; remainder of head excluding eyes subrec- tangular; ecdysial lines ochreous, Y-shaped, stem wider than arms, arising from poste- VOLUME 101, NUMBER 2 Figs. 1-2. Platyvelia brachialis. 1, First instar. 2, Second instar. Scale bar mm. LoS) WwW 374 rior margin of head, arms extending to an- teroinferior margin of eyes. Eyes globose, red, separated by ca. 1.6X width of eyes; ocular setae absent. 3 pairs of trichobothria present, Ist pair just posterior to antecly- peus, 2nd pair posterolateral to Ist pair, 3rd pair just medial to inner margin of eyes. Antenna 4-segmented, brown with seg- ments | and 2 darker than 3 and 4; segment 1 curved, segment 2 shortest, segment 4 longest and tapered apically. Beak 4-seg- mented, segments 1-3 ochreous, 4 brown; segment 2 shortest, segment 3 longest, seg- ments | and 4 subequal. Thoracic nota each with pair of sclero- tized, brown subrectangular plates covering most of dorsum, each wider laterally, sep- arated medially by an ochreous ecdysial membrane which widens posteriorly; pro- notal plates extending further laterally than meso- and metanota, mesonota extend only slightly further laterally than metanota. Dark fovea present on each pronotal plate roughly in line with anteromedial corner of eye; shallow oblique depression lateral to fovea extending anterolaterally. Meso- and metanota with dark longitudinal markings in line with posteromedial corner of eyes. Thoracic pleura and sterna ochreous, mem- branous; supracoxal lobes sclerotized and pigmented brown. Legs brownish dorsally, ochreous ven- trally, setose throughout; mesothoracic leg longest. Prothoracic leg somewhat raptorial; coxa globose distally, subequal in length to trochanter; trochanter ventral margin longer than dorsal, subequal in length to tarsus, 5— 7 spines on ventral surface distally; femur slightly curved, ochreous at distal tip, ca. 1 43% longer than tibia, proximal *%4 of ventral margin with double row of spines; tibia slightly curved, row of short, thick spines anteroventrally, longer, thinner spines pos- teroventrally, tuft of hairs anterodistally, grooming comb distally and posteroventral; tarsi straight, 1-segmented, subequal in length to trochanter, short, paired claws subapical. Mesothoracic coxa globose dis- tally, subequal in length to trochanter; tro- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON chanter without ventral spines, ca. % length of tarsus; femur slightly curved, ochreous at distal tip, double row of spines absent, subequal to slightly longer than tibia; tibia straight, ventral spines absent, anterodistal hair tuft present but less dense than on pro- tibia, grooming comb present; tarsus straight, ca. % length of tibia, subapical claw slender, longer than protarsal claw. Metathoracic coxa elongate, subrectangular, constricted proximally but not rounded dis- tally, shorter than trochanter; trochanter without ventral spines, subequal in length to tarsus; femur slightly curved, ochreous at distal tip, subequal to slightly longer than tibia, single row of 4—6 short, thick spines ventral; tibia straight, single row of 13-15 thin recurved spines on ventral surface, an- terodistal hair tuft absent, grooming comb present, tarsus straight, ca. 4% length of tibia, subapical claw similar to mesotarsal claw. Abdomen ochreous. Tergum | with pair of rectangular mediotergites widely sepa- rated by medial ecdysial membrane, later- otergites absent. Terga 2-8 with pair of rectangular mediotergites, lateral margins slightly lateral to medial margins of 1st me- diotergites, narrowly separated by medial ecdysial membrane; two pairs of lateroter- gites present, the first small, narrowly and irregularly elliptical, found on anterior mar- gin of segments, the second minute, located intrasegmentally, distal to first, both absent on segment 8; distance between medio- and laterotergites decreases posteriorly. Spira- cles present dorsally on segment 1, laterally on segments 2—7. Minute pleurites present on segments 2—7. Venter mostly membra- nous, pair of sclerites present medially on segment 8. Segment 9 cone-shaped, scler- otized dorsally and laterally. Segment 10 only visible ventrally, enclosed dorsally and laterally by segment 9. Second instar (Fig. 2).—Length, 1.70 mm + 0.05 mm; width, 0.74 mm + 0.04 mm. Ground color slightly darker than first. Antennal tubercles more distinct. Pair of dark foveae present on each pro- VOLUME 101, NUMBER 2 notal plate, medial fovea in line with an- teromedial margin of eye, lateral fovea pre- sent in medial end of oblique depression, in line with lateral margin of base of antennae, area between foveae of each plate slightly depressed. Meso- and metanotal plates with dark, narrowly U-shaped longitudinal markings in middle, open anteriorly. Ventral surface of legs brownish. Protho- racic femur ca. 1.4 length of tibia; faint yellow band in distal 4%. Tibia with faint ochreous band in middle third. Mesothorac- ic trochanter with pair of denticles on dis- toventral aspect. Femur with faint yellow band in distal third. Tibia with faint ochre- ous band in middle third; double row of spines present, shorter and fewer than that of profemur. Tarsus ca. 0.8X length of tibia. Metathoracic trochanter with 3—4 short spines on distoventral aspect. Femur with double row of 7—9 ventral spines, dorsal row longer, more robust; faint yellow band in distal half. Tibia with faint ochreous band in middle %. Abdominal mediotergites of segment 1 lightly pigmented, subrectangular, narrowly separated by ecdysal line; mediotergites of segments 2—8 greatly reduced, taking on an irregular elliptical shape, located anteriorly along intersegmental fold; all mediotergites forming a parallel row in line with the me- dial prothoracic foveae. Laterotergites pres- ent on segment 1 as small elliptical disks along intersegmental membrane with meta- thorax. Segment 9 with ecdysial line. Otherwise similar to first instar. Third instar (Fig. 3).—Length, 2.10 mm + 0.16 mm; width, 0.95 mm + 0.04 mm. Ground color darker. Eyes separated by ca. 1.2 width of eye. Antennal segment | subequal to segment 4. Posterior margin of pronotum extends posterior at midline. Lateral foveae in line with posteromedial margin of eyes. Meso- notal wing pads present, extending poste- riorly ca. ¥ length of metanotum. Metanotal wing pads extend ca. % length of first ab- dominal segment. Leg bands more distinct; proximal % of S78) tarsi ochreous. Prothoracic trochanter with 8—12 spines on distoventral surface; femur ca. 1.6X longer than tibia, spines extend nearly length of ventral surface. Mesotho- racic tarsus ca. 0.7X length of tibia. Meta- thoracic femur with double row of 9-11 short, thin spines; femur slightly longer than tibia. Sternite on abdominal segment 8 heavily setose. Otherwise similar to second instar. Fourth instar (Fig. 4).—Length, 3.04 mm + 0.18 mm; width 1.13 mm + 0.10 mm. Body more elongate, greatest width at metathorax. Ground color darker. Eyes sep- arated by ca. 1.6 width of eyes. First an- tennal segment ca. 1.4 length of fourth segment; darker in proximal 4%. Second an- tennal segment with faint yellow band in proximal %. Mesothoracic wing pad nearly extends to posterior border of metanotum; metathorac- ic wing pad nearly extends to posterior bor- der of first abdominal segment. Mesothoracic tarsus ca. 0.6 length of tibia. Metathoracic tarsus ca. 0.4 length of tibia. Thin patch of silvery pubescence present on distolateral margins of abdominal seg- ments 2-7; ecdysal line forming ovate patch at anterior margin of segment 7, a smaller pair of ochreous spots lateral to ec- dysal patch between the medio- and lateral tergites. Medial % of dorsal surface segment 8 ochreous; sexual differentiation apparent on ventral sternite of segment 8 Otherwise similar to third instar. Fifth instar (Fig. 5).—Length, 4.16 mm + 0.23 mm; width, 1.48 mm, + 0.10 mm. Ground color darker, patterns more dis- tinct. Antennal tubercles more distinct, nearly black; anteclypeus, postclypeus, and posterior margin of head nearly black. Eyes separated by 1.4 width of eyes. First an- tennal segment ca. 1.6 length of segment 4; segments 2 and 3 subequal. Pronotum with dark brown markings ex- tending anterolaterally from lateral foveae to anterior margin of pronotum and poste- 376 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 3—4. riorly % the length of the lateral margin. Posterior % raised, quadrate and widened laterally. Midline elevated, reaching highest point at posterior margin. Wing pad darker, more developed. Me- sonotal pad with a broad, setose, semilunar elevation in medial %4; pads extend poste- Platyvelia brachialis. 3, Third instar. 4, Fourth instar. Scale bar = 1 mm. riorly ca. %4 length of abdominal segment 1, nearly covering metanotal pad. Distal tip of metanotal pad curved laterally, extending just distal to mesonotal pad. Prothoracic femur ca. 1.2 length of tib- ia. Tibia ca. 2.5 length of tarsi, yellow band on posterior of proximal %4; anterior VOLUME 101, NUMBER 2 Bi Fig. 5. surface yellow. Mesothoracic tibia with yel- low band on posterior of proximal '4; an- terior surface yellow. Metathoracic tibia ca. 4.3X length of metatarsus; yellow band in proximal 4%. Pubescence present along distolateral margins of abdominal segments 2—7 more distinct; additional pubescence present in random patterns along dorsal surface of segments 3—7. Segment 7 with dark yellow ovate spot present along anterior portion of ecdysal line. Segment 8 with anterior mar- gin dark yellow, expanding along ecdysal line posteriorly; ventral aspect completely covered by sternites. Segment 9 usually with most of dorsal surface yellow. Otherwise similar to fourth instar. DISCUSSION Polhemus and Polhemus (1993) list three apomorphies for the genus Platyvelia. The first, opposing metasternal and mesoaceta- bular tubercles, are not present on immature specimens of P. brachialis. This is there- fore an adult character. The second apo- Platyvelia brachialis. fitth instar. Scale bar = | mm. morphy, silvery setae on the abdominal dorsal surface (and on the hemelytra of winged adults), begins in the 4th instar, but only as small intersegmental areas between segments 2—7. Larger areas of silvery setae appear on the dorsal abdominal surface of instar 5, but these are not present in a dis- tinct pattern. The final apomorphy is the lack of ocular setae, a trait shared with Stei- novelia. We found no evidence of ocular setae in any instar. Thus, of three characters considered diagnostic for adult Platyvelia only one, patches of silvery setae on the abdominal dorsum, was observed on im- matures of P. brachialis. ACKNOWLEDGMENTS We thank John T. Polhemus and an anon- ymous reviewer for their critical review of the manuscript. LITERATURE CITED Andersen, N. M. 1982. The Semiaquatic Bugs (He- miptera, Gerromorpha): Phylogeny, Adaptations, Biogeography, and Classification. Entomonograph 378 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 13. Scandinavian Science Press, Klampenborg, the World (Hemiptera: Heteroptera). Cornell Uni- Denmark. 455 pp. versity Press, Ithaca. 336 pp. Polhemus, J. T. and D. A. Polhemus. 1993. Two new Smith, C. L. 1988. Family Veliidae Amyot and Ser- genera for New World Veliinae (Heteroptera: Ve- ville, 1843, pp. 734-742. In Henry, T. J. and R. liidae). Journal of the New York Entomological C. Froeschner, eds. Catalog of the Heteroptera, or Society 101(3): 391-398. True Bugs, of Canada and the Continental United Schuh, R. T. and J. A. Slater. 1996. True Bugs of States. E. J. Brill, Leiden. xix + 958 pp. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 379-381 CRASPEDOLEPTA EAS (MCATEE) AND TRIALEURODES PHLOGIS RUSSELL (HEMIPTERA: STERNORRHYNCHA: PSYLLIDAE AND ALEYRODIDAE): NEW DISTRIBUTIONAL AND HOST-PLANT RECORDS OF TWO LITTLE-KNOWN PHLOX SPECIALISTS A. G. WHEELER, JR. Department of Entomology, Clemson University, Clemson, SC 29634-0365, U.S.A. (e-mail: awhlr@clemson.edu) Abstract.—The psyllid Craspedolepta eas (McAtee) and the whitefly Trialeurodes phlogis Russell are seldom-collected, univoltine specialists on Phlox species (Polemoni- aceae), especially moss phlox, P. subulata L., in shale barrens. Craspedolepta eas is reported for the first ttme from Maryland, North Carolina, and South Carolina, with ad- ditional records given for Virginia and West Virginia. Phlox nivalis Lodd. is a new host of C. eas. The whitefly 7. phlogis, previously known only from the type locality in Virginia, is newly recorded from Maryland, Pennsylvania, South Carolina, and West Vir- ginia, with additional records given for Virginia. Phlox nivalis also is a new host of this aleyrodid. Key Words: Moss phlox (Phlox subulata L.; Pole- moniaceae) is a mat-forming, somewhat woody (suffruticose) perennial that harbors a diverse insect fauna, especially in mid- Appalachian shale barrens and outcrops. Its fauna includes two recently described spe- cies (Henry 1979, Russell 1993) and sev- eral undescribed species, as well as spe- cialist herbivores whose association with phlox became known only with recent at- tention to this plant (Wheeler 1994; 1995a,b). Herein, I provide new distribu- tional and host-plant data for two little- known sternorrhynchans that are phlox spe- cialists. Craspedolepta eas (McAtee) This psyllid was described from Mary- land in the vicinity of Plummers Island and from Great Falls (McAtee 1918). No addi- tional records were available, nor were host relationships known, until Wheeler (1994) Insecta, monophagy, distribution, shale barrens reported it from Phlox species in Illinois, Missouri, Pennsylvania, Virginia, and West Virginia. Craspedolepta eas develops on the narrow-leaved P. subulata in shale bar- rens and on broader-leaved phloxes, P. di- varicata L. and P. stolonifera Sims, of more erect growth habit in moist woods. Nymphs of this univoltine psyllid overwin- ter at the base of their hosts and in spring resume feeding on stems near ground level. Adults begin to appear in early to mid-April and are present until mid- to late May. In- fested hosts are most readily detected by looking for the psyllid’s white, waxy secre- tions and honeydew in the crowns of moss phlox, or on basal stems in the case of more erect, herbaceous species of Phlox (Wheeler 1994). The numbers in parentheses below refer to adults. Records.—_MARYLAND: Allegany Co., High Germany shale barren E. of Little Or- 380 leans, 6 May 1994 (1); Oldtown shale bar- ren, 6 May 1994 (2). Washington Co., Boy Scout shale barren, Sideling Hill Wildlife Management Area, 6 & 15 May 1994 (>10). NORTH CAROLINA: Granville Co., Rt. 15, 0.6 km S. of Bullock, 29 Apr. 1997 (4). SOUTH CAROLINA: York Co., Blackjacks Heritage Preserve, S. of Rock Hill, 20 Apr. 1997 (3). VIRGINIA: Meck- lenburg Co., Rt. 15 at State Line Rd., 1.3 km N. of North Carolina state line, 29 Apr. 1997 (5); Rockingham Co., Forest Service Rd. 87, George Washington National For- est, W. of Fulks Run, 18 May 1994 (1); Shenandoah Co., Edinburg Gap, 4 km E. of Edinburg, 30 Apr. 1994 (3). WEST VIR- GINIA: Pendleton Co., SE. of Upper Tract, 24 Apr. 1994 (1). Remarks.—Most of the new records of C. eas are from the Valley and Ridge Phys- iographic Province except those from North Carolina, South Carolina, and Mecklenburg County, Virginia, which extend the known distribution to the Piedmont. The host plant in the Piedmont was the narrow-leaved P. nivalis Lodd., a new host record for this psyllid. For all other new distributional rec- ords, the host was P. subulata. Trialeurodes phlogis Russell This whitefly was described from the Short Mountain shale barren near Mount Jackson, Va., on the basis of pupae I col- lected on P. subulata in mid-April 1991 and 1992 (Russell 1993). Trialeurodes phlogis has remained known only from the type lo- cality. Taxonomy of the Aleyrodidae is based on pupae; adults, therefore, cannot be iden- tified with certainty. In the case of T. phlo- gis, the adults I collected from mat-forming phloxes are almost certainly those of this species. My initial observation of adults on P. subulata at Shanks, W. Va., in late April led Louise Russell to suspect that the white- fly involved might be an uncommon spe- cies; an early-season emergence of adults is atypical in the genus Trialeurodes (L.M. Russell, personal communication). Trialeu- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rodes phlogis is the only aleyrodid known from wild phloxes, Britton’s (1902) record of the greenhouse whitefly, 7. vaporarior- um (Westwood), from Phlox likely pertain- ing to plants in a greenhouse or garden (Russell 1993). Moreover, adults collected from phloxes of prostrate growth habits are gray, with dark markings on the forewings, matching the appearance of those I reared from pupae on foliage of P. subulata at the type locality of 7. phlogis. The numbers in parentheses below refer to adults unless otherwise noted. Records—-MARYLAND: Allegany Co., High Germany shale barren, E. of Little Or- leans, 6 May 1994 (1). PENNSYLVANIA: Chester Co., Unionville serpentine barrens, NE. of Unionville, 14 May 1994 (1). SOUTH CAROLINA: Pickens Co., nr. Todds Creek S. of Six Mile, 11 Apr. 1998 (2); York, .Co.;/Rt- 77. Nz of junc. RE .o0ik S. of Rock Hill, 18 Apr. 1992 (1). VIRGIN- IA: Alleghany Co., Rt. 18, 17 km S. of Covington, 6 May 1990 (1); Bath Co., Rt. 678, Fort Lewis shale barren nr. Cowpas- ture River, 30 Apr. (2) & 20 May 1994 (3); Mecklenburg Co., Rt. 15 at State Line Rd., 1.3 km N. of North Carolina state line, 29 Apr. 1997 (1). WEST VIRGINIA: Green- brier Co., Kates Mountain, S. of White Sul- phur Springs, 6 May 1990 (2), 12 May 1991 (1 pupal case), 1 May 1994 (2); Whites Draft Rd., E. of Alvon, | May 1994 (3); Hampshire Co., Rt. 220, S. of Purgits- ville, 14 May 1989 (2); Rt. 50, Shanks, 29 Apr. (>20) & 14 May 1989 (>10). Remarks.—The host plant from which adults were collected was P. subulata ex- cept in Mecklenburg County, Virginia, and in South Carolina, where P. nivalis was the host. This plant is a new host record for 7. phlogis. Trialeurodes phlogis is a characteristic, though obscure, insect of mid-Appalachian shale barrens and shale outcrops. I also found it on P. subulata in a Pennsylvania serpentine barren. Phlox nivalis was a host in the Piedmont of Virginia and South Car- olina. The pupae are difficult to find on the VOLUME 101, NUMBER 2 upper or lower surfaces of the needlelike leaves of P. nivalis or P. subulata. Adults can be collected by shaking mats of host phloxes over a pan or tray. Unlike the psyl- lid C. eas, which occurs on phloxes of pros- trate and erect growth habits, 7., phlogis has been found only on mat-forming phlox- es. On the basis of my observations at the type locality in Virginia, 7. phlogis is a uni- voltine whitefly. It overwinters as a third- stage larva (and possibly also as a pupa), with the adults emerging during the latter half of April. Adults are present until mid- to late May. ACKNOWLEDGMENTS I am grateful to Louise Russell (USDA, Systematic Entomology Laboratory) for de- scribing the new whitefly species found on phlox and to Gary Miller (USDA, System- atic Entomology Laboratory) for depositing voucher material of both species in the Na- tional Museum of Natural History, Belts- ville, Md. For providing directions to sites with Phlox nivalis or P. subulata, facilitat- ing the process of obtaining collecting per- mits, or accompanying me in the field, I thank Steve Bennett (South Carolina De- partment of Natural Resources), Ashton Berdine (The Nature Conservancy of Mary- land), Richard Hoffman (Virginia Museum 381 of Natural History), Carl Keener (Pennsyl- vania State University), John Nelson (Uni- versity of South Carolina), Tom Rawinski (Massachusetts Audubon Society), and Doug Samson (The Nature Conservancy of Maryland). My colleague Peter Adler (Clemson University) provided useful sug- gestions for improving the manuscript. LITERATURE CITED Britton, W. E. 1902. The white-fly or plant-house A/- eyrodes. Connecticut Agricultural Experiment Station Bulletin 140 (Entomological Series 8): 1— 17s Henry, T. J. 1979. Descriptions and notes on five new species of Miridae from North America (Hemip- tera). Melsheimer Entomological Series 27: 1—10. McAtee, W. L. 1918. Psyllidae of the vicinity of Wash- ington, D.C., with description of a new species of Aphalara (Hom.). Entomological News 29: 220— 224. Russell, L. M. 1993. A new species of Trialeurodes (Homoptera: Aleyrodidae) from Phlox. Proceed- ings of the Entomological Society of Washington 95: 583-586. Wheeler, A. G., Jr. 1994. Craspedolepta eas: distri- bution, hosts, and habits of a phlox specialist (Ho- moptera: Psylloidea: Aphalaridae). Proceedings of the Entomological Society of Washington 96: 91— 97. . 1995a. Insects of moss phlox (Phlox subula- ta): unexpected diversity in Appalachian shale barrens. Virginia Journal of Science 46: 148 (Ab- stract). . 1995b. Plant bugs (Heteroptera: Miridae) of Phlox subulata and other narrow-leaved phloxes in eastern United States. Proceedings of the En- tomological Society of Washington 97: 435—451. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 382-390 A NEW SPECIES OF CRASPEDOXANTHA BEZZI FROM TANZANIA AND A REVISED PHYLOGENY FOR THE GENUS (DIPTERA: TEPHRITIDAE) AMNON FREIDBERG Department of Zoology, The George S. Wise Faculty of Life Sciences, Tel Aviv Uni- versity, Tel Aviv 69978, Israel (e-mail: afdipter@ post.tau.ac.il) Abstract.—Craspedoxantha yarivi, n. sp., is described from specimens collected on, and reared from, Vernonia calvoana ssp. usambarensis C. Jeffrey (Asteraceae), in the western Usambara Mountains, Tanzania. The species is included in a refined phylogeny for the genus in which the previously proposed arrangement of two species groups remains stable. Characters of immature stages are described and compared to those of two other species in the genus. Key Words: Freidberg (1985) revised Craspedoxan- tha Bezzi (Tephritidae: Tephritinae: Terel- liini) and included nine species. Freidberg and Mathis (1990) described a tenth species and analyzed the phylogenetic relationships within the genus using Hennig86© (see Fitzhugh 1989 for description). Their cla- distic analysis resulted in two well-resolved trees (consistency index 86), comprising two well-supported species groups: the marginalis group, with four Afrotropical species (marginalis (Wiedemann), milleri Freidberg, polyspila Bezzi, unimaculata Bezzi); and the manengubae group, with four Afrotropical species (bafut Freidberg and Mathis, manengubae Speiser, vernoniae Freidberg, yaromi Freidberg) and two Ori- ental species (indica Zaka-ur-Rab, octo- punctata Bezzi). During two recent trips to the western Usambara Mountains, in northeastern Tan- zania, Yariv Malihi and I collected and reared a new species of Craspedoxantha from its host plant, Vernonia calvoana ssp. usambarensis C. Jeffrey (Asteraceae: Ver- nonieae). Morphologically, this species of Craspedoxantha at first appeared to be in- Craspedoxantha, C. yarivi n. sp., fruit flies, phylogeny, Vernonia termediate between the two species groups of Craspedoxantha. It was therefore of in- terest to include it in a revised cladistic analysis and to see whether its addition would affect the overall phylogenetic pic- ture. The new species is described below, and a cladistic analysis follows. Comments on two other species of Craspedoxantha are included. MATERIALS AND METHODS Adult flies were swept from host plants and reared from flower heads placed in or- gandy bags. Abdomens of some males and females were dissected, relaxed in 10% KOH, washed in water and preserved in glycerin inside microvials pinned with the specimens. While embedded in glycerine gel, the terminalia were studied and drawn, using a camera lucida. Puparial characters were studied and drawn in a similar manner. The procedures for the cladistic analysis are explained in the appropriate section below. Terminology follows Freidberg (1985) and Freidberg and Mathis (1986, 1990). The ho- lotype and most other specimens are de- posited in the entomological collection, De- VOLUME 101, NUMBER 2 Figs. 1-3. male, dorsal view. partment of Zoology, Tel Aviv University (TAU). RESULTS Craspedoxantha yarivi Freidberg, new species (Figs. 1-13) Diagnosis.—Craspedoxantha yarivi dif- fers from all its congeners by the following unique combination of characters: Scutel- lum with black spots, and anepisternum with two setae (characters of the marginalis species group); femora not densely setulose ventrally, and dorsocentral seta aligned an- terior of anterior supra-alar seta (characters of the manengubae species group). Other unique characters are blackish stripe bor- dering postpronotum dorsally (lacking in all other species), a distinct infuscation bor- dering crossvein dm-cu (lacking in all other species, except C. milleri, which has a more or less well defined crossband over this vein), and vein M ratio (ratio between api- cal and penultimate sections of this vein) about 1.5 (larger than 2 in all other species). Description.—Specimens of this species Characters of Craspedoxantha yarivi. 1, Head, lateral view. 2, Thorax, dorsal view. 3, Abdomen, were compared to the detailed generic de- scription available for Craspedoxantha (Freidberg 1985). Discrepancies are noted below, accompanied by other characters considered significant. Specimens were brown in life. Chaetotaxy conforms with that of the genus and tribe (Freidberg 1985): Head (Fig. 1): Head structure quite dif- ferent from generic description and Fig. | of Freidberg (1985): Head relatively longer, with frons and frontofacial margin rounded, ventral facial margin strongly projecting and postgena more prominent; eye only about 1.6 times as high as long; face rela- tively low; antenna relatively short; Ist flagellomere 1.5 times as long as wide. Thorax (Fig. 2): Presutural supra-alar seta aligned with anterior notopleural seta; dorsocentral seta aligned half way between anterior supra-alar seta and transverse su- ture; two anepisternal setae present. Scutal setulae fine, extending laterally to level of anterior supra-alar seta. Pattern of dark lat- eral spots on scutum and scutellum differs from all other species: anterior supra-alar 384 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Wing of Craspedoxantha yarivi. spot lacking, | additional blackish micro- tomentose spot present, extending from an- terior margin of scutum to presutural seta, bordering postpronotum dorsally; small blackish spot present around base of apical scutellar seta, sometimes extended to, or nearly to, base of basal scutellar seta and to tip of scutellum. Following spots similar to those of other species: presutural, 2 pos- terolateral spots (between wing base and scutellum), dorsocentral and acrostichal spots. Scutellum less convex, lacking black- ish setulae, with sparse whitish setulae es- pecially laterally; pleura mostly yellowish, with darker, rather inconspicuous longitu- dinal stripe at dorsal % of pleural region, extending from propleuron, which is usu- ally with distinct small blackish spot (such spot lacking in other species), to anatergite, which is often also with small blackish spot; katepisternum similarly darker on ventral %. Legs: Of elongate type (Freid- berg 1985, Fig. 5); femora ventrally not densely setulose. Wing (Fig. 4): Vein M ratio about 1.5; point of cell cup small; anterobasal part of wing hyaline, except small blackish spot at middle of cell c; marginal band extends from base of pterostigma and fork of veins R,,, and R,,,; to slightly beyond end of vein M; crossvein dm-cu, including distal sec- tion of vein CuA,, covered by distinct in- fuscate spot; posterior part of crossvein r-m also covered by infuscate spot; wing oth- erwise generally grayish. Length: 6-8 mm. Abdomen (Fig. 3): Predominantly yellow, but with unique pattern in male: Syntergite 1+2 laterally with small blackish spot; T3 and T4 laterally with similar, though larger, spots; each of these tergites with additional small median spot; TS with black, marginal, ring-shaped pattern, leaving greater central part and posteromesal margin yellow. Black pattern in female present or lacking; at maximum expression, syntergite 1+2 with rather large, median, triangular spot; T2 and T3 with small lateral spots; T3—-T6 each with small median spot. Terminalia (Figs. 5-11): Male: Epan- drium rounded in posterior view (Fig. 5), with convex posterior margin in lateral view (Fig. 6); cerci triangular or diamond shaped (Fig. 7), sclerotized and bifurcate apically. Distiphallus (Fig. 8) with long tube spinose, preaedeagal swelling profuse- ly spinulose, and otherwise rather similar to that of C. vernoniae. Female: Aculeus (Figs. 9-10) and spermatheca (Fig. 11) without overt features. Material examined.—Holotype 2: TAN- ZANIA: Usambara Mts. [Mountains], Gol- ogolo, 1,900 m, 23.vili.1996, A. Freidberg, ex flowerhead Vernonia calvoana ssp. usambarensis, ix. 1996. Paratypes: same collection data as holotype (36 6, 30 @); same data, but not reared (1 6, 7 2). Ad- ditional paratypes: TANZANIA: Usambara Mts., Rt. [Route] B124, Gologolo, 1,800 m, 12—13:ix.1992,. A; Breidberg, 2 654249): same data, but ex flowerhead Vernonia cal- voana ssp. usambarensis, September 1992 (3.6, 3. 2). The holotype is in excellent condition, pinned directly through the tho- rax, and is deposited in TAU. Paratypes will be distributed to the National Museum of Natural History, Smithsonian Institution, Washington, D.C., The Natural History Museum, London, National Collection of Insects, Pretoria, and National Museum of Kenya, Nairobi. Host plant, biology and immature stag- es.—The host plant, Vernonia calvoana ssp. usambarensis C. Jeffrey (Asteraceae: Vernonieae), was described from Lushoto District, Usambara Mountains, and is en- VOLUME 101, NUMBER 2 Ww ie.) N / 10 Figs. 5-11. Craspedoxantha yarivi, terminalia. 5, Epandrium, posterior view. 6, Epandrium, lateral view. 7, Cerci, male, posterior view. 8, Distiphallus. 9, Aculeus, whole. 10, Aculeus tip. 11, Spermatheca. 386 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON demic to Tanzania (Jeffrey 1988). It is the host of at least three additional species of tephritids, all apparently undescribed (Freidberg, unpublished observations). Ver- nonia calvoana (Hook. f.) Hook. f. is wide- ly distributed over tropical Africa and con- sists of seven geographically more or less restricted subspecies (Jeffrey 1988). It was recorded as host of C. vernoniae, C. yaromi (Freidberg 1985), C. bafut and possibly C. manengubae (Freidberg and Mathis 1990), but the host subspecies were not indicated. Identification of the relevant host subspe- cies may show that speciation in some Craspedoxantha species may correspond to the level of subspecies of V. calvoana. The same pattern was also observed in some schistopterine tephritids infesting V. cal- voana and other Vernonia Schreb. species (Freidberg, unpublished observations). Hence, if C. yarivi is indeed specific to the subspecies usambarensis, it may also be en- demic to the same small geographical range of its host. Adults of C. yarivi began to emerge from flower heads several days after the plants were collected and put in mesh bags, and emergence continued at least one week. Some puparia were found inside the bags, hence it is assumed that at least some mag- gots may usually leave the flower heads to pupate in the ground. These puparia formed Figs. 12-13. the basis for the study of immature char- acters reported below. A total of ten puparia were studied, some of them lacking parts of the operculum. The puparium is yellow, with distinct segmen- tation. Some puparia were entirely wrin- kled, others were rather smooth, especially dorsally on the central segments. Wrinkles were circular, more or less parallel, some- what irregular, with up to about 10 wrinkles on a single segment. Length: 4.75-6 mm (average: 5.6 mm); width: 2.3-3 mm (av- erage: 2.75 mm). Cephalopharyngeal skel- eton (Fig.12) about 0.9 mm long. Mandible without teeth, although concave (ventral) edge not entirely smooth, otherwise similar to that of C. polyspila and C. marginalis (Janzon 1985). Anterior spiracle (Fig. 13; N = 16) consisting of 9-11 digits (average 10; only 7 in C. polyspila), each 0.05—0.06 mm long. Digits, unlike in C. polyspila, are not arranged in a tight row. In unprepared puparia, digits appear to be 2—3 times lon- ger than in the drawing, but this might be because the roots of the digits are visible under the surface of the cuticle. Posterior spiracle generally similar to that of C. po- lyspila but requires more careful study. Etymology.—The species epithet, yarivi, is a genitive patronym in honor of my stu- dent, Mr. Yariv Malihi, who reared a large number of specimens, which comprise most of the type series. 13 Craspedoxantha yarivi, puparium. 12, Cephalopharyngeal skeleton. 13, Anterior spiracle. VOLUME 101, NUMBER 2 Craspedoxantha bafut Freidberg and Mathis Craspedoxantha bafut Freidberg and Math- is 1990: 325. This species was described from Came- roon and Nigeria based on 13 specimens, some of them reared, and has not been re- corded subsequently. A recent checking of the type material revealed that the holotype (Cameroon, Rt. N6, Bali-Batibo, W. of Ba- menda, 20.xi.1987, A. Freidberg, deposited in TAU) is a male, not a female, as stated in the original description. Craspedoxantha milleri Freidberg Craspedoxantha milleri Freidberg 1985: 195. This species was described based on a single male collected on the Drakensberg, Natal, South Africa. A second male from the same general region is now available for study. It is labeled: Blue Mountain Pass, Makhaleng Valley, Maloti Mountains, 2,150—2,525 m, 12-14 Jan. 1963. Another label: Maseru District, Basutoland, B. & P. Stuckenberg (National Collection of In- sects, Pretoria). This specimen fits the orig- inal description well, except that it is small- er than the male holotype (only 4.5 mm, instead of 5 mm), and the transverse band over the crossveins is narrower and inter- rupted, especially over vein M. The epan- drium has a concave posterior margin in lat- eral view (not studied in the holotype). CLADISTIC ANALYSIS OF CRASPEDOXANTHA For a detailed explanation of the cladistic methodology and its application to Cras- pedoxantha, see Freidberg and Mathis (1990). Trees were calculated from the character data using the “implicit enumer- ation” option of Hennig86. The only dif- ferences from the character matrix of Freid- berg and Mathis (1990) are the addition of Craspedoxantha yarivi (the new species) and Chaetostomella cylindrica Robineau- Desvoidy (as a second outgroup), and some changes, mostly additions, to the characters 387 Table 1. doxantha and two outgroup species. Character matrix for species of Craspe- Taxon (Cl ters) Orellia punctata 0000 100 Chaetostomella cylindrica 001 Craspedoxantha bafut ard Craspedoxantha indica 0211 Craspedoxantha manengubae Craspedoxantha marginalis Craspedoxantha milleri 0100011200001 000001100000? Craspedoxantha octopunctata 020111010000: Craspedoxantha polyspila 0000011210000 Craspedoxantha unimaculata 010001120?00? Craspedoxantha vernoniae ial bab ioab al Oalal Craspedoxantha yarivi 0110000212121 Craspedoxantha yaromi 12d MOOT OOD used in the analysis. A brief presentation of the 13 characters used in the analysis and coding of the character states are given be- low. The character matrix is in Table 1. All characters with three character states were coded ‘‘additive.”’ Thorax 1. Size of presutural black spots: about as large as dorsocentral spots (0); distinct- ly smaller than dorsocentral spots (1). 2. Number of black spots on scutellum: 3—4 (0); 2 (1); O (2). 3. Alignment of dorsocentral seta: with anterior supra-alar seta (0); distinctly anterad of anterior supra-alar seta (1). 4. Number of anepisternal setae: 2 (0); 1 Ci): 5. Lateral extension of scutal setulae: to level of anterior supra-alar seta (0); to level of presutural seta (1). 6. Coloration of scutellar setulae: all or most setulae yellow (0); all or most se- tulae black (1). Legs 7. Structure and vestiture of femora: slen- der and lacking dense investment of se- tulae (0); swollen and densely setulose ventrally (1). 388 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Character 1 2 3 4 5 Steps 2 4 2 2 2 Consistency index 50 50 50 50 50 Retention index 66 75) 80 80 80 Wing 8. Additional band or spot present over crossvein dm-cu and connected or dis- connected to marginal band (0); such band or spot lacking, but marginal band approaching crossvein r-m (1); such band or spot lacking, and marginal band not approaching crossvein r-m (2). 9. Color of cell cup: distinctly yellow (0); indistinctly yellow (1). Abdomen 10. Shape of posterior margin of the epan- drium (the surface from which the cerci arise) in lateral view: concave (0); straight (1); convex (2). 11. Long tube of distiphallus: not spinose (O); spinose (1). 12. Preaedeagal swelling: not spinulose (O); slightly spinulose (1); profusely spinulose (2). Biology 13. Host plants: Host plant associations that are in part or exclusively with- Vernonia Schreb. (tribe Vernonieae) are hypothesized as the derived condi- tion (1). I was unable to hypothesize a complete transformation series for the other host associations, whether they include plants of the tribe Cardueae (hosts of most Terelliini), Lactuceae (host of Orellia Robineau-Desvoidy species) or others, and I therefore cod- ed them all as primitive (0). The results of the cladistic analysis are presented below (Fig. 14), and the analysis of the characters is given in Table 2. DISCUSSION The main purpose of this study was to describe a new species of Craspedoxantha Character analysis for species of Craspedoxantha and two outgroup species. 6 7 8 9 10 11 12 13 3 i} 5 3 p) 2 2 2 33 100 40 33 100 50 100 50 33 100 S77 60 100 10) 100 66 and to revise the phylogeny of this genus. The new species, C. yarivi, possesses most adult characters of this rather homogenous genus. By its morphology, it appears to be an intermediate species between the two previously recognized species groups. However, in the cladistic analysis above, it appears as a terminal clade of the manan- gubae group, not upsetting the basic divi- sion into the two species groups, rather sup- porting it. Yet, the monophyly of these spe- cies groups is now primarily substantiated by two characters (the alignment of the dor- socentral setae and the structure and vesti- ture of the femora), rather than the four characters that were available before C. ya- rivi was discovered (Freidberg 1985). In the cladistic analysis I first used the “implicit enumeration” option. Six trees re- sulted from this option, but only one (Fig. 14) was similar to the Nelson consensus tree of Freidberg and Mathis (1990: fig. 5) in supporting the same (marginalis and ma- nengubae) species groups. Two of the other trees did not support any species grouping, and the other three trees progressively sup- ported a stronger marginalis group, culmi- nating with a resolved group containing C. marginalis, C. unimaculata and C. polys- pila. However, in all these five trees C. mil- leri branched at the base, forming an out- group to all other species of Craspedoxan- tha. The successive weighting technique was then used, resulting in one tree that is identical to the above mentioned tree (Fig. 14) of the “‘implicite numeration” option. This tree is fully resolved and includes C. yarivi as the sister species of C. vernoniae in the manengubae group. Two noteworthy autapomorphies of C. yarivi are the completely whitish-setulose scutellum and the infuscated spot on cross- vein dm-cu. Blackish setulae on the scutel- VOLUME 101, NUMBER 2 (3.1, 4.1) (951513:1) Fig. 14. bers refer to the cladistic section. lum were one of the main diagnostic char- acters of Craspedoxantha (see key to ter- elliine genera in Freidberg 1985), and the entirely whitish setulae on the scutellum of C. yarivi renders the monophyly of this ge- nus less substantiated. The second note- worthy apomorphy, viz. an infuscated spot restricted to crossvein dm-cu, is unique to C. yarivi, although a similar character is found in C. milleri, in which a cross band near the middle of the wing extends over both crossveins r-m and dm-cu (Freidberg 1935). The monophyly of Craspedoxantha is now supported by only two unequivocal characters: the relatively high eye (1.5—2 times as high as long) and the uninterrupted yellow costal band on the wing (spotted with black). The monophyly of the genus may also be supported by zoogeography, as Craspedoxantha is the only tropical genus of Terelliini, and it is disjunct or nearly dis- All (10.2 389 Orellia punctata Chaetostomella cylindrica Cr. margina! Cr. unimaculata Cr. polyspila Cr. milleri . octopunctata . indica (6.0) Cr. yaromi . bafut Cr. manengubae Cr. vernoniae (6.0)— = Cr. yarivi Reconstructed phylogeny for species of Craspedoxantha and two outgroup species. Character num- junct from the other terelliine genera (which are essentially Palearctic and/or Ne- arctic). I suggest continuing to treat Cras- pedoxantha as a valid genus at least until this validity is tested by a cladistic analysis of the entire tribe. The comparison between puparial char- acters of C. yarivi and larval characters of C. polyspila and C. marginalis (Janzon 1985) indicated that characters of immature stages may be significant for the classifi- cation of Craspedoxantha. Some differenc- es in the shape of the cephalopharyngeal skeleton and anterior spiracles were noted. These differences may represent supraspe- cific categories. However, additional spe- cies should be studied before this assump- tion can be substantiated. Additional new species of Craspedox- antha should be expected in Africa, and perhaps also in southeast Asia, especially if all subspecies of Vernonia calvoana and 390 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON other Vernonia species are searched for these flies. ACKNOWLEDGMENTS I am grateful to my student, Yariv Ma- lihi, who diligently helped me during a re- cent collecting trip to Tanzania, especially in rearing the large series of specimens that formed the basis to this paper. I am also grateful to Wayne N. Mathis (Smithsonian Institution) and Allen L. Norrbom (System- atic Entomology Laboratory, USDA, % Smithsonian Institution) for their help and useful suggestions, to Bernhard Merz, ETH, Zurich, for reviewing a final draft of the manuscript, to Fini Kaplan, for continuous encouragement, to Walter Ferguson for the drawings and to Amikam Shoub for the photograph. LITERATURE CITED Fitzhugh, K. 1989. Cladistics in the fast lane. Journal of the New York Entomological Society 97: 234— 241. Freidberg, A. 1985. The genus Craspedoxantha Bezzi (Diptera: Tephritidae: Terelliinae). Annals of the Natal Museum 27: 183-206. Freidberg, A. and W. N. Mathis. 1986. Studies of Ter- elliinae (Diptera: Tephritidae): A revision of the genus Neaspilota Osten Sacken. Smithsonian Contributions to Zoology 439: 1-75. . 1990. A new species of Craspedoxantha and a revised phylogeny for the genus (Diptera: Te- phritidae). Proceedings of the Entomological So- ciety of Washington 92: 325-332. Janzon, L.-A. 1985. Descriptions of 3rd instar larvae of Craspedoxantha polyspila Bezzi and C. mar- ginalis (Wiedemann) (Diptera: Tephritidae) from Gambia. Entomologica Scandinavica 16: 227- 229. Jeffrey, C. 1988. The Vernonieae in east tropical Af- rica. Kew Bulletin 43: 195—277. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 391-411 A NEW GENUS CLYPEOLONTHA LI AND YANG, FOR THE GENUS MELOLONTHA FABRICIUS (COLEOPTERA: SCARABAEOIDEA: MELOLONTHINAE) FROM SOUTHEASTERN ASIA CHUN-LIN LI AND PING-SHIH YANG Laboratory of Insect Conservation, Department of Entomology, National Taiwan Uni- versity, Taipei 10764, Taiwan, Republic of China (e-mail: psyang@ccms.ntu.edu.tw) Abstract.—Clypeolontha, new genus, is described with Melolontha alboplagiata Bren- ske, new combination, as the type species. In addition, three species are herein recognized as new: C. siamensis, C. bertiae, and C. laosensis. The following information is provided for each species, when appropriate: literature review, diagnosis, description or redescrip- tion, illustration of important external characters, data for material studied, geographical distribution, and taxonomic remarks. A key to females and a distribution map to the Clypeolontha species are given. A preliminary discussion on the systematics of Melolon- tha sensu lato is presented. Key Words: As the type genus of the subfamily Mel- olonthinae, Melolontha Fabricius, 1775, has attracted attention because of their diversity and agricultural importance of its species. Burmeister (1855) first treated this genus and assigned Hoplosternus Guérin-Méne- ville, 1838 and the newly defined Schoen- herria as members of Melolontha, although both are later widely accepted as indepen- dent genera. Reitter (1902) proposed four groups mainly based on their geographical distributions in his division of Melolontha, including Tocama, the first subgenus from mainland China. Medvedev (1951) desig- nated the second subgenus, Apropyga, un- der a strict concept of Melolontha in his faunal review. Since these beetles are widely distributed throughout Eurasia, it is surprising that no revision encompassing the some 40 con- stituent species within Melolontha has ever been furnished to date. Most taxonomic works for Melolontha consist of isolated contributions. Medvedev (1951) is probably Melolonthinae, Melolontha, Clypeolontha, new genus the most comprehensive study so far, par- ticularly his subgeneric treatment, which in- cludes all species of Melolontha from the former Soviet Union and adjacent regions. Nomura (1952) reviewed the northeastern Asian Melolontha species and described a related genus, Tricholontha, endemic to the Okinawa Islands, Japan. Baraud (1992) pro- vided a identification key as well as a de- tailed literature review for each of the nine nominal European species of Melolontha. Neither of these works discussed the sys- tematic problems of Melolontha. Since the first Melolontha was described, several species have been removed, or es- tablished as the type species of new genera and more cases will likely be reappraised in the future. Apparently, the broad concept of Melolontha is a result of all allied genera being lumped together. Not only do they all share the common character of a 7-seg- mented antennal club, but sometimes, par- ticularly at the earlier taxonomic works, merely an enlarged antennal club. The type oS) \O N species of the genus Cyphochilus Water- house, 1867, C. candidus (Olivier 1789), is an obvious example with a 3-segmented en- larged club in males and previously consid- ered a Melolontha. Thus, a reasonable clas- sification system within the whole subtribe Melolonthina also needs further construc- tion and analysis. Most recently, Baraud (1992) provided two other diagnostic char- acters, namely the membranous margin of the elytra and the number of antennal club segments in females, to seperate Melolon- tha from Polyphylla Harris, 1941. These two characters and the ratio between eye and interocular width used by Nomura (1952) are characters (in addition to male antennal segments) employed to distinguish Melolontha from other related taxa. In fact, most of those genera closely re- lated to Melolontha (1.e., Polyphylla Harris, Schoenherria Burmeister, 1855 and Exolon- tha Reitter, 1901) are most diverse and are mainly distributed in or restricted to East and Southeast Asia. However, we will pro- vide further information with broad evolu- tionary implications to those, described or undescribed, taxa closely related with Mel- olontha sensu lato which based on cladistic analysis within the next complementary work and this is the first part we refer to the taxonomic assessment of Melolontha sensu lato (see also Systematics). MATERIAL AND METHODS Specimens used in this study were bor- rowed from and deposited in the institutions referred to in the section of material or type. The acronyms follow Arnett et al. (1993) and are listed below. BMNH: The Natural History Museum, London, U.K.; Malcolm Kerley. Institut Royal des Sciences Na- turelles de Belgique, Bruxelles, Belgium; Konjev Desender and Marcel Cludts. Museum of Nature and Human Activities, Hyé6go, Japan; Yo- shihisa Sawada. ISNB: MNHA: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MNHN: Muséum national d’ Histoire na- turelle.* Paris.” France: Jean’ J: Menier and Nicole Berti. National Museum of Natural History, Taichung, Taiwan; Mei- Ling Chan. Insect Museum, Department of Entomology, National Taiwan University, Taipei, Taiwan; Tung-Ching Hsu. Taiwan Agricultural Research In- stitute, Taichung, Taiwan; Liang- Yih Chou. Museum ftir Naturkunde der Humboldt-Universitat, Berlin, Germany; Hella Wendt and Joa- chim Schulze. NMNH: INGE: TARI: ZMNB: Observation and measurements of exter- nal characters were made using an ocular micrometer. Male genitalia were dissected and cleaned using 10% KOH solution for few days, then stored in glycerine in micro- vials and attached to specimens from which they have been removed. The specimen label data has been abbre- viated to indicate as a handwritten (H) ora printed (P) data respectively. Separate la- bels are indicated by double slashes from another while specimens with two more ones. The information on geographical dis- tribution is referred from label data. The measurements and ratios that are considered to be useful in the separation of Clypeolontha species are briefly summa- rized in Table |. Abbreviations for charac- ters and mensural procedures are listed as follows: BL body length, measured from an- terior margin of clypeus to the apex of elytra. BW body width, measured across elytral humeri. BW/HW ratio of body width to head width across eyes in female. MPR ratio of maximum length of fe- male maxillary palpi 2—4. ASR ratio of length of male antennal basal segments 1-3. VOLUME 101, NUMBER 2 393 Table 1. Summary of selected descriptive measurements and ratios for species of Clypeolontha: range, mean and standard deviation. Body length Body width Taxa (n) (BL), mm (BW), mm BL/BW MFL/W PeWiI C. siamensis 16.4-19.6 8.0—9.0 2.05—2.21 2.93-3.21 1 .08-1.22 Male (21) 18.4 (0.84) 8.65 (0.31) 2.13 (0.06) 3.06 (0.06) 1.15 (0.38) Ditto, female (8) 19.7—20.7 9.3-10.0 2.04—2.17 2.34-2.56 1.17-1.25 20.2 (0.22) 9.6 (0.32) 2.09 (0.05) 2.45 (0.08) 1.21 (0.06) C. alboplagiata 16.8—17.0 7.8-8.0 2.13-2.15 2.42—2.50 1.00—1.03 Male (4) 16.9 (0.1) 7.9 (0.1) 2.14 (0.01) 2.46 (0.04) 1.02 (0.02) Ditto, female (3) 19.8—20.1 8.8-9.0 2.23—2.25 2.08—2.18 1.11-1.14 19.95 (0.15) 8.9 (0.1) 2.24 (0.01) 2.13 (0.05) 1125 (0:01) C. bertiae (5) 17.9-18.5 7.8-7.9 2.28—2.36 2.34-2.45 (n = 4) 1.08—1.12 18.2 (0.3) 7.9 (0.06) 2.36 (0.01) 2.40 (0.08) 1.10 (0.02) C. laosensis (1) Wie2 es) 2.29 2.61 1.18 PTR ratio of maximum length of ic placement but will be discussed at a later protarsomeres 1-5. date. MFL/W ratio of length of metafemur to Through examinations of 25 of 41 cur- its maximum width in ventral rently valid species in Melolontha and 20 aspect. of 29 species in Hoplosternus, we present PgW/H ratio of height of pygidium to’ the following diagnostic characters, based its maximum width in dorsal aspect. SYSTEMATICS The taxonomic legacy surrounding the members of the genus Melolontha sensu lato has sufferred many changes in position. These situations may become more compli- cated as more new taxa are discovered. We consider that the lack of systematic research related to the genera of Melolontha, espe- cially Hoplosternus, and lack of taxonomic characters that clearly delimit all closely re- lated genera, are apparently the reasons for confusion regarding the classification of melolonthine taxa. Since we recently ac- quired many related taxa from various geo- graphical areas and the senior author has seen many types and other specimens from several European institutions, it was felt time is appropriate to make a new definition of Melolontha. Therefore, the establishment of new genus Clypeolontha is herein pro- posed to accommodate those questionable taxa, in part, presently assigned to Melolon- tha sensu stricto. Several other taxa are also in need of a reassessment in their taxonom- mainly on the type species, Melolontha melolontha L. for Melolontha sensu lato: (1) presence of metallic coppery with some green or purple coloration on head, prono- tum, scutellum and femora when surface se- tae are removed; (2) overall punctation of pronotum usually coarser and sparser with varied distribution; (3) each elytron with 5 discal costae including 1 along epipleural margin; (4) lateral sides of abdominal ster- na 1—6 usually with a lighter maculation, although on sternite 6 sometimes less de- veloped; and (5) male genitalia in general symmetrical, apex of parameres usually obliquely truncate and swollen when viewed laterally, in frontal aspect with trap- ezoid to broad bean-like swelling, dorsal portion with hook apically. In the time past, Melolontha and Hoplosternus were differ- entiated based only on the appearance and development of mesosternal process. Arrow (1913) first argued against this character when he wrote that the male of M. guttigera Sharp lack the mesosternal process while the female has one. However, we have ex- amined many determined specimens of M. guttigera and found that both sexes did 394 Fig. |. Clypeolontha siamensis. have the mesosternal process although the male has a remarkably less-developed one than female. Regardless, it is our view that the genus Melolontha is probably paraphy- letic when Hoplosternus is treated as an in- dependent derivative from Melolontha. The only distinctive character, the mesosternal process, used to separate Hoplosternus from the other taxa is subject to a broad transitional intra/inter-specific variation, and it is also commonly found in many gen- era of Scarabaeidae. Thus, this character, a plesiomorphic feature, is useless for rec- ognition of Hoplosternus as an independent genus and we will present a formal taxo- nomic treatment in the future. Furthermore, as will be proposed cladistically elsewhere (Li in preparation), members of the true Melolontha lineage are defined by the above-mentioned synapomorphies (1), (2), (4), and (5). All of these characters are shared by Melolontha and Hoplosternus. It is unlikely that these structures evolved twice because they are found in the same geographical distributions of Eurasia with a sympatric distribution pattern and have not been discovered elsewhere. Brenske (1898) described M. alboplagia- ta (here referred to as Clypeolontha), based only on a single female specimen. Even lacking a male, Brenske still assigned this species to a member of Melolontha proba- bly because its wide maculation continu- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ously located along the lateral sides of ab- dominal sternites 1—6 makes it morpholog- ically closed to Melolontha than the other genera. However, this character is not shared by Melolontha and is one of auta- pomorphic characters defining our new ge- nus. Fortunately, we obtained a series of both sexes collected from the type locality and neighboring areas. These were com- pared with the type and other related taxa from Southeast Asia respectively. There- fore, we here propose the new genus, Cly- peolontha for M. alboplagiata together with three other new species. Clypeolontha Li and Yang, new genus Type species. Melolontha alboplagiata Brenske 1898: 236, here designated. Diagnosis.—The following combination of characters separate Clypeolontha from all other related genera within the subtribe Melolonthina. Body oblong, subparallel- sided medially, smaller (16.4—19.6 mm), and absence of metallic coloration or re- flection on body surface, antennae, and legs. Head surface densely rugose, clypeus shallowly depressed, emarginate centrally exposing labrum when viewed dorsally. Pronotal surface densely punctate, punc- tures evenly distributed, each with a seta subequal in length, lateral margins very weakly and incompletely serrate; pro- and mesosternal processes vestigial. Pretarsus ey) Ke) Nn VOLUME 101, NUMBER 2 Figs. 2-5. Dorsal habitus. 2, Clypeolontha siamensis, holotype male. 3, C. alboplagiata, male. 4, C. sia- mensis, female. 5, C. alboplagiata, holotype female. 396 small, subapical tooth one-half to two- thirds length of apical claw. Metepisternum and metepimeron densely covered with scales. Six visible abdominal sternites with lateral maculation on ventrites | through 6, narrowest on ventrite 1, expanding oblique- ly towards middle through to about ventrite 4 and decreasing 5 in width through to 6, which maculation consisting of dense white scales with faint iridescent-tinged reflection continuous throughout the lateral edges. Paramere of male genitalia asymmetrical. Etymology.—tThe prefix of the generic name is derived from the Latin combining form clypeo-, reflecting distinctive charac- ter of clypeus on both sexes among species, and the suffix is partly taken from the genus Melolontha showing their close relation- ship. The gender is feminine. KEY TO FEMALES OF CLYPEOLONTHA The key is based on females only; males material were not available for two species in this study. However, we consider that the distinguishing characters employed herein are constant throughout each species. Ad- ditionally, we provide a diagnosis to the species with male specimens compared. 1. Body size larger (BL = 19.7 mm, BW 2 8.8 mm, BL/BW = 2.25; surface usually clothed with yellowish-brown to whitish-yellow setae .. 2 — Body size smaller; pronotal and elytral surfaces clothed with pale white to yellow setae .... 3 2. Labrum triangular (Fig. 11); apical maxillary palpomere spindly elongate, longer than com- bined segments 2—3 (Fig. 43) (mean MPR = 1.6:1.3:3.7); pronotal midline not visible, an- gles obtuse (Fig. 13); mean BW/HW = 1.64; WwW PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gidium broadened basally with apex rounded and inflated (Fig. 29); PgW/H = 1.17-1.25; northern Thailand ........ apical maxillary palpomere spindly elongate, equal to combined length of segments 2-3 (Fig. 44) (mean MPR = 2.0:1.5:3.5); pronotal disc moderately convex with finely punctate, evenly distributed, midline not visible, angles weakly protuberant (Fig. 14); mean BW/HW = 1.73; protibial spur subequal to one-fourth length of first pretarsomere; hind femur round- ly inflated (Fig. 20) (MFL/W = 2.34-2.45); posterior edges of abdominal sternites 5 and 6 smooth, sternite 6 with feeble irregular serra- tion (Fig. 26); pygidium clothed with pale white to yellowish-brown setae, moderately elongate with apex sharpened (Fig. 32); PgW/ H = 1.08—1.12; western Laos and northeastern Thailand Clypeus moderately depressed anteriorly; api- cal maxillary palpomere spindly stout, shorter than combined length of segments 2—3 (Fig. 45) (MPR = 1.8:1.3:2.7); pronotal disc weakly convex with irregularly distributed punctures, midline shallowly depressed, bearing tiny brownish setae, anterior angle acute, posterior angles obstuse (Fig. 15); BW/HW = 1.91; pro- tibial spur subequal to one-third length of first pretarsomere; hind femur slightly transversely inflated (Fig. 21) (MFL/W = 2.61); posterior edge of abdominal sternite 5 distinctly point- curved, sternite 6 strongly arc-curved medially (Fig. 27); pygidium clothed with yellowish brown setae, slightly elongate with apex round- ed (Fig. 33); PgW/H = 1.18; central north- CANES ANILAOS 255000 case ane C. laosensis, n. sp. Clypeolontha siamensis Li and Yang, new species (Figse 2 as Se lO aaliGsel (ae2 25a. 28; 29, 3455s SOyO7ea2) C. siamensis, n. sp. . Clypeus very shallowly depressed anteriorly; C. bertiae, n. sp. hind femur broadly stout (Fig. 19) (MFL/W = 2.08—2.18); abdominal sternite 5 with posterior edge smooth (Fig. 25); apex of pygidium somewhat sharpened (Fig. 31); PgW/H = 1.11—1.14; Sikkim and Bhutan Labrum rounded (Fig. 10); apical maxillary palpomere elongate, bulged centrally, subequal to combined segments 2—3 (Fig. 42) (mean MPR = 1.9:1.4:3.5); pronotal midline shallow- ly depressed, angles acute (Fig. 12); mean BW/ HW = 1.84; hind femur stout (Fig. 17) (MFL/ W = 2.34—2.56); abdominal sternites 5 and 6 anteriorly point-curved medially (Fig. 25); py- C. alboplagiata Brenske Melolontha alboplagiata Brenske: sensu Itoh 1995: 202 (distribution, new record). Type series.—Holotype d¢ (deposited in NTUI as DPPE-9702): N. THAILAND: Chiang Mai Prov., Fang, 9.V.1995. Paraty- pes (18 3, 7 2) as follows: same data as holotype? 906324 Pin 25sIVAI995- thie remaining 6 d and 2 @, Chiang Rai Prov., Wiang Pa Pao, all on 1.V.1995 but 1 d on 5.1V.1995 (2 6 paratypes deposited in BMNH; 2 ¢ in TARI; 3 6 and 2 & in NMNH; 3 6 and 2 2 in ZMHB; 3 ¢ in VOLUME 101, NUMBER 2 397 Figs. 6—7. ISNB); 1 ¢o dated May 27, 1990 from Wiang Papso, N. Thailand and 1 @ dated May 1, 1988 from Samoeng, near Chiang Mai, N. Thailand in Takeshi Itoh’s collec- tions; | d dated May 10-13 from Doi Song, near Chiang Mai in MNHA The re- maining paratypes, 3 d and 2 2 are depos- ited in NTUI and the authors’ collections, respectively. Male diagnosis.—Antennal club 2.3—2.7 times length of stem; mean ASR = 2.1:0.7: 1.3; labrum shape rounded; pronotal mid- line very shallowly depressed, angles acute; protarsomere | shorter than combined length of protarsomeres 2 and 3 (PTR = Br Ose/Al-641 .6:2:7);0 hind “femur “stout (MFL/W = 2.93-3.21); abdominal sternite 5 with posterior edge moderately arc- shaped centrally; apex of pygidium distinct- ly prolonged, longitudinally convex, PgW/ H = 1.08—1.22; paramere apex irregularly broadened, right paramere with a inward spine on basal one-fourth. Dorsal habitus. 6, Clypeolontha laosnesis, holotype female. 7, C. bertiae, holotype female. Description.—Male. Body: Oblong (Figs. 1, 2), sides subparallel medially. Dorsal sur- face densely covering with short yellowish- brown to whitish-yellow setae, same in length. BL = 16.4-19.6 mm; BW = 8.0— 9.0 mm; BL/BW = 2.05—2.21. Basal color blackish brown to yellowish brown. Head: Surface densely to confluently punctate, punctures moderately large, each bearing a short seta. Antenna 10-segmented with 7-segmented club; lamellae slightly outwardly curved, 2.3—2.7 times length of stem; first basal segment wider than third one; mean ASR = 2.1:0.7:1.3. Clypeus transverse, shallowly depressed anteriorly, weakly emarginated; sides gradually round- ing to biarcuate apex; middle of anterior margin shallow and smooth (Fig. 8); cly- peo-frontal suture moderately developed; frons slightly narrowed, making eyes large. Labrum weakly to moderately grooved with respect to clypeus; transversally symmetri- cal, rounded and strongly bilobed, one-third 398 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 8-11. mensis, front view. 11, C. alboplatiata, front view. Scale line = 2.0 mm. width of clypeal base, depressed laterally (Fig. 10); visible dorsally, moderately pro- tuberant. Maxillary palpus light yellowish brown, 4-segmented; apical palpomere spindly elongate, truncate, centrally bulged, longer than combined length of palpomeres 2-3, each with a dorsal concavity. Labial palpus 3-segmented; apical palpomere sub- cylindrical and glabrous. Pronotum: Moderately transverse to su- bquadrate, widest at middle, slightly nar- rowing posteriorly, weakly convex. Surface densely punctate, punctures evenly distrib- uted, each with a short seta; anterior margin weakly emarginate. Midline very shallowly depressed and always covered with less stout and somewhat shorter setae than lat- eral. Discal sides of pronotum irregularly convex at middle, each with a varied pit (Fig. 12). Laterally, marginal serration poorly developed; both anterior and poste- rior angles acute, moderately protuberant (Fig. 12). Scutellum wider than long, api- cally rounded, moderate in size. Labrum. 8, Clypeolontha siamensis, dorsal view. 9, C. alboplagiata, dorsal view. 10, C. sia- Elytron: Randomly distributed with zero to seven small dark tubercles, sometimes only vestigial. Surface with 5 parallel cos- tae including epipleural and sutural margin. Costae 2 and 3 fused on apical knobs. Costa 4 very feebly developed and sometimes hardly visible. Intervals slightly impressed and with setiferous punctures throughout. Humeral knobs moderately swollen. Epi- pleuron with aligned row of entire setae, broadest at middle. Lateral and apical mar- gin membranous. Thoracic sternites: Surface hairy with except metepisternum and metepimeron densely covered with whitish-yellow scales with slightly iridescent tinge (with illumi- nation and magnification). Mesosternum transverse, surface of disc and mesometas- ternal suture depressed. Metasternum large subquadrate, weakly depressed along mid- dle groove. Legs: Protibia tridentate, tooth color dis- tinctly darker than disc; basal tooth some- what vestigial; anterior spur movable, un- VOLUME 101, NUMBER 2 399 Figs. 12-15. Pronotum. 12, Clypeolontha siamensis. 13, C. alboplagiata. 14, C. bertiae. 15, C. laosensis. Scale line = 1.0 mm. 400 Set aim : —. Figs. 16-21. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON aie <8 “Ss Outlined hind femur. 16, Clypeolontha siamensis, male. 17, C. siamensis, female. 18, C. al- boplagiata, male. 19, C. alboplagiata, female. 20, C. bertiae, female. 21, C. laosensis, female. Scale line = 2.0 mm. dersized, subequal to one-fourth length of first protarsomere. Claws symmetrical, abruptly curved apically with a vertical subapical tooth at middle, one-half to two- thirds length of apical tooth. Protarsomere 1 shorter than combined length of protar- somerés? 2-33) PIR. = 3:029.721.621.6:2.7. Pro- and mesofemora transversely flattened, surface densely clothed with long, yellow- ish white pile. Hind femur stout (Fig. 16) with shorter pile than that on pro- and me- sofemora, about 2.5 times as wide as tibia, broadest at middle; MFL/W = 2.93-3.21. Tibiae and tarsi moderately clothed with pale white setae. Meso- and metatibia with two elongate, apical spurs, outer spur of metatibiae sharper and slightly longer than length of metatarsomere 1, inner spur curved apically. Meso- and metatarsi light yellowish brown with tarsomeres 1—5 sub- equal in length. Abdomen: Sternites moderately punc- tate; segments 1—6 continuous with wide maculation along lateral edges consisting of white scales with weakly iridescent tinge. Intervening surface shallowly setiferously punctate, with less stout and more sparsely distributed setae than those on dorsum; transversely sparsely intermixed with lon- ger setae. Sternites 2—5 weakly fused me- dially, not well defined. Sternite 5 with pos- terior margin moderately arc-shaped cen- trally sometimes exposing intersegmental membrane (Fig. 22). Pygidium triangular; apex distinctly prolonged, longitudinally convex, somewhat abruptly declivious on apical one-fourth when viewed laterally (Fig. 28); width approximately equal to length; PgW/H = 1.08—1.22; marginate lat- erally and becoming smooth apically. Dor- sal and ventral surface densely covered with tiny setae same as on dorsum, sparsely intermixed with longer setae which becom- ing denser on apical margin. Male genitalia: Parameres strongly asymmetrical with apices irregularly broad- VOLUME 101, NUMBER 2 EI eS S5 22 24 26 401 25 27 Outlined abdominal sternites 5—6 in ventral view. 22, Clypeolontha siamensis, male. 23, C. siamensis, female. 24, C. alboplagiata, male. 25, C. alboplagiata, female. 26, C. bertiae, female. 27, C. laosensis, female. S = sternite; P = pygidium. ened, contracted dorsomedially, and fused at base (Fig. 34); right paramere with a in- ward spine on basal fourth and not fused when viewed ventrally (Fig. 35). In lateral aspect parameres asymmetrically concave and sharpened forwardly (Figs. 36, 38). Basal piece and paramere approximately equal in length. Female.—BL = 19.7—20.7 mm.; BW = 9.3-10.0 mm; BL/BW = 2.04-2.17. Simi- lar to male except with stouter and more robust shape (Fig. 4); BW/HW = 1.76— 1.94, mean = 1.84; antennal club 6-seg- mented, compact and club length longer than basal segments 2—4, the first club seg- ment one-half to one-third length of rest, three basal segments each one-half as wide as first one; apical segment of maxillary palpus subequal to combined length of seg- ments 2—3; mean MPR = 1.9:1.4:3.5 (Fig. 402 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 42); protibia broader and more robust, an- terior tooth broadened; mean PTR: 3.0:1.2: 1.1:1.2:2.5; hind femur stouter and more rounded (Fig. 17); MFL/W = 2.34—2.56; the longer metatibial spur more or less stouter; abdomen dorsoventrally inflated; posterior edges of abdominal sternites 5 and 6 anteriorly with point-curved medially (Fig. 23); pygidium broadened basally with apex roundly inflated, slightly elongate and concave in lateral view (Fig. 29); PgW/H = 1.17-1.25. Distribution.—Montane areas of northern Thailand near the border of Burma (Myan- mar) (Fig. 46). Remarks.—Itoh (1995) misidentified several specimens of this species as C. al- boplagiata which were collected from the neighboring areas of the type locality, al- though he noted differences of the relative length of the first antennal club segment and the remainder in the female. Etymology.—The specific epithet is named for Siam, the former name of col- lecting place of this species. Clypeolontha alboplagiata (Brenske), new combination (Bigs.3 5.09: Ail ltss eee 9. 24~ 25-30, 31, 37,. 39, 40, 41,43) Melolontha alboplagiata Brenske 1898: 236 (nec Itoh, 1995: 202); Dalla Torre 1912: 267 (catalog); Sabatinelli 1993: 615 (catalog). Male diagnosis.—Male: Antennal club about 2.2 times length of stem; mean ASR =) 1.3:0.5:1.8; labrum shape triangular; pronotal midline not observed, angles ob- tuse; protarsomere | longer than combined length of protarsomeres 2 and 3 (PTR = S202 1225.14122>2-5)3 hind; femur broadly stout (MFL/W = 2.42-2.50); abdominal sternite 5 with posterior edge slightly arc- shaped centrally; apex of pygidium mod- erately prolonged; PgW/H = 1.00-—1.03; paramere strongly curved laterally at basal one-fourth, oblique ridge along right para- mere, apex turned inwardly and thinned apically. Description.—Male. Body: Oblong (Fig. 3), sides subparallel medially Dorsal sur- face densely covered with short yellowish- brown to whitish-yellow setae, all similar in length. BL = 16.8—-17.0 mm; BW = 7.8— 8.0 mm; BL/BW = 2.13-—2.15. Basal color reddish brown. Head: Surface densely to confluently punctate, punctures moderately large, each bearing a short seta. Antenna 10-segmented with 7-segmented clubs; lamellae slightly outwardly curved, about 2.2 times length of stem; first basal segment wider than third one; mean ASR = 1.3:0.5:1.8. Clypeus transverse, shallowly depressed anteriorly, weakly emarginated; sides gradually round- ing to biarcuate apex; middle of anterior margin inwardly depressed at tip (Fig. 9); clypeo-frontal suture moderately devel- oped; frons slightly narrowed making eyes large. Labrum moderately grooved with re- spect to clypeus; transversally symmetrical, strongly bilobed and each triangular, one- third width of clypeal base, depressed lat- erally (Fig. 11), visible dorsally, moderately protuberant. Maxillary palpus light yellow- ish brown, 4-segmented; apical palpomere spindly elongate, truncate, subequal in length to palpomeres 2—3, each with a dor- sal concavity. Labial palpus 3-segmented; apical palpomere subcylindrical and gla- brous. Pronotum: Moderately transverse, wid- est at middle, slightly narrowing posterior- ly, weakly convex. Surface densely punc- tate, punctures evenly distributed, each with a tiny seta; anterior margin weakly emar- ginate. Midline not visible. Discal sides of pronotum irregularly convex at middle, each with a vestigial concavity (Fig. 13). Laterally, marginal serration poorly devel- oped with feeble emargination, anterior and posterior angles obtuse, less protuberant (Fig. 13). Scutellum wider than long, api- cally rounded, moderate in size. Elytron: Surface with 5 parallel costae including along epipleural margin and su- VOLUME 101, NUMBER 2 tural margin. Costae 2 and 3 fused on apical knobs of elytron. Costae 4 very feebly de- veloped and sometimes hardly visible. In- tervals slightly impressed and with setifer- ous punctures throughout. Humeral knobs moderately swollen. Epipleuron with aligned row of entire setae, broadest at mid- dle. Lateral and apical margins membra- nous. Thoracic sternites: Surface hairy except metepisternum and metepimeron densely covered with whitish-yellow, slightly iri- descent scales (with illumination and mag- nification). Mesosternum transverse, sur- face of disc and mesometasternal suture de- pressed. Metasternum large, subquadrate, weakly depressed along middle groove. Legs: Protibia tridentate, tooth color dis- tinctly darker than disc; basal tooth some- what vestigial; anterior spur movable, un- dersized, subequal to one-fourth length of first protarsomere. Claws symmetrical, abruptly curved apically with a vertical subapical tooth at middle, one-half to two- thirds length of apical tooth. Protarsomere 1 longer than combined length of protar- someres 2 and 3; mean PTR = 3.0:1.2:1.1: 1.2:2.5. Femora of front and middle legs transversely flattened, surface densely clothed with long, yellowish-white pile. Hind femur more stout broadly (Fig. 18) with shorter pile than that on pro- and me- sofemora; about 2.5 times as wide as tibia, broadest at middle; MFL/W = 2.42-2.50. Tibiae and tarsi moderately clothed with pale white setae. Meso- and metatibiae with two elongate, apical spurs, outer spur of metatibia sharper and slightly longer than length of metatarsomere 1, inner spur curved apically. Meso- and metatarsi light yellowish brown with tarsomeres 1—5 sub- equal in length. Abdomen: Sternites moderately punc- tate; segments 1—6 continuous with wide maculation along lateral edges consisting of white scales with weakly iridescent tinge. Intervening surface shallowly setiferously punctate, with less stout and more sparsely distributed setae than those on dorsum; 403 transversely sparsely intermixed with lon- ger setae. Sternites 2-5 weakly fused me- dially, not well defined. Sternite 5 with pos- terior edge deeply, angularly emarginate (Fig. 24). Pygidium triangular; apex mod- erately prolonged and somewhat weakly de- pressed on apical one-third when viewed laterally (Fig. 30); width subequal to length; PgW/H = 1.00-—1.03; marginate iaterally and becoming smooth apically. Surface densely covered with tiny setae as on dor- sum, sparsely intermixed with longer setae and becoming denser on apical margin. Male genitalia: Paramere moderately asymmetrical, strongly curved laterally at basal one-fourth and fused at base; oblique ridge along right paramere to fused base (Fig. 40). Apex turned inwardly and thinned apically when viewed ventrally (Fig. 41). In lateral aspect, paramere at bas- al one-third concave and sharpened anteri- orly (Figs. 37, 39). Basal piece and para- mere approximately equal in length. Redescription of female.—Body and legs dark brown to reddish brown; clothed with pale white setae (Fig. 5). BL = 19.8—20.1 mm.; BW = 8.8-9.0 mm; BL/BW = 2.23-— 2.25. Sexual dimorphism in stouter shape with dorsoventrally convex; BW/HW = 1.62—1.66, mean = 1.64; surface clothed with stouter pale white setae; antennal club 6-segmented, compact length subequal to basal segments 2—4, first club segment one- fourth length of rest, third basal segments each half as wide as first; apical segment of maxillary palpus longer than the combined length of segments 2—3; mean MPR = 1.6: 1.3:3.7 (Fig. 43); protibia broader and more robust; anterior tooth longer than in male and broadened; mean PTR = 3.0:1.1:1.1: 1.2:2.8; anterior and posterior angle of pronotum obtuse, less protuberant; hind fe- mur stouter and roundly inflated (Fig. 19), MFL/W = 2.08-2.18; longer metatibial spur somewhat stouter than in male; abdo- men dorsoventrally inflated; posterior edge of abdominal sternite 5 smooth, sternite 6 shallowly biarcuate (Fig. 25); pygidium broadened basally with apex somewhat 404 sharpened; slightly elongate and concave in lateral view (Fig. 31); PgW/H = 1.11-—1.14. Material examined.—Holotype @ at ZMHB labeled separately as follows: ““Type(P)//India, Sikkim, ex coll. Fruhstor- fer(P)//coll. Brenske(P)//Melolontha albo- plagiata, type, Brsk, (H)//Zool. Mus Ber- lin(P). 2 d and 1 2 are placed in ISNB labeled as: Sikkim, Kurseong, R. P. Bretau- deau, 1894(P)//ex Museo Oberthur(P)(1 ¢). British Bootang, L. Durel, 1899(P)//ex Mu- seo Oberthur(P)(1 ¢ and 1 ¢). 2 6 and 1 2? in BMNH labeled as: Himalaya(H)// Bowring. 63 47*(P)//Melolontha(S. G. Schonherria Burm. n. sp.)(H)(1_ d). Alla- hauad(W)?//Bowring. 63 47*(P)//. Deter- mined from description. G. J. A.(P) Melo- lontha alboplagiata Brsk(W)(1 @). Atkin- son Coll. 92-3.(P)(1 ¢). Distribution.—Eastern Himalayan areas, including Sikkim and Bhutan (new record) (Fig. 46). Remarks.—In his original description, Brenske (1898) placed that C. alboplagiata with the typical Melolontha species and dis- tinguished them from M. albidiventris Fair- maire, M. cochinchinae Brenske, M. rubi- ginosa Fairmaire, and M. costata Nonfried by differences of the clypeus, pygidium, and first antennal club segment of the fe- males and therefore considered them as an unique group. We agree that C. alboplagia- ta should be recognized as an distinct group but we reject the concept of so-called typ- ical Melolontha applied at that time because it does not satisfy today’s systematic re- quirement and may cause more taxonomic uncertainty. However, after careful compar- ison of the types and other material, the sys- tematic position of the above-mentioned species by Brenske (1898) within Melolon- tha suggest further revisionary work. Clypeolontha bertiae Li and Yang, new species (Figs. 7, 14, 20, 26, 32, 44) Type series.—Holotype @ (deposited in NMNH) with label data in handwritting as follows: Pukhieo, Chiaya poon, NE Thai- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON land, 5-V-1986, P, EK-Amnuay. 4 @ para- types: one with the same data as holotype deposited in MNHN;; one in ISNB, with la- bel data as follows: Laos, Luang Prabang: Ban Na Gnan. 20. V. 1920. R. V. de Sal- vaza(P). Specimen (in ISNB) condition: right protarsus, middle and hind legs miss- ing; anterior two segments of left protarsus missing. The remaining two paratypes is in BMNH with identical printed label data as follows: At light//N.E. Thailand: 800m., Phu Khieo Wildlife Sanctuary, 16°30'N, 101°46’E//Chaiyaphum Province, Khon San. 13-15. V.1988//Evergreen rain forest. M. J. D. Brendell. B. M. 1988-183. Description.—Female. Body: Oblong (Fig. 7), dorsoventrally convex and sides subparallel medially. BL = 17.9-18.5 mm; BW = 7.8-7.9 mm; BL/BW = 2.28-2.35. Head: Deeply rufous brown to lightly red- dish brown. Surface distinctly punctate, punctures large, each bearing a tiny, pale white to yellowish-brown seta, brighter on sides of eyes. Antenna 10-segmented with 6-segmented club; first club segment half to two-thirds length of rest; first basal segment subequal in length to third. Clypeus trans- verse, very shallowly depressed anterior to marginated edge; sides gradually rounding to biarcuate apex; middle of anterior margin inwardly depressed at tip; clypeo-frontal su- ture moderately developed. Labrum mod- erately grooved with respect to clypeus; transversally symmetrical, strongly bilobed, moderately declivous laterally, one-third width of clypeal base, visible dorsally, moderately protuberant. Maxillary palpus yellowish-brown; 4-segmented; apical pal- pomere spindly elongate, truncate, equal to combined length of segments 2—3 (Fig. 44); mean MPR = 2.0:1.5:3.5. Labial palpus 3- segmented; apical palpomere subcylindrical and glabrous. Pronotum: Rufous brown. Moderately transverse, widest at middle, slightly nar- rowing posteriorly, disc moderately convex. Surface densely finely punctate, punctures evenly distributed, each with a pale white to yellow seta; anterior margin weakly VOLUME 101, NUMBER 2 emarginate. Midline not visible. Discal sides of pronotum irregularly convex at middle, each with a shallow concavity (Fig. 14). Laterally, marginal serration poorly de- veloped with feeble emargination; anterior and posterior angle weakly protuberant (Fig. 14). Scutellum wider than long, api- cally rounded, moderate in size. Elytron: Brightly rufous yellow to ru- fous brown. Mean BW/HW = 1.73. Surface clothed with pale white to yellow setae; bearing 5 parallel costae on each elytron in- cluding along epipleural margin and sutural margins. Costae 2 and 3 fused on the apical knobs of elytron. Costae 4 very feebly de- veloped and hardly visible. Intervals slight- ly impressed and with setiferous punctures throughout. Humeral knobs moderately swollen. Epipleuron with aligned row of smaller entire setae, broadest at middle. Lateral and apical margin membranous. Thoracic sternites: Surface hairy except metepisternum and metepimeron densely covered with whitish-yellow scales with slight iridescent tinge (with illumination and magnification). Mesosternum trans- verse, surface of disc and mesometasternal suture depressed. Metasternum large, su- bquadrate, weakly depressed along middle groove. Legs: Protibia tridentate, tooth color darker than disc; basal tooth somewhat ob- solete; anterior spur movable, undersized, subequal to one-fourth the length of first protarsomere. Claws symmetrical, abruptly curved apically with a vertical subapical tooth at middle, two-thirds length of apical tooth. Pretarsus reddish brown; broadened laterally; protarsomere | longer than com- bined length of protarsomeres 2 and 3; PTR = 3.3:1.4:1.2:1.4:2.2 (n = 4). Pro- and me- sofemora transversely flattened, surface densely clothed with long, yellowish-white pile. Meso- and metatibiae and tarsi lightly yellowish brown; metatarsal spurs reddish brown. Hind femur roundly inflated (Fig. 20) with shorter pile than those on pro- and mesofemora; MFL/W = 2.34-—2.45. Abdomen: Dorsoventrally inflated. Ster- 405 nites transversely rugose, moderately punc- tate; segments 1—6 continuous with wide maculation along lateral edges consisting of white scales with weakly iridescent tinge. Intervening surface shallowly setiferously punctate with setae same as on dorsum, transversely intermixed with very sparse longer setae. Sternites 2—5 weakly fused medially, not well defined. Posterior edges of sternites 5 and 6 smooth, sternite 6 with feeble irregular serration (Fig. 26). Pygidi- um deeply reddish to lightly rufous brown; densely clothed with pale white to yellow setae sparsely intermixed with longer setae; triangular; in lateral view apex moderately elongate, smoothly declivous apically with apex sharpened (Fig. 32); marginate later- ally then decreased apically. PgW/H = 1.08—1.12. Apex dorsoventrally with a ter- minal tuft of denser setae. Distribution—Some 200 km north of Vientiane, Laos, and the low montane area of northeastern Thailand (Fig. 46). Remarks.—We describe Clypeolontha bertiae from only five females because those diagnostic characters found in the fe- male types of C. siamensis, referring to both C. siamensis and C. alboplgiata, are useful and stable enough to separate inter- specific females. Those distinguishing char- acters are given in the key. Etymology.—Named for Dr. Nicole Berti who helped the senior author at Muséum national d’ Histoire naturelle, Paris, in 1997. Clypeolontha laosensis Li and Yang, new species (Bigs. 6, 155,21, 273-33; 49) Type.—Holotype ¢ (deposited in ISNB) labeled as follows: Laos(P), Nam Tien(H), le(P), 14-IV(H), 191(P)8(H), R. Vitalis de Salvaza(P). Description.—Female. Body: Oblong (Fig. 6), dorsoventrally convex and sides subparallel medially. BL = 17.2 mm; BW = 7.5 mm; BL/BW = 2.29. Basal color dark rufous brown. Head: Surface distinctly punctate, punctures large, each bearing a tiny, yellowish-brown seta, brighter on 406 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 29 31 33 Figs. 28-33. Pygidium, right lateral view. 28, Clypeolontha siamensis, male. 29, C. siamensis, female. 30, C. alboplagiata, male. 31, C. alboplagiata, female. 32, C. bertiae, female. 33, C. laosensis, female. Scale line = 1.0 mm. VOLUME 101, NUMBER 2 407 ee. Male genitalia. 34, 35, 36, 37, Clypeolontha siamensis. 38, 39, 40, 41, C. alboplagiata. 34, Figs. 34-41. 40, Dorsal view. 36-37, Right lateral view. 38-39, Left lateral view. 35, 41, Ventral view. Scale line = 1.0 mm. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 42-45. Female outlined maxillary palpomeres 2-4. 42, Clypeolontha siamensis. 43, C. alboplagiata. 44, C. bertiae. 45, C. laosensis. Scale line = 0.5 mm. 408 VOLUME 101, NUMBER 2 (] Ly = -* Fig. 46. C. bertiae; star = C. laosensis. sides of eyes. Antenna 10-segmented with 6-segmented club; first club segment half length of rest; first basal segment subequal in length to third. Clypeus transverse, mod- erately depressed anterior to marginated edge; sides gradually rounding to biarcuate apex; middle of anterior margin inwardly depressed at tip; clypeo-frontal suture mod- erately developed. Labrum moderately grooved with respect to clypeus; transver- sally symmetrical, strongly bilobed, mod- erately declivous laterally, one-third width of clypeal base, visible dorsally, moderately protuberant. Maxillary palpus dark yellow- ish brown; 4-segmented; apical palpomere spindly stout, truncate, shorter than com- bined length of segments 2—3 (Fig. 45); MPR = 1.8:1.3:2.7. Labial palpus 3-seg- mented; apical palpomere subcylindrical and glabrous. Pronotum: Moderately transverse, wid- @ THAILAND fan. we Distribution. Solid square = Clypeolontha siamensis; solid circle = C. alboplagiata; open circle = est at middle, slightly narrowing posterior- ly, weakly convex. Surface densely punc- tate, punctures more or less irregularly dis- tributed, each with a pale white seta; ante- rior margin weakly emarginate. Midline very shallowly depressed, clothed with very tiny, brownish setae. Discal sides of pro- notum irregularly convex at middle, each with a vestigial concavity (Fig. 15). Later- ally, marginal serration poorly developed with feeble emargination; anterior angle acute, protuberant; posterior angle obstuse (Fig. 15). Scutellum wider than long, api- cally rounded, moderate in size. Elytron: BW/HW = 1.91. Surface clothed with pale white setae, tending yel- low basally; bearing 5 parallel costae on each elytron including along epipleural margin and sutural margin. Costae 2 and 3 fused on the apical knobs of elytron. Costae 4 very feebly developed and hardly visible. 410 Intervals slightly impressed and with seti- ferous punctures throughout. Humeral knobs moderately swollen. Epipleuron with aligned row of smaller, brownish entire se- tae, broadest at middle. Lateral and apical margins membranous. Thoracic sternites: Surface hairy except metepisternum and metepimeron densely covered with whitish-yellow scales with slight iridescent tinge (with illumination and magnification). Mesosternum transverse, sur- face of disc and mesometasternal suture de- pressed. Metasternum large, subquadrate, weakly depressed along middle groove. Legs: Protibia tridentate, tooth color dis- tinctly darker than disc; basal tooth somewhat obsolete; anterior spur movable, undersized, subequal to one-third length of first protar- somere. Claws symmetrical, abruptly curved apically with a vertical subapical tooth at middle, two-thirds length of apical tooth. Pre- tarsus dark yellowish brown; broadened lat- erally; protarsomere | longer than combined length of protarsomeres 2 and 3; PTR = 3.0: 1.4:1.2:1.2:2.3. Femora dark rufous brown. Pro- and mesofemora transversely flattened, surface densely clothed with long, yellowish- white pile. Hind femur slightly transversely inflated (Fig. 21) with shorter pile than those on pro- and mesofemora; MFL/W = 2.61. Tibiae and tarsi moderately clothed with pale white setae. Meso- and metatibia with 2 elon- gate, apical spurs, outer spur of metatibiae sharper and slightly longer than length of me- tatarsomere 1, inner spur curved apically. Meso- and metatarsi dark yellowish brown with tarsomeres 1—5 subequal in length. Abdomen: Dorsoventrally inflated. Ster- nites transversely rugose, moderately punc- tate; segments 1—6 continuous with wide maculation along lateral edges consisting of white scales with weakly iridescent tinge. Intervening surface shallowly setiferous punctate with whitish-yellow setae; trans- versely intermixed with very sparsely lon- ger setae. Sternites 2—5 weakly fused me- dially, not well defined. Posterior edge of sternite 5 with distinct point-curve, sternite 6 deeply, broadly emarginate (Fig. 27). Py- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gidium reddish brown; wider than long; tri- angular; densely clothed with yellowish- brown setae sparsely intermixed with lon- ger setae; in lateral view, apex slightly elon- gate, smoothly declivous apically with apex rounded (Fig. 33); marginate laterally and smooth apically. PEW/H = 1.18. Apex dor- soventrally with a terminal tuft of denser setae. Distribution.—Northern Laos, roughly 19°34'N, 103°42’E (Fig. 46). Remarks.—Clypeolontha laosensis is de- scribed from a single female. Geographi- cally and morphologically, it is close to C. bertiae, but it may be separated from it by the diagnostic characters in the key. Etymology.—tThe specific epithet is from the country of collection. ACKNOWLEDGMENTS We greatly appreciate the help of Keith Philips (The Ohio State University, Colum- bus) and Kerry Katovich (University of Wisconsin, Madison) for reading and com- menting on a primary version of this man- uscript. We also thank the following cura- tors or researcher of the museum or insti- tution for kindly loaning invaluable mate- rial which made this study possible: Manfred Uhlig, Hella Wendt, and Joachim Schulze (ZMHB); Konjev Desender and Marcel Cludts (ISNB); Malcolm Kerley (BMNH). Special thanks are also due to Takeshi Itoh (Osaka) for his kind assis- tance. Finally, we are especially grateful to Robert Alexis (Belgium) and his wife for their warm assistance during the senior au- thor’s visits to Europe in 1997. This work was supported partly by grant NSC 87- 2313-13-002-054 from the National Sci- ence Council of the Republic of China. LITERATURE CITED Arnett, R. H., G. A. Samuelson, and G. M. Nishida. 1993. The Insect and Spider Collections of the World, 2nd ed. Flora and Fauna Handbook 11, 310 pp. Gainesville, FL. Arrow, G. J. 1913. Notes on the Lamellicorn Coleop- tera of Japan and descriptions of a few new spe- VOLUME 101, NUMBER 2 cies. Annals and Magazine of Natural History 8(12): 394—408. Baraud, J. 1992. Faune de France 78. Coléoptéres Scarabaeoidea d’Europe. 1x + 856 pp. Lyon. Brenske, E. 1898. Einige neue Melolonthiden. Stetti- ner Entomologische Zeitung 59: 236-248. Burmeister, H. 1855. Handbuch der Entomologie Bde. IV, 2, Coleoptera Lamellicornia Phyllophaga Chaenochela. x + 570 pp. Berlin. Dalla Torre, K. W. von. 1912. Scarabaeidae: Melolon- thinae III, pp. 135—290. In W. Junk & S. Schenk- ling, eds. Coleopterorum Catalogus Pars 49, Vol- ume 20. Itoh, T. 1995. Melolonthidae (Coleoptera) from Thai- land, II. Elytra 23(2): 195-204. Medvedev, S. I. 1951. Fauna SSSR, Coleoptera. Vol. 411 X, 1, Scarabaeidae Melolonthinae. 375 pp. Mos- cow. Nomura, S. 1952. Uber die Melolontha-Arten aus Ja- pan und dessen Umgebung, mit den Beschreibun- gen einen neuen Gattung und einiger neuer Arten. T6h6 Gakuho 2: 24-34. Reitter, E. 1902. Bestimmungs-Tabelle thidae aus der europdischen fauna und den an- grenzenden Landern. III: Pachydemini, Sericini und Melolonthini. Verhandlungen des Naturfor- schenden Vereines in Briinn 50: 93-303. Sabatinelli, G. 1993. Catalogue of the Scarabaeoidea Melolonthidae from Himalaya and North India: Macrodactylini, Hoplini, Diplotaxini, Melolonthi- ni and Sericini Coleoptera. Memorie della Societa Entomologica Italiana, Genova 71(IL): 593-636. der Melolon- PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 412-413 DASYMUTILLA TOMBERLINI, A NEW SPECIES OF VELVET ANT (HYMENOPTERA: MUTILLIDAE) FROM NEW MEXICO DONALD G. MANLEY Department of Entomology, Clemson University, Pee Dee Research and Education Cen- ter, 2200 Pocket Road, Florence, SC 29506-9706, U.S.A. (e-mail: dmanley @ clemson.edu) Abstract.—Dasymutilla tomberlini is a new species of velvet ant (Hymenoptera: Mu- tillidae) collected in extreme southwestern New Mexico. It is distinguished by the presence of only yellow pubescence, by the complete absence of black pubescence, and by the smooth anterior surface of the first gastral tergite. Eleven specimens were examined, all collected in pitfall traps. Key Words: In March of 1996, specimens of velvet ants (Hymenoptera: Mutillidae) from the Carnegie Museum of Natural History col- lection were sent to me for identification by C. W. Young. Among those specimens was a series of 11 females of Dasymutilla Ash- mead from extreme southwestern New Mexico that was determined to be a new species. A diagnosis and description follow. Dasymutilla tomberlini Manley, new species Diagnosis.—This species has the integ- ument reddish throughout and is densely clothed over most of the body with yellow- ish pubescence. All pubescence is of this color. A scutellar scale is present and the antennal scrobes are carinate. There is no genal carina and the genae are relatively smooth and shining. The anterior face of the first abdominal tergite is smooth and shining, devoid of punctation or pubes- cence. The pygidium is finely rugose, al- most granulate. Description.—Female: Length, 10.6— 15.0 mm. Head reddish, densely clothed with yellowish recumbent pubescence; Dasymutilla tomberlini, new species, Mutillidae, Hymenoptera some long, erect yellow pubescence on ver- tex; mandible acute at tip, with an incon- spicuous inner tooth about one-third dis- tance from tip; clypeus evenly convex on anterior margin, but concealed by yellowish pubescence; scape weakly carinate, smooth and shining, clothed with yellowish pubes- cence; first flagellomere long, about length of second and third united, remaining fla- gellomeres subequal in length; antennal scrobes distinctly carinate; front and vertex coarsely punctate, but with dense yellow pubescence concealing sculpture; gena smooth and shining, with shallow, well-sep- arated punctures, lacking a genal carina, and concealed with yellow pubescence; head width 2.1—2.6 mm; relative width of head to thorax 0.75. Thorax reddish, densely clothed with erect and recumbent yellow pubescence; dorsum of thorax longer than broad (3.2— 4.3 mm long X 2.7—3.5 mm wide); scutellar scale present and conspicuous, as well as a transverse, sinuate carina immediately an- terior to scutellar scale; cephalic margin of pronotum evenly rounded, not emarginate medially; entire thorax densely covered VOLUME 101, NUMBER 2 with yellow pubescence, obscuring punc- tation. Abdomen reddish, densely clothed with erect and recumbent yellow pubescence, except anterior face of first gastral tergite smooth and shining, devoid of pubescence except for apical fringe; pygidium devoid of pubescence, and disk of second sternite with only a few scattered hairs; disk of sec- ond tergite smooth and shining, with sparse, shallow punctures; remainder of sculpture concealed by dense yellow pubescence; py- gidium finely rugose, almost granulate; first sternite with a distinct carina about one- third its length, elevated distally to form a tooth. Legs reddish, smooth, and shining, con- spicuously clothed with yellow pubescence. Most type specimens (7) longer than 13.0 mm. Thorax in each specimen longer than broad, with width:length ratio generally about 0.8:1.0. Head width distinctly nar- rower than thoracic width. Male: Unknown. Holotype.—?, Antelope Pass, Peloncillo Mountains, Hidalgo County, New Mexico, July 1990 (Barney Tomberlin), collected by pitfall trap, deposited in the collection of the Carnegie Museum of Natural History. Paratypes.—10 2, same data as holo- type. Paratypes deposited in the Carnegie Museum of Natural History (7), and with the author (3). Etymology.—Named after Barney Tom- berlin who collected the series of speci- mens. Discussion.—This species superficially resembles Dasymutilla magna (Cresson). However, under microscopic examination, it is easily distinguished from D. magna. The latter has the genae as coarsely punc- tate as the front and vertex and possesses a genal carina. In D. tomberlini, although ob- scured by the dense pubescence, the genae are relatively smooth and shining, much less coarsely punctate than the front and vertex, and a genal carina is lacking. In Mickel’s (1928) key to Dasymutilla, this species keys to couplet 81, which includes 413 D. satanas Mickel and D. sackenii (Cres- son). In Mickel’s more recent (1936) key to Dasymutilla, it keys to couplet 87, which includes the same two species. This species can be distinguished from ali of the above- mentioned species by the tota! lack of black pubescence and by the smooth and shining anterior margin of the first gastral tergite. This species is known only from the type series, collected in pitfall traps designed to collect reptiles, near the Arizona border in extreme southwestern New Mexico. In spite of extensive personal collecting and exam- ination of numerous collections including specimens collected from the same general locale, no other specimens of this species have been encountered. Due to the collec- tion method, only females were collected. None of the species of Dasymutilla listed in Krombein (1979) for which only the male is known seem to be likely candidates as the male of this species. Although D. candida Mickel is somewhat similar in col- or and geographic range (being found in southeastern Arizona), it is much smaller (length 8—9 mm) than D. tomberlini (10—15 mm). It seems more likely that D. candida represents the unknown male of D. thetis (Blake) than the present species. ACKNOWLEDGMENTS I thank C. W. Young and the Carnegie Museum of Natural History, Pittsburgh, Pa., for loan of the specimens for study. This is Technical Contribution No. 4202 of the South Carolina Agricultural Experiment Station, Clemson University. LITERATURE CITED Krombein, K. V. 1979. Family Mutillidae, p. 1276— 1314. In Krombein, K.. V., PD: Hurd, Jr; D..R. Smith, and B. D. Burks, eds. Catalog of Hyme- noptera In America North of Mexico, Volume 2. Smithsonian Institution Press, Washington, D.C. Mickel, C. E. 1928. Biological and taxonomic inves- tigations on the mutillid wasps. United States Na- tional Museum Bulletin 143: 1-351. . 1936. New species and records of Nearctic mutillid wasps of the genus Dasymutilla (Hyme- noptera). Annals of the Entomological Society of America 29: 29-60. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 414-427 LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF TRUPANEA WHEELERI CURRAN (DIPTERA: TEPHRITIDAE) ON ASTERACEAE IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN AND JEFFREY A. TEERINK Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: rgoeden @ucracl.ucr.edu) Abstract.—Trupanea wheeleri Curran is a multivoltine, florivorous fruit fly (Diptera: Tephritidae) infesting flower heads of a wide variety of Asteraceae in California and the western half of the United States. Four new host-plant genera and five new species records are reported. To date, 7. Wheeleri is known from seven tribes, 15 subtribes, 28 genera, and 47 species of hosts. The egg, first through third instar larvae, and puparium are described and figured for the first time. The egg pedicel has one or two rows of aeropyles. The interspiracular processes of the first instar are large, broad, and multibranched. The anterior thoracic spiracles of the second instar each bear 7—8 papillae, more than any previously studied congeneric species. The lateral spiracular complexes of the third instar are identical to that of 7. imperfecta (Coquillett), which is the first report of two species of Trupanea that share the same type and number of sensilla in their metathoracic and abdominal, lateral spiracular complexes. The life cycle of 7. wheeleri in southern Cali- fornia is of the aggregative type. The eggs are inserted alongside or into the corollas of florets and ovules upon which the first instars feed in closed, preblossom flower heads. Second instars feed mainly on ovules and florets of preblossom flower heads and soft achenes of open flower heads; whereas, third instars feed on soft achenes in open and postblossom flower heads. Pupariation occurs inside the mature flower heads, from which the adults emerge about the time that the achenes are shed. Several generations are pro- duced on a variety of hosts during the spring, summer, and fall, and overwintering is as long-lived adults. Seven species of chalcidoid Hymenoptera were reared from individual puparia and mature flower heads bearing puparia of 7. wheeleri as solitary, primary, larval-pupal endoparasitoids: Eurytoma n. sp.? (Eurytomidae), Eurytoma obtusiventris Ga- han (Eurytomidae), Eurytoma veronia Bugbee (Eurytomidae), Eupelmus sp. (Eupelmi- dae), Mesopolobus sp. (Pteromalidae), Pteromalus sp. (Pteromalidae), Torymus sp. (To- rymidae). Key Words: Insecta, Trupanea, Asteraceae, nonfrugivorous Tephritidae, biology, tax- onomy of immature stages and adults, flower-head feeding, host-plant range, parasitoids This is the penultimate paper in our re- portance (Foote and Blanc 1963, Foote et cent series on Trupanea, one of the larger al. 1993). Trupanea remained little known and more widespread genera of nonfrugi- (Foote 1960, Foote et al. 1993) until we vorous fruit flies in North America and Cal- published detailed life histories of ten spe- ifornia, though of little or no economic im- cies from southern California (Cavender VOLUME 101, NUMBER 2 and Goeden 1982; Goeden 1987, 1988; Goeden and Teerink 1997b, 1998, 1999; Goeden et al. 1998a, b; Headrick and Goe- den 1991; Knio et al. 1996b), along with descriptions of their immature stages (Cav- ender and Goeden 1982; Goeden and Teer- ink 1997b, 1998, 1999; Goeden et al. 1998a, b; Headrick and Goeden 1991; Knio et al. 1996a; Teerink and Goeden 1998, 1999). In this paper we describe the life his- tory and immature stages of an eleventh species, 7. wheeleri Curran. MATERIALS AND METHODS This study was based in large part on dis- sections of subsamples of flower heads of Asteraceae infested by JT. wheeleri from samples collected since 1990 in southern California in the manner described by Goe- den (1985, 1992). One-liter samples of ex- cised, immature and mature flower heads containing eggs, larvae, and puparia were transported in cold-chests in an air-condi- tioned vehicle to the laboratory and stored under refrigeration for subsequent dissec- tion, photography, description, and mea- surement. Twenty-two eggs, 15 first-, 10 second-, and 12 third-instar larvae, and sev- en puparia dissected from flower heads were preserved in 70% EtOH for scanning electron microscopy (SEM). Additional pu- paria were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity chambers at room temper- ature for adult and parasitoid emergence. Specimens for SEM were hydrated to dis- tilled water in a decreasing series of acid- ulated EtOH. They were osmicated for 24 h, dehydrated through an increasing series of acidulated EtOH and two, 1-h immer- sions in Hexamethlydisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, and studied with a Philips XL30-FEG scanning electron mi- croscope in the Institute of Geophysics and Planetary Physics, University of California, Riverside. Most adults reared from isolated puparia were individually caged in 850-ml, clear- 415 plastic, screened-top cages with a cotton wick and basal water reservoir and provi- sioned with a strip of paper toweling im- pregnated with yeast hydrolyzate and su- crose. These cages were used for longevity studies in the insectary of the Department of Entomology, University of California, Riverside, at 25 + 1°C, and 14/10 (L/D) photoperiod. Virgin male and female flies obtained from emergence vials also were paired (n = 3) in clear-plastic petri dishes provisioned with a flattened, water-moist- ened pad of absorbant cotton spotted with honey (Headrick and Goeden 1991, 1994) for observations of their courtship and cop- ulation behavior. Plant names used in this paper follow Hickman (1993) and Bremer (1994); tephritid names and adult terminology follow Foote et al. (1993). Terminology and telegraphic format used to describe the immature stages follow Goeden and Teerink (1997a, b; 1998, 1999), Goeden et al. (1998a, b), Headrick and Goeden (1991), Knio et al. (1996a), Teerink and Goeden (1998, 1999), and our earlier works cited therein. Means + SE are used throughout this paper. Voucher specimens of 7. wheeleri and its parasitoids reside in the research collections of RDG; preserved specimens of eggs, lar- vae and puparia are stored in a separate col- lection of immature Tephritidae acquired by JAT and now maintained by RDG. RESULTS AND DISCUSSION Taxonomy Adult.—Trupanea wheeleri was first de- scribed by Curran (1932) from a female ho- lotype from San Diego Co., California, as Trypanea wheeleri. Trypanea is a misspell- ing of Trupanea. Curran (1932), Foote (1960), Foote and Blanc (1963), and Foote et al. (1993) pictured the wing pattern of the female and male, which, unlike those of several North American 7rupanea spp., 1s not overtly sexually dimorphic (Foot et al. 1993). However, sex-related, wing-pattern variations were found by RDG in T. whee- 416 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Incidences (%) of absence/complete-/incompleteness of proximal and distal rays in cell dm in wings of 423 6 and 376 2 T. wheeleri reared from flower heads of native Asteraceae in southern California during 1989-1997. Left wing Right wing Proximal ray Distal ray Proximal ray Distal ray M: B: Cc: M B e M B G M B Cc Males 205 186 32 O 19] 232 201 183 39 ] 189 1333} (48.5) (44.0) (7.5) (0.0) (45.2) (54.8) (47:5) (43.3) (9.2) (0.2) (44.7) (55.1) Females 117 197 62 0) 87 289 114 200 62 0) 90 286 Gi) ©2224)" 16:5) lO) 3a), 6:9) G03)) 4632) cts) (OO) (@32) (7s) 4M = missing, B = broken, C = complete. leri. Foote et al. (1993, p. 446) cautioned that the “‘... identification of this species must be approached with care, as the prox- imal ray in cell dm varies from completely present to completely absent.’’ RDG quan- tified the variation in this key character in reared specimens of 7. wheeleri from Cal- ifornia in his research collection (Table 1). The data in Table | show that the proximal ray is much more prone to be missing than complete than is the distal ray in both wings of males as well as females. In nearly half of both wings examined in males, the prox- imal and distal rays were broken, a condi- tion which Foote et al. (1993, p. 446) de- scribed for the proximal ray as, ‘“‘... the two ends of this ray are visible as outcrop- pings in the dark area around vein r-m and anteriorly-directed extensions of the distal end of the elongate marking on vein CuA,.”” Among females, however, the prox- imal ray was twice as likely to be broken than the distal ray (Table 1), and unlike the proximal ray, this broken ray in both sexes usually appeared as a posteriorly-directed outcropping arising midway between cross- veins r-m and dm-cu along vein M. Foote et al. (1993) also noted that the holotype has the proximal ray complete in one wing, but almost completely missing in the other. Indeed, in 99 (23.4%) of the 423 males and 69 (18.4%) of the 376 females examined (Table 1), the proximal and distal rays of one wing differed as to whether they were missing, broken, and complete in the other wing of the same individuals. Finally, 32 (7.6%) of all males and 17 (4.5%) of all females lacked the elongated darkening along vein CuA, or had this infuscation re- duced, sometimes to a spot, as in some T. actinobola (Loew), T. jonesi Curran, T. tex- ana Malloch, and 7. vicina (Wulp) (Foote et al. 1993), yet all of these variants were reared along with 7. wheeleri from the same samples of flower heads of known hosts of T. wheeleri. Misidentifications of such variants may explain the limited rec- ords in Foote et al. (1993) for T. texana from southern California, where we have not yet reared this species. This variation may also help to explain difficulties expe- rienced in distinguishing some females of T. actinobola from T. wheeleri (Goeden et al. 1998b). In most instances, however, the infuscation along vein CuA, combined with the dark ray connecting the pterostigma with vein r-m that is narrower than the length of the pterostigma serve to distin- guish 7. wheeleri from all North American congeners (Foote et al. 1993). Immature stages.—The eggs, larvae, and puparium heretofore have not been de- scribed nor illustrated. Egg: Twenty-seven eggs of 7. wheeleri dissected from field-collected flower heads were white, opaque, smooth, elongate-ellip- VOLUME 101, NUMBER 2 Acc V Dy SSS 100kV 30 136x 170 Spot Magn 200 jim Ficsell: soidal, 0.63 + 0.003 (range, 0.58—0.64) mm long, 0.18 + 0.003 (range, 0.16—0.20) mm wide, smoothly rounded at tapered basal end (Fig. 1A); peg-like pedicel 0.02 mm long, with 1—2 rows of aeropyles (Fig. 1B). The egg of 7. wheeleri is similar to that of T. bisetosa in possessing one or two rows of aeropyles, all other species of Trupanea previously studied have only one row (Goe- den and Teerink 1997b, 1998, 1999; Goe- den et al. 1998a, b; Headrick and Goeden 1991; Knio et al. 1996a; Teerink and Goe- den 1998, 1999). The egg of 7. bisetosa is much longer, however, than that of 7. whee- leri (Knio et al. 1996a). The 7. wheeleri egg also is slightly larger than the egg of T. pseudovicina (Goeden and Teerink 1998a), wider than that of 7. arizonensis (Goeden and Teerink 1998b), but smaller than that of T. imperfecta (Teerink and Goeden 1998). First instar: White, elongate-cylindrical, rounded anteriorly and posteriorly (Fig. 2A), minute acanthae circumscribe inter- segmental lines; gnathocephalon smooth, lacking rugose pads (Fig. 2B); dorsal sen- sory organ a dome-shaped papilla (Fig. 2B- 1, 2C-1); anterior sensory lobe (Fig. 2B-2), bears terminal sensory organ (Fig. 2C-2), pit sensory organ (Fig. 2C-3), lateral sen- sory organ (Fig. 2C-4) and supralateral sen- sory organ (Fig. 2C-5); stomal sense organ reduced and ventrad of anterior sensory 417 AccV SpotMagn WD (— 20 um B 100kV 30 1791x171 Egg of Trupanea wheeleri: (A) habitus, anterior end to left: (B) pedicel, aeropyles. lobe (Fig. 2B-3); mouth hook bidentate (Fig. 2B-4, 2D-1); median oral lobe later- ally flattened (Fig. 2B-5, D-2); labial lobe with two pore sensilla (Fig. 2D-3); a pair of integumental petals dorsad of mouth hooks (Fig. 2B-6, 2D-4); pit sensillum lat- erad of mouth lumen (Fig. 2D-5); minute acanthae ventrad of mouth lumen; anterior thoracic spiracle absent; caudal segment with two stelex sensilla dorsad and ventrad of posterior spiracular plates; posterior spi- racular plate bears two ovoid rimae, Ca. 0.008 mm in length (Fig. 2E-1), and four interspiracular processes, each with 1-2, multi-dentate branches, longest measuring 0.007 mm (Fig. 2E-2); intermediate sensory complex with a stelex sensillum (Fig. 2F- 1) and a medusoid sensillum (Fig. 2F-2). The first instar is similar in general hab- itus and sensory structures to previously studied Trupanea species (Goeden and Teerink 1997b, 1998, 1999; Goeden et al. 1998a, b; Knio et al. 1996a; Teerink and Goeden 1998, 1999). However, the inter- spiracular processes are large, broad, and apically multi-branched, and thus are far more elaborate than those of, for example, T. arizonensis (Goeden and Teerink 1998) and 7. conjuncta (Teerink and Goeden 1998). Second instar: White, elongate-cylindri- cal, tapering anteriorly, rounded posteriorly (Fig. 3A), minute acanthae circumscribe in- 418 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON AccV Spot Magn WD /-——————-+4 200 jim AccV Spot Magn WD 5x 171 - 0 245 2 -_ \ oe z ¥ ; = g AccV Spot Magn | a AccV SpotMagn WD /————=104m 10.0 kV 3.0 6041x# 22. oi sti 100kV30_ 1976x_ 22 ¥ / : i) ‘ a4 f ai af fy rs / Acc.\V SpotMagn WD —- = 10.0kV 3.0 11923x 22.4 » Fig. 2. First instar of Trupanea wheeleri: (A) habitus, anterior end to left; (B) gnathocephalon, anterior view, 1—dorsal sensory organ, 2—anterior sensory lobe, 3—stomal sense organ, 4—mouth hook, 5—median oral lobe, 6—integumental petal; (C) anterior sensory lobe, |1—dorsal sensory organ, 2—terminal sensory organ, 3— pit sensory organ, 4—lateral sensory organ, 5—supralateral sensory organ; (D) gnathocephalon, ventral view, 1—mouth hook, 2—median oral lobe, 3—labial lobe pore sensilla, 4—integumental petal, 5—pit sensillum; (E) posterior spiracular plates, 1—rima, 2—interspiracular process; (F) intermediate sensory complex, 1—stelex sensillum, 2—medusoid sensillum. VOLUME 101, NUMBER 1) Acc.V Spot Magn : AccV SpotMagn WD 10.0 kV 3.0. 68x 3 10.0kV 30 857x278 Acc.V Spot Magn™8WD 10.0kV 3.0 1310x 19.9 Fig. 3. Second instar of Trupanea wheeleri: (A) habitus, anterior end to left: (B) gnathocephalon, anterior view, 1—serrated rugose pad, 2—dorsal sensory organ, 3 anterior sensory lobe, 4—stomal sense organ, 5— pit sensillum; (C) anterior sensory lobe, 1—serrated rugose pads, 2—dorsal sensory organ, 3—terminal sensory organ, 4—pit sensory organ, 5—lateral sensory organ, 6—supralateral sensory organ, 7—stomal sense organ; (D) gnathocephalon, anterior view, 1—mouth hook, 2—median oral lobe; (E) anterior thoracic spiracle; (F) caudal segment, I—stelex sensillum, 2—verruciform sensillum, 3—rima, 4—interspiracular process, 5—inter- mediate sensory complex, stelex sensillum, 6—intermediate sensory complex, medusoid sensillum. 420 tersegmental lines; gnathocephalon conical (Fig. 3B); rugose pads laterad of anterior sensory lobe serrated on ventral margin (Fig. 3B-1, C-1); dorsal sensory organ a dome-shaped papilla (Fig. 3B-2, 3C-2); an- terior sensory lobe (Fig. 3B-3), bears ter- minal sensory organ (Fig. 3C-3), pit sen- sory organ (Fig. 3C-4), lateral sensory or- gan (Fig. 3C-5), and supralateral sensory organ (Fig. 3C-6); stomal sense organ ven- trolaterad of anterior sensory lobe (Fig. 3B- 4, 3C-7); mouth hook bidentate (Fig. 3D- 1); median oral lobe laterally flattened (Fig. 3D-2); pit sensilla circumscribe gnatho- cephalon (Fig. 3B-5); minute acanthae cir- cumscribe anterior margin of prothorax; ru- gose pads and two rows of verruciform sen- silla circumscribe prothorax; anterior tho- racic spiracle bears 7—8 ovoid papillae (Fig. 3E); verruciform sensilla circumscribe me- sothorax; lateral spiracular complexes not seen; caudal segment with two stelex sen- silla, dorsad and ventrad of posterior spi- racular plate (Fig. 3F-1); two verruciform sensilla dorsolaterad of posterior spiracular plate (Fig. 3F-2); posterior spiracular plate bears three ovoid rimae, ca. 0.018 mm in length (Fig. 3F-3), and four interspiracular processes, each with 3—6 branches, longest measuring 0.01 mm (Fig. 3F-4); interme- diate sensory complex with a medusoid sensillum (Fig. 3F-5) and a stelex sensillum (Fig. 3F-6). The second instar bears serrated rugose pads laterad of the mouth lumen similar to the second instars of T. nigricornis and T. pseudovicina (Knio et al. 1996a, Goeden and Teerink 1998). The second instars of T. imperfecta and T. jonesi lack serrated ru- gose pads, although the third instars of both species bear serrated rugose pads (Goeden et al. 1998a, Teerink and Goeden 1998b). In the second instar of 7. wheeleri, the an- terior spiracle bears 7—8 papillae, more than any previously studied congeneric species (Goeden and Teerink 1997b, 1998, 1999; Goeden et al. 1998a, b; Headrick and Goe- den 1991; Knio et al. 1996a; Teerink and Goeden 1998, 1999). The interspiracular PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON processes are not as elaborately branched as those of the first instar. Third instar: White, barrel-shaped, ta- pering anteriorly, rounded posteriorly, mi- nute acanthae circumscribe intersegmental lines (Fig. 4A); gnathocephalon conical (Fig. 4B), rugose pads laterad of mouth lu- men serrated on ventral margin; dorsal sen- sory organ a dome-shaped papilla (Fig. 4C- 1); anterior sensory lobe bears terminal sen- sory organ (Fig. 4C-2), pit sensory organ (Fig. 4C-3), lateral sensory organ (Fig. 4C- 4), and supralateral sensory organ (Fig. 4C- 5); stomal sense organ ventrolaterad of an- terior sensory lobe (Fig. 4C-6); mouth hooks hidden in all prepared specimens; prothorax circumscribed anteriorly with mi- nute acanthae (Fig. 4B-1); rugose pads cir- cumscribe prothorax posteriorad to minute acanthae (Fig. 4B-2); two rows of verruci- form sensilla circumscribe prothorax pos- teriorad to rugose pads (Fig. 4B-3); stelex sensillum located dorsomedially (Fig. 4B- 4); anterior spiracle on posterior margin of prothorax bears 3—5 rounded papillae (Fig. 4B-5, 4D); mesothorax circumscribed an- teriorly with verruciform sensilla (Fig. 4B- 6); metathoracic lateral spiracular complex consists of a spiracle (Fig. 4E-1), a stelex sensillum (Fig. 4E-2), and two verruciform sensilla (Fig. 4E-3); abdominal lateral spi- racular complex consists of a spiracle (Fig. 4F-1) and two verruciform sensilla (Fig. 4F-2); caudal segment circumscribed by minute acanthae; two stelex sensilla, dorsad and ventrad of posterior spiracular plates (Fig. 4G-1); two verruciform sensilla dor- solaterad of posterior spiracular plates (Fig. 4G-2); posterior spiracular plate bears three ovoid rimae, ca. 0.034 mm in length (Fig. 4G-3), and four interspiracular processes, each with 3—6 branches, longest measuring 0.018 mm (Fig. 4G-4); intermediate sen- sory complex (Fig. 4G-5), with a medusoid sensillum (Fig. 4H-1), and a stelex sensil- lum (Fig. 4H-2). The third instar bears serrated rugose pads similar to the third instars of T. im- perfecta, T. jonesi, T. nigricornis, T. pseu- VOLUME 101, NUMBER 2 dovicina and T. signata (Goeden and Teer- ink 1997b, 1998; Goeden et al. 1998a; Knio et al. 1996a; Teerink and Goeden 1998). Compared to the second instar, the third in- star bears only 3—5 papillae on the anterior spiracle. As with the high number of papil- lae in the second instar, so large a reduction in the number of papillae between instars has not been reported in any other T7rupa- nea species (Goeden and Teerink 1997b, 1998, 1999; Goeden et al. 1998a, b; Head- rick and Goeden 1991; Knio et al. 1996a; Teerink and Goeden 1998, 1999). These atypical characteristics call the identity of the second instars examined into question, but this stage was described from 10 rep- licates obtained from three different host species; which would reduce the probability that the same contaminent occurred in all four samples from which second instars were obtained. The mesothorax alone is circumscribed by verruciform sensilla, similar to 7. nigri- cornis (Knio et al. 1996a); whereas both the meso- and metathorax are circumscribed by verruciform sensilla in 7. arizonensis and T. imperfecta (Goeden and Teerink 1998, Teerink and Goeden 1998). The third instar lateral spiracular complex is identical to that in 7. imperfecta (Teerink and Goeden 1998). This is the first instance in which two species of Trupanea have shared the same type and number of sensilla in the metathoracic and abdominal lateral spirac- ular complexes (Goeden and Teerink 1997b, 1998, 1999; Goeden et al. 1998a, b; Headrick and Goeden 1991; Knio et al. 1996a; Teerink and Goeden 1998, 1999). Other pairs of congeneric species in other genera are known to share the same number and type of sensilla in the lateral spiracular complexes, e.g., Procecidochares kristineae and P. lisae; Aciurina idahoensis and A. mi- chaeli; A. thoracica and A. trixa (Goeden and Teerink 1996a, b, 1997a; Headrick and Goeden 1993; Headrick et al. 1997). Puparium: Black, elongate-cylindrical, minute acanthae circumscribe intersegmen- tal lines (Fig. 5A); anterior end bears the 421 invagination scar (Fig. 5B-!) and anterior thoracic spiracles (Fig. 5B-2); caudal seg- ment circumscribed by minute acanthae (Fig. 5C-1), two stelex sensilla, dorsad and ventrad of posterior spiracular plates (Fig. 5C-2); two verruciform sensilla dorsolater- ad of posterior spiracular plates (Fig. 5C- 3); posterior spiracular plate bears three ovoid rimae (Fig. 5C-4), and four interspi- racular processes, each with 3—6 branches (Fig. 5C-5); intermediate sensory complex with a medusoid sensillum and a stelex sen- sillum. Ninety-four puparia averaged 2.53 + 0.03 (range, 1.71—3.17) mm in length; 1.24 + 0.02 (range, 0.85—1.72) mm in width. DISTRIBUTION AND Hosts The distribution of 7. wheeleri in North America north of Mexico as mapped by Foote et al. (1993) included Arizona, Cali- fornia, New Mexico, Oregon, Texas, and Utah in the western United States and a sin- gle record from near the border in adjacent western Canada. Wasbauer (1972) and Goeden (1985, 1986, 1992) reported T. wheeleri from five tribes, 14 genera, and 29 species of host plants in North America. Five new rearing records for T. wheeleri are listed below in the manner of Goeden (1992), which along with taxonomic changes in Hickman (1993) and Goeden et al. (1998b), increase the re- ported host range to include seven tribes, 15 subtribes (Bremer 1994), 28 genera and 47 species. All flies were reared from ca. 1- liter samples of mature flower heads from California. New host genera.—Lasthenia, Machaer- anthera, Psathyrotes, Senecio New host records.—Erigeron aphanactis (Gray) Greene, 3 ¢ and 1 2, SE of Mission Springs at 2480-m elevation, San Bernar- dino Nat. Forest (N section), San Bernar- dino Co., 17.vii.1997; Lasthenia glabrata Lindley, 1 ¢, San Jacinto Wildlife Area at 390 m, Lakeview, Riverside Co., 2.1v.1997; Machaeranthera canescens (Pursh) Gray, | 6 and 1 @, N of Jenks Lake and S of Bar- 422 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Acc.V Spot Magn ; 10:0 KV33.0 | 1607) © ye i ; LC s2 Acc¥ “Spot Magn wD /———4 50um NV , ) Me 2Po \cc.V Spot Magn WD ee te um 10.0KV.30" 477k 20.9 \10.0°kV 3.0 _1179x 26.9 AccV Spot Magn WD /———— 54m 2 Acc.V poh aay) 1P.0kV.3.0 4302x 27.6 10.0 KV 3.0 _ 339x, *. Fig. 4. Third instar of Trupanea wheeleri: (A) habitus, anterior to right; (B) gnathocephalon, prothorax, anter- iolateral view, 1—minute acanthae, 2—rugose pad, 3—verruciform sensillum, 4—stelex sensillum, 5—anterior thoracic spiracle, 6—verruciform sensillum; (C) anterior sensory lobe, |—dorsal sensory organ, 2—terminal sen- sory organ, 3—pit sensory organ, 4—lateral sensory organ, 5—supralateral sensory organ, 6—stomal sense organ; (D) anterior thoracic spiracle; (E) metathorax, 1—spiracle, 2—stelex sensillum, 3—verruciform sensilla; (F) first VOLUME 101, NUMBER 2 ton Flats at 1830-m, San Bernardino Nat. Forest (N_ section), San Bernardino Co., 9.viii.1995; Psathyrotes ramosissima (Tor- rey) Gray, 2 2, Painted Canyon at 155 m, Riverside Co., 19.11.1996; Senecio canus Hooker, 2 6 and 1 2, 1.6 km S of Beach Meadow at 2430 m, Sequoia Nat. Forest (N section), Tulare Co., 15.vii.1993. All of the rearing records for the 45 re- ported hosts of 7. wheeleri are from Cali- fornia, and all but four of these were con- firmed by us or are ours or RDG’s records, including nine recently reassigned from T. actinobola (Goeden et al. 1998b). Of the eight valid host names that Wasbauer (1972) listed, we have confirmed four. As known to date, most hosts for 7. wheeleri belong to the tribe Astereae (26 spp.), with good representation from the Helenieae (9 spp.) and Heliantheae (4 spp.), and token representation from Eupatorieae (2 spp.), Senecioneae (2 spp.), Anthemideae (1 sp.), and Mutiseae (1 sp.) (Munz 1974, Hickman 1993, Bremer 1994). Similarly, the subtribe Solidagininae (21 spp.) is best represented, with additional representation in the sub- tribes Asterinae (5 spp.), Verbesininae (3 spp.), Alomiinae (2 spp.), Baeriinae (2 spp.), Chaenactidinae (2 spp.), Madiinae (2 spp.), Achilleinae (1 sp.), Gaillarediinae (1 sp.), Nassauviinae (1 sp.), Pectidinae (1 sp.), Peritylinae (1 sp.), Senecioninae (1 sp.), Tussilagininae (1 sp.) (Bremer 1994). BIOLOGY Trupanea wheeleri is a difficult species to study in flower heads because it usually co-occurs with other tephritids in the same host-plant species (symphagy) (Goeden 1997), including congeners, and only cer- tain host species sampled at certain loca- tions contained mixes of immature stages of recognizable species. He 423 Egg.—A total of 27 eggs was recovered from preblossom flower heads of Coreth- rogyne filaginifolia (Hooker and Arnott) Nuttall, Ericameria brachylepis (A. Gray) H. M. Hall (=Haplopappus propinquus S. FE Blake) and Hazardia (formeriy Haplo- pappus) squarrosus (Hooker and Arnott) Greene, for an average of 2.5 + 0.2 (range, 1—4) eggs per flower head. All eggs were inserted pedicel-last, mostly through the phyllaries or between their appressed api- ces, perpendicular to or at 30° to 60° to the receptacle, and alongside or into a corolla or ovule of a floret on the periphery or at the center of a closed, immature flower head (Fig. 6A, B). Most eggs were ovipos- ited singly (Fig. 6A, B) or side-by-side, in pairs, or rarely, in threes by single females. Larva.—Upon eclosion, first instars tun- neled into and fed mainly on corollas of florets and on ovules in preblossom flower heads. No receptacle within flower heads was pitted by first-instar feeding in a total of 10 heads of E. brachylepis and H. squar- rosus (Fig. 6C). Second instars fed mainly on ovules and florets of preblossom flower heads and soft achenes of open flower heads (Fig. 6D). Re- ceptacles of a total of 13 flower heads of C. filaginifolia, Ericameria palmeri (Gray) Hall, and H. squarrosus containing second instars averaged 1.9 + 0.3 (range, 0.9-3.7) mm in diameter and none was pitted. These flower heads contained an average of 1.7 + 0.2 (range, 1—3) second instars that had fed upon an average of 5.3 + 0.7 (range, 3—10) florets/ovules/achenes, or 31% (range, 9— 67%) of an average total of 21 + 4.0 (range, 8—58) florets/ovules/achenes per head. Third instars fed on soft achenes at the centers, and less commonly to the margins, of open or postblossom flower heads (Fig. abdominal segment, 1—spiracle, 2—-verruciform sensilla; (G) caudal segment, 1—stelex sensillum, 2—verruciform sensillum, 3—rima, 4—interspiracular process, 5—intermediate sensory complex; (H) intermediate sensory com- plex, 1—medusoid sensillum, 2—stelex sensillum. 424 AccV SpotMagn WD /————-4 100kV 30 44x 208 500 pm » Acc.V = Spot Magn E=10.0 KV 3:0) -248x- ais LLL) ji) GLE. 4 Acc.V Spot Magn WD ‘ 10.0 kV 3.0 247. ~~ 258 C Fig. 5. Puparium of Trupanea wheeleri: (A) hab- itus, anterior end to left; (B) anterior end, 1—invagi- nation scar, 2—anterior thoracic spiracle; (C) caudal segment, I—minute acanthae, 2—stelex sensillum, 3—-verruciform sensillum, 4—rima, 5—interspiracular process. 6E, F). In a total of 28 flower heads of Acourtia (formerly Perezia) microcephala deCandolle, C. filaginifolia, E. palmeri, and H. squarrosus averaging 1.8 + O.1 (range, 1.1—2.6) mm in diameter and containing an PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON average of 1.4 + 0.1 (range, 1—4) third in- stars and 18.8 + 1.9 (range, 6—36) achenes, an average of 6.8 + 1.0 (range 1—23) soft achenes were damaged. Most third instars fed with their long axes oriented perpendic- ular to and mouthparts directed towards the receptacles, within the upper parts of the soft achenes, and well above the receptacles (Fig. 6E). Consequently, only three (11%) receptacles of flower heads that respectively contained three, two, and one third instars were pitted in the 28 flower heads exam- ined. Upon completing feeding, the larvae oriented with their anterior ends away from the receptacles, retracted their mouthparts, and pupariated (Fig. 6F). Pupa.—As noted with other florivorous, congeneric tephritids studied (Goeden and Teerink 1998, 1999: Goeden et al. 1998a, b), flower heads containing puparia (Fig. 6G) had the greatest amount of damage that the seed-feeding larvae of T. wheeleri caused within flower heads sampled. The receptacles of 90 infested, blossom and postblossom flower heads of A. microceph- dla, C. filaginifolia, E. palmeri, and H. squarrosus containing puparia averaged 1.7 + 0.1 (range, 0.9—2.9) mm in diameter and bore an average of 15.2 + 1.1 (7-32) soft achenes, of which an average of 6.9 + 0.6 (range, 1-16) soft achenes or 45% (range, 19—-100%) were damaged. About half of the receptacles were pitted and about half of the puparia were found next to the phyllaries at the margins of flower heads; the remaining flower heads contained puparia at or near their centers (Fig. 6G). All puparia had their anterior ends facing away from the recep- tacles, and their long axes were perpendic- ular to the receptacles (Fig. 6G). Adult.—Adults emerged from mature flower heads, and were long-lived under in- sectary conditions, as 20 unmated males (Fig. 6H, I) averaged 91 + 11 (range, 32-— 230) days, and 23 virgin females (Fig. 6H) averaged 68 + 8 (range, 21—181) days. Like T. actinobola, T. arizonensis, T. jonesi, and T. pseudovicina (Goeden and Teerink 1998, 1999; Goeden et al. 1998a, b), these flies VOLUME 101, NUMBER 2 425 ng Sete, ee Tm, ee i t i i i ' Fig. 6. Life stages of Trupanea wheeleri: (A) egg inserted in corolla of floret of Machaeranthera canescens; (B) egg inserted in corolla of floret of Corethrogyne filaginifolia; (C) first instar in ovules in flower head of M. canescens; (D) second instar tunneling in ovules in flower head of C. filaginifolia; (E) third instar in center of open flower head of C. filaginifolia; (F) late third instar in center of open flower head of Erigeron foliosus Nuttall; (G) puparium in flower head of Erigeron foliosus; (H) mating pair, ventral view; (I) mating pair, side view. Lines = | mm. 426 are among the longer average and maxi- mum adult longevities that we have record- ed for native species of nonfrugivorous Te- phritidae from southern California. Such lengthy longevities are consistent with the aggregative type of life cycle ascribed be- low to this tephritid. The premating and mating behaviors of T. wheeleri were briefly studied in the lab- oratory; however, the petri dish arenas found to be so useful with many other Te- phritinae species (Headrick and Goeden 1994), but unsatisfactory with most 7Tru- panea, facilitated only two matings of 5 and 7 minutes duration each observed between 14:00 and 15:00 h PDST (Fig. 6H, I). See Headrick and Goeden (1994), Knio et al. (1996b), Goeden et al. (1998a, b) for de- scriptions of premating and mating behav- iors of congeneric California species. Seasonal history.—The life cycle of T. wheeleri in southern California follows an aggregative pattern in which the long-lived adults in reproductive diapause overwinter and aggregate to mate on preblossom host plants in the spring (March—April) (Head- rick and Goeden 1994). They reproduce first in the Colorado Desert, then in the higher-elevation Mojave Desert, interior valleys, and coastal areas (Headrick and Goeden 1994). Like T. jonesi (Goeden et al. 1998a) and T. nigricornis (Knio et al. 1996b), reproduction by subsequent gener- ations of these multivoltine tephritids con- tinue thereafter throughout the spring, sum- mer, and fall on a wide range of alternate host plants, as flowering of Asteraceae con- tinues at ever higher elevations and more northerly latitudes in California. Natural enemies.—Seven species of chal- cidoid Hymenoptera were reared from in- dividual puparia and mature flower heads bearing puparia of 7. wheeleri as solitary, primary, larval-pupal endoparasitoids: Eur- ytoma n. sp.? (Eurytomidae), Eurytoma ob- tusiventris Gahan (Eurytomidae), Eurytoma veronia Bugbee (Eurytomidae), Eupelmus sp. (Eupelmidae), Mesopolobus sp. (Pter- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON omalidae), Pteromalus sp. (Pteromalidae), Torymus sp. (Torymidae). ACKNOWLEDGMENTS We thank Andrew C. Sanders, Curator of the Herbarium, Department of Botany and Plant Sciences, University of California, Riverside, for identifications of plants men- tioned in this paper. The parasitoids were identified by Harry E. Andersen, Huntington Beach, California, Michael Gates, Depart- ment of Entomology, University of Califor- nia, Riverside, and John C. Luhman, when he was in the Department of Entomology at Riverside. We also are grateful to Kris- tine Gilbert for technical assistance and to D. H. Headrick for his helpful comments on an earlier draft of this paper. LITERATURE CITED Bremer, K. 1994. Asteraceae Cladistics & Classifica- tion. Timber Press, Inc. Portland, Oregon. Cavender, G. L. and R. D. Goeden. 1982. Life history of Trupanea bisetosa (Diptera: Tephritidae) on wild sunflower in southern California. Annals of the Entomological Society of America 75: 400— 406. Curran, C. H. 1932. New species of Trypanidae, with a key to the North American genera. American Museum Novitates 556: 1-19. Foote, R. H. 1960. A revision of the genus Trupanea in America north of Mexico. U.S. Department of Agriculture Technical Bulletin 1214. 29 p. Foote, R. H. and F L. Blanc. 1963. The fruit flies or Tephritidae of California. Bulletin of the Califor- nia Insect Survey 7, 117 p. Foote, R. H., E L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York. Goeden, R. D. 1985. Host-plant relations of Trupanea spp. (Diptera: Tephritidae) in southern California. Proceedings of the Entomological Society of Washington 87: 564—571. . 1986. New records of Tephritidae (Diptera) from Santa Cruz Island, California. Pan-Pacific Entomologist 62: 326-328. 1987. Life history of Trupanea conjuncta (Adams) on Trixis californica Kellogg in southern California (Diptera: Tephritidae). Pan-Pacific En- tomologist 63: 284-291. 1988. Life history of Trupanea imperfecta (Coquillett) on Bebbia juncea (Bentham) Greene in the Colorado Desert of southern California VOLUME 101, NUMBER 2 (Diptera: Tephritidae). Pan-Pacific Entomologist 64: 345-351. . 1992. Analysis of known and new host rec- ords for Trupanea from California (Diptera: Te- phritidae). Proceedings of the Entomological So- ciety of Washington 94: 107-118. . 1997. Symphagy among florivorous fruit flies (Diptera: Tephritidae) in southern California. Chapter 3 /n: K. Dettner, G. Bauer, and W. Volkl (eds.). Vertical Food Web Interactions: Evolution- ary Patterns and Driving Forces. Ecological Stud- ies 130: 39-52. Goeden, R. D. and J. A. Teerink. 1996a. Life histories and descriptions of adults and immature stages of two cryptic species, Aciurina ferruginea (Doane) and A. michaeli, new species (Diptera: Tephriti- dae), on Chrysothamnus viscidiflorus (Hooker) Nuttall in southern California. Proceedings of the Entomological Society of Washington 98: 415— 438. ———.. 1996b. Life history and descriptions of adults and immature stages of Aciurina idahoensis Steyskal (Diptera: Tephritidae) on Chrysothamnus viscidiflorus (Hooker) Nuttall in southern Califor- nia. Proceedings of the Entomological Society of Washington 98: 681—694. . 1997a. Notes on life histories and descriptions of adults and immature stages of Procecidochares kristineae and P. lisae new species (Diptera: Te- phritidae) on Ambrosia spp. in southern Califor- nia. Proceedings of the Entomological Society of Washington 99: 67-88. 1997b. Life history and description of im- mature stages of Trupanea signata Foote (Diptera: Tephritidae) on Gnaphalium luteo-album L. in southern California. Proceedings of the Entomo- logical Society of Washington 99: 747-754. . 1998. Life history and description of imma- ture stages of Trupanea pseudovicinia Hering (Diptera: Tephritidae) on Porophyllum gracile Bentham (Asteraceae) in southern California. Pro- ceedings of the Entomological Society of Wash- ington 100: 361-372. . 1999. Life history and description of imma- ture stages of Trupanea arizonensis Hering (Dip- tera: Tephritidae) on Trixis californica Kellog var. californica (Asteraceae) in southern California. Proceedings of the Entomological Society of Washington 101: 75-85. Goeden, R. D., J. A. Teerink, and D. H. Headrick. 1998a. Life history and description of immature stages of Trupanea jonesi Curran (Diptera: Te- phritidae) on native Asteraceae in southern Cali- fornia. Proceedings of the Entomological Society of Washington 100: 126-140. 427 1998b. Life history and description of im- mature stages of Trupanea actinobola (Loew) (Diptera: Tephritidae) on Acamprtopappus sphaer- ocephalus Harvey and Gray} Gray (Asteraceae) in southern California. Proceedings of the Entomo- logical Society of Washington !00: 674-688. Headrick, D. H. and R. D. Goeden. 1991. Life history of Trupanea californica Malloch (Diptera: Te- phritidae) on Gnaphalium spp. in southern Cali- fornia. Proceedings of the Entomological Society of Washington 93: 559-570. ———. 1993. Life history and description of imma- ture stages of Aciurina thoracica (Diptera: Te- phritidae) on Baccharis sarothroides in southern California. Annals of the Entomological Society of America 86: 68-79. 1994. Reproductive behavior of California fruit flies and the classification and evolution of Tephritidae (Diptera) mating systems. Studia Dip- terologica 1(2): 194-252. Headrick, D. H., R. D. Goeden, and J. A. Teerink. 1997. Taxonomy of Aciurina trixa Curran (Dip- tera: Tephritidae) and its life history on Chryso- thamnus nauseosus (Pallas) Britton in southern California; with notes on A. bigeloviae (Cocker- ell). Proceedings of the Entomological Society of Washington 99: 415-428. Hickman, J. C. (ed.) 1993. The Jepson Manual. Uni- versity of California Press. Berkeley and Los An- geles. Knio, K. M., R. D. Goeden, and D. H. Headrick. 1996a. Descriptions of immature stages of Tru- panea nigricornis and T. bisetosa (Diptera: Te- phritidae) from southern California. Annals of the Entomological Society of America 89: 1-11. ———.. 1996b. Comparative biologies of the cryptic, sympatric species, Trupanea nigricornis and T. bisetosa (Diptera: Tephritidae) in southern Cali- fornia. Annals of the Entomological Society of America 89: 252-260. Munz, P. A. 1974. A Flora of Southern California. Uni- versity of California Press, Berkeley and Los An- geles. Teerink, J. A. and R. D. Goeden. 1998. Description of the immature stages of Trupanea conjuncta (Ad- ams). Proceedings of the Entomological Society of Washington 100: 431—438. . 1999. Description of the immature stages of Trupanea imperfecta (Coquillett). Proceedings of the Entomological Society of Washington 101: Wasbauer, M. W. 1972. An annotated host catalog of the fruit flies of America north of Mexico (Dip- tera: Tephritidae). California Department of Ag- riculture, Bureau of Entomology Occasional Pa- pers 19, 172 pp. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 428-437 TWO NEW SPECIES OF APHIS L. (HEMIPTERA: APHIDIDAE) FROM ARGENTINA LIVING ON ASTERACEAE M. PILAR MIER DURANTE AND JAIME ORTEGO (MPMD) Departamento de Biologia Animal, Universidad de Leon, E-24071 Leon, Spain (e-mail: dbajnn@unileon.es); (JO) AER Malargiie, Instituto Nacional de Tecnologia Agropecuaria, Saturnino Torres 862, 5613 Malargiie (Mza), Argentina Abstract.—Two new Argentinean aphid species are described: Aphis (Aphis) coridifo- liae, living on Baccharis coridifolia from Cordoba Province, and Aphis (Aphis) melosae, living on Grindelia chiloensis from Mendoza Province. The apterous and alate viviparous females of both species are described, and two forms of A. (A.) melosae apterae are distinguished: ‘“‘big’’ and ‘“‘dwarf.” The differences between the new species and other closely related species are given. Resumen.—‘*Dos nuevas especies de Aphis (Hemiptera: Aphididae) propias de Aster- aceae de Argentina’. Se decriben las hembras viviparas apteras y aladas de dos nuevas especies de pulgones de Argentina: Aphis (A.) coridifoliae, de la provincia de Cérdoba, sobre Baccharis coridifolia, y Aphis (A.) melosae, de la provincia de Mendoza, sobre Grindelia chiloensis. Se distinguen dos tipos de viviparas apteras de A. (A.) melosae: las grandes y las enanas. Se discuten las diferencias entre las nuevas especies y otras pro- ximas. Key Words: The genus Aphis Linnaeus, 1758 and its nominotypical subgenus are the largest ge- nus and subgenus of Aphididae (Remaudi- ére and Remaudiére 1997), and they are mostly distributed in the northern territo- ries. There are few Aphis s. st. species record- ed in the southern territories of the World: 8 in Australia (Eastop 1966), 16 in Sub- Saharan Africa (Millar 1994), approximate- ly 29 in India (Ghosh 1975, 1977, Ray- chaudhuri, Ghosh and Basu 1980) although Numann-Etienne and Remaudiére (1995) recorded seven species in Pakistan, which are not known in India. In South America 25 species of the sub- aphid, new species, pulg6n, especie nueva genus Aphis are known (and two species of the subgenus Protaphis Borner, 1952) (Or- tego and Mier Durante 1997). This number may increase, because of the presence of large areas with favorable climatic condi- tions for this subgenus, especially the southern part of South America. Moreover, a high proportion of recorded species there are endemic: 11/25 species (Remaudieére 1994, Ortego and Mier Durante 1997). Four Aphis s. st. species live on Astera- ceae in South America: Aphis coreopsidis (Thomas 1878) is also known in North America and Africa, A. helianthi (Monell 1879) is also known in North America, and two endemic species A. senecionicoides VOLUME 101, NUMBER 2 Blanchard 1944 and Aphis sp. unpublished (Ortego 1998). Moreover the polyphagous and more or less cosmopolitan A. cracci- vora Koch, 1854, A. fabae Scopoli, 1763, and A. spiraecola Patch, 1914, can live on Asteraceae, and, in fact, they have been re- corded on Asteraceae in South America. Two new species of Aphis s. st. have been found in Argentina on the South American Asteraceae Grindelia and Bac- charis, and these are described here. Abbreviations used in the text and tables are as follows: abd.seg.I to VIII = abdom- inal segment I to VIII; ant. II, IV, V = an- tennal segments III, IV, V; ant.VIb and ant. VIpt = base and processus terminalis of antennal segment VI; BL = body length; D = basal diameter of ant.III; d = diameter of trochanter-femoral joint of hind legs; h.t.II = second segment of hind tarsus; u.r.s. = ultimate rostral segment. Values in pa- rentheses () are exceptional values. Aphis (A.) coridifoliae Mier Durante and Ortego, new species (Fig. 1) Apterous viviparous female (n = 142; 16 measured) (Fig. 1 A—I).—Body 1.00 to 1.70 mm. long, 7.38 to 10.00 (mostly 8 to 9) times siphunculus. Light or greenish yellow when alive, with white waxy powder, retic- ulated, with apex of antenna and legs, si- phunculus and cauda dark brown to black and frequently with lateral dark brown spots on abdomen (intersegmental scler- ites). Prepared specimens light in general with head, most of antenna and legs, ros- trum and sometimes postsiphuncular, and dorso-abdominal VII and VIII sclerites (paler), smoky, and apex of ant.V, antennal segment VI, apex of tibiae, tarsi, sometimes apex of hind femur III, intersegmental sclerites, siphunculus, cauda and genital and anal plates dark brown to black. Dorsal cuticle slightly and irregularly re- ticulated. Setae pale, short (Table 1), acute or dorsal ones slightly blunt. Large and low domelike marginal papillae on prothorax 429 (bigger) and abd.seg.I and VII (exception- ally absent on abd.seg.VII); 2 to 7 (fre- quently 4 to 6) marginal papillae similar in shape, but smaller, on abd.seg.II, III, ['V and VI. Frontal profile convex, slightly sinuate. Antenna 5 or 6 segmented (without corre- spondence with BL), (0.57) 0.62 to 0.90 mm, (0.42) 0.480 to 0.59 times BL; anten- nal segment lengths (in mm): ant.III+IV (5 segmented antenna) = 0.21 to 0.33; ant.III (6 segmented antenna) = 0.10 to 0.25; ant.IV (also 6 segmented) = 0.07 to 0.14; ant.V = 0.08 to 0.15; ant.VIb = 0.08 to 0.12; ant.VIpt = 0.10 to 0.14; ant.III 1.87 to 2.95 (5 segmented antenna) or 0.87 to 1.79 (6 segmented antenna) times longer than ant.VIpt, which is (0.91) 1.14 to 1.47 times longer than ant.VIb. Antennal setae few: 2 to 4 (6) and | to 3 respectively on ant.III and IV (6 segmented antenna) or on their correspondent parts (5 segmented an- tenna). Rostrum (0.33 to 0.40 mm long) reaching hind coxae; BL. 3.03 to 4.56 times length of rostrum; u.r.s. 0.09 to 0.11 mm long, 2.0 to 2.5 (2.7) times as long as its basal width, 0.95 to 1.11 longer than h.t.II, (0.92) 1.00 to 1.25 times ant.VIb, with sides slightly concave and 2 accessory lateral setae. Hind tibia (0.32) 0.37 to 0.44 times BL. First tarsal segment with 3.3.2. setae, as is normal in Aphis (Eastop 1966); h.t.If 0.09 to 0.11 mm long. Conspicuous intersegmental sclerites; a narrow bar across dorsum of abd.seg. VIII in front of setae and sometimes small spinal on abd.seg.VII and sometimes postsiphun- cular sclerites. Abd.seg.I to VI with 2 (rare- ly 3) marginal setae each side and 2 spinal ones; only two setae on abd.seg.VIII. Si- phunculus more or less cylindrical, slowly enlarged on basal third, rough, 0.11 to 0.22 mm long, 2.80 to 4.00 times its width in middle, and 0.89 to 1.26 times cauda. Sub- genital plate with 2 anterior and 4—10 pos- terior setae. Cauda fingerlike, 0.12 to 0.18 430 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Aphis (A.) coridifoliae. A-1, Apterous viviparous female. J-K, Alate viviparous female. A, Habitus. B, Ultimate rostral segment. C, Prothoracic marginal papillae. D, Hind trochanter. E, Hind tarsus. Fk, Marginal part of the abd.seg.VI and VIL. G, Siphunculus. H, Subgenital plate. I, Cauda. J, Abdomen. K, Antennal segments III (above) and VI. Scale bar = 0.615 mm (A, J), 0,40 mm (H), 0,20 (B, D, EB. Gs, 1, K),.05125smm (Cab): Illustrations by Marfa Nieto Gonzalez. VOLUME 101, NUMBER 2 Table 1. foliae, apterous viviparous [ap. viv.] and alatae vivip- arous [al. viv.] females. Setae measurements of Aphis (A.) coridi- ap. Viv. al. viv. ant.II wm long 7-13 10-13 Dtimes 0.5-0.9 0.8—1.0 vertex yum long 10-18 13 D times 0.7-1.4 0.9-1.0 hind trochanter wm long 22-30 20-25 dtimes 0.5—0.8 0.6—0.8 hind femur: dorsal wm long 10-18 10-13 D times 0.7-1.4 0.8-1.0 abd.seg.III: spinal pm long 15-20 12-18 Dtimes 0.9-1.6 1.0—-1.4 abd.seg.III: marginal pmliong 10-20 12-18 D times 0.7—1.4 1.0-1.4 abd.seg. VII: spinal pm long 17-28 20-28 D times 1.2-2.2 1.6—2.2 abd.seg. VIII ym long 20-30 22-28 D times 1.5-2.4 1.8-2.2 mm long, 1.19 to 1.52 times its basal width, with 4 to 6 setae. Alate viviparous female (n = 8; 5 mea- sured) (Fig. 1J, K).—Body 1.22 to 1.48 mm long. Alive and mounted similar to apterae, but darker on head, antenna (ant.I, % distal ant.III, % distal ant.V and VI dark; ant.TII, other parts of ant.IIIJ and V and ant.IV smoky), thorax and legs (excepted basal % of tibiae). Four to eight large and rounded secondary sensoria on ant.III. Marginal sclerites on abd.seg.II to IV present; other dorsal sclerites on abdomen also similar to those in apterae. Other metric and meristic characters (se- tae included, Table 1) very similar to ap- terae, but with following differences: ant.III up to 0.235 mm length and 1.48 to 2.24 times ant.VIpt, siphunculus (narrower than apterae) 3.14 to 3.75 times its medial width, and cauda 1.05 to 1.55 its basal width. Type material.—Holotype: apterous vi- viparous female (measured specimen num- ber 16) collected on Baccharis coridifolia DC at Villa Dolores (Cordoba province, Ar- gentina; 32°00’S, 65°10’W, 540 m), 1-XI- 96, J. Ortego leg., in collection Universidad de Leon (Departamento de Biologia Ani- mal). Paratypes: 141 apterous and 8 alate 431 viviparous females found (J. Ortego leg.) on the same host-plant at the same locality, 3-IX-95 and 1-XI-96, deposited in the au- thors’ collections (Universidad de Le6n, and INTA-Malargiie) and in The Natural History Museum, London and Viuséum Na- tional d’Histoire Naturelle, Par: Etymology.—tThe specific name is an ad- jective used as a substantive in the genitive case derived from the specific name of the aphid’s hostplant: coridifolia (1.C.Z.N., ar- ticle 11 (h) (4) (4), International Commision of Zoological Nomenclature 1985). Biology and distribution.—Aphis (A.) coridifoliae is possibly monoecious and holocyclic on Baccharis coridifolia and perhaps on other related species of Bac- charis (Asteraceae). It forms small and dense colonies on the stems of the host plant, which is distributed, from the center of Argentina to Bolivia, Paraguay and southern Brazil; these territories constitute the potential area of distribution of the new aphid species. Discussion.—Aphis (A.) baccharicola Hille Ris Lambers, 1974 is the only other Aphis species specific on Baccharis; it lives on Baccharis pilularis DC. in California (U.S.A.). It belongs to the Aphis helianthi Monell group (Hille Ris Lambers 1974), and it is a very different species from A. (A.) coridifoliae. Apterous viviparous fe- males of A. (A.) baccharicola have margin- al sclerites, have not marginal papillae on abd.II to VI, and have longer setae (SO—60 wm and 70 wm on abd.seg. III and VIII re- spectively, and setae on ant.seg.III twice as long as D) longer ant.VIpt, and longer si- phunculus, which frequently have setae. Aphis (A.) coridifoliae can be differenti- ated from the majority of species in the sub- genus Aphis recorded in South America, by the short ant.VIpt (1.47 times ant.VIb at most) in both apterous and alate viviparous females. The ant.VIpt is shorter than 1.5 times ant.VIb only in A. (A.) danielae Remaudi- ére, 1994, A. (A.) mulinicola Hille Ris Lambers, 1974 and A. (A.) senecionicoides 432 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Blanchard, 1944. In A. (A.) danielae apter- ous viviparous females, living on Lycium sp. (Solanaceae), the marginal papillae on abd.seg.II to VI are absent, the u.rs. is rel- atively long (at least 0.11 mm) and the dis- cal plate on the abdomen is frequently pre- sent. In A. (A.) mulinicola apterous vivipa- rous females, living on Mulinum (Api- aceae), the discal plate on the abdomen is present and the cauda has 8—10 setae. In A. (A.) senecionicoides apterous viviparous fe- males, living on Senecio (Asteraceae), the marginal papillae on abd.seg. II to VI are absent and the u.rs. is longer (approximate- ly 0.16 mm). Sometimes, the apterous and alatae fe- males of Aphis (A.) schinifoliae Blanchard, 1939 and the apterous females of A. (A.) craccivora Koch, 1854, have the ant.VIpt shorter than 1.5 times ant.VIb. A. (A.) schinifoliae apterae have light and outward- ly curved siphunculi and lack marginal pa- pillae on abd.seg.II to VI and A. (A.) crac- civora apterae have a discal plate on the abdomen and 3 marginal papillae on abd.seg.II to VI at most. Aphis schinifoliae lives on Schinus spp. (Anacardiaceae) and A. craccivora is polyphagous. Aphis (A.) melosae Mier Durante and Ortego, new species (Fig. 2) Apterous viviparous female (n = 301; 34 measured) (Fig. 2A—K).—Two forms are distinguished. (1) “‘big’”’ ones: BL = (1.45) 1.52 to 2.05 mm, dark brown to blackish brown; shining, with abdominal plate and mainly with secondary sensoria on ant.III, and (2) “‘dwarf’’ ones: BL = 0.97 to 1.45 mm, light brown to dark green, more or less opaque, with an incomplete or without ab- dominal plate and without secondary sen- soria. Mounted specimens ‘big’? are dusky light in general, with head, ant.I, II, V (apex) and VI, rostrum, coxae, dorsal part of hind femur (sometimes also front and middle femora) and dorsal plates smoky to light brown, apex of tibiae, tarsi, siphun- culus (darkest), cauda and anal and genital plates dark brown to black; “‘dwarf”’’ spec- imens with ant.II and V and femora, paler than “‘big’’ ones. Dorsal cuticle more or less reticulated. Setae hard, acute and pale (measurements in Table 2). Marginal papillae present on prothorax and abd.seg.I and VII, elevated domelike to ovoidal in shape but different in size, prothoracic ones are largest and ones on abd.seg.VII are smallest and some- times absent in “‘dwarf’’ specimens; (1) 3 to’ 6 (“bis® specimens) Yor’ Oto 2-6) (‘‘dwarf’’) marginal papillae in all on abd.seg.II, HI and IV. Front moderately sinuate, with shallow laterofrontal sinuses. Antenna 6 (“‘big”’ and “‘dwarf’’) or 5 (‘““dwarf’’) segmented; 0.92 to 1:25 (big) or'05S to 0901 GCG dwar) mm long and 0.52 to 0.67 times BL, mea- surements of antennal segments on Table 3; antennal setae few: 5 to 8 (12) (“‘big’’) or 2 to 8 (‘‘dwarf’’) on ant.seg.III (6 seg- mented antenna) or on its correspondent part (5 segmented antenna). “Big” speci- mens with 0 to 6 rounded secondary sen- soria, placed on line on ant.III; sensoria per antenna (34 antennae examined)—O: 26% antennae, 1—3: 50% and 4—6: 24%. In “big” specimens rostrum reaching hind coxae, it is 0.50 to 0.65 mm long and 0.28 to 0.41 times BL; and u.rs. is 0.14 to 0.17 mm long, 2.07 to 3.33 times as long as its basal width and 1.11 to 1.32 longer than h.t.II. In ‘“‘dwarf’’ specimens rostrum reaching up to abd.seg.V, it is 0.44 to 0.57 mm long and 0.34 to 0.47 times BL; and u.rs. is 0.11 to 0.14 mm long, 2.25 to 3.00 times as long its basal width and 1.20 to 1.37 h.t.Il. In both kinds, u.rs. with sides slightly concave and 2 accessory lateral se- tae. Hind tibia 0.43 to 0.58 times BL. First tarsal segment with 3.3.2 setae; h.t.II 0.09 (“dwarf”) or 0.11 (“‘big’’) to 0.14 mm long. Abdomen of “‘big’’ specimens with a spi- no-pleural plate from mesothorax to VOLUME 101, NUMBER 2 43 ee) Fig. 2. Aphis (A.) melosae. A-K, Apterous viviparous female “‘big.’”” L—-M, Alate viviparous female. A, Habitus (on left with pigmentation, on right without pigmentation). B, Antennal segment III. C, Ultimate rostral segment. D, Prothoracic marginal papillae. E, Hind trochanter. FE. Hind tarsus. G, Marginal part of abd.seg.I. H, Marginal part of abd.seg. VII. I, Siphunculus. J, Subgenital plate. K, Cauda. L, Abdomen. M, antennal flagellum. Scale bar = 1 mm (A, L), 0,40 mm (I, J, M), 0,27 mm (E), 0,25 mm (C, F), 0,20 mm (B, D, G, H, K). Illustrations by Maria Nieto Gonzalez. 434 Table 2. alatae viviparous [al. viv.] females. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Setae measurements of Aphis (A.) melosae apterous viviparous [ap. viv.] ““big’’ and *‘dwarf’’ and ap. VIV. “big? ““dwarf”’ al. viv ant. III pm long 15-25 10-23 15-23 D times 0.7-1.4 0.6-1.3 0.8-1.2 vertex wm long 25-33 20-30 25-35 D times 1.3—2.0 1.3—2.2 1.1-2.0 hind trocanter ym long 30-43 25-40 27-37 d times 0.5—0.8 0.6—0.9 0.5-0.7 hind femur: dorsal wm long 25-40 15-25 20-28 D times 1.1—2.0 1.0—2.0 1.1-1.5 abd.seg.I: spinal wm long 22-33 17-30 20-33 D times 1.1—2.0 1.1-—2.0 1.1-1.9 marginal ym long 25-38 17-30 22-33 D times 1.1-2.2 1.1—2.2 1.2—2.0 abd.seg. VII: spinal ym long 30-43 22-38 20-40 D times 1.5—2.3 1.6—2.6 1.1—2.3 abd.seg. VIII wm long 30-45 25-38 32-43 D times 1.7—2.5 1.8—3.0 1.8-2.8 abd.seg.IV or V (from prothorax to abd.seg.VI in early spring) more or less coalescents with marginal sclerites (mostly independent) and frequently with interseg- mental breaks; a transverse bar with differ- ent degree of development on posterior seg- ments. Abdomen of “dwarf” specimens at most with marginal sclerites, a lobulated spino-pleural plate from mesothorax to abd.seg. IV (frequently fragmented) and a narrow transversal bar on posterior seg- ments; but sometimes only several small marginal sclerites and a bar on abd.seg. VIII. Two (exceptionally 1, 3 or 4) setae on abd.seg.VIII and 2 marginal setae Table 3. “dwarf? and alatae viviparous [al. viv.] females. at each side on presiphuncular abdominal segments. Siphunculus more or less cylindrical or slowly cone-truncated, rough, in specimens 0.20 to 0.33 mm long (2.32) 3.42 to 4.71 (5.46) longer its width at mid- dle and 1.26 to 1.65 times the cauda, in ‘dwarf’? specimens 0.09 to 0.20 mm long, (2.22) 2.60 to 4.00 (4.34) longer than its width and 0.95 to 1.18 (1.39) times the cau- da. Subgenital plate with 2 anterior setae and 10-17 (“‘big’’) or 8—13 (“‘dwarf’’) pos- terior setae. Cauda broad fingerlike, in ‘big’? specimens 0.15 to 0.21 mm long, 1.00 to 1.21 (1.38) times its basal width and Antennal segments measurements of Aphis (A.) melosae apterous viviparous [ap. viv.] ““big’” and ap. viv. *‘big”’ ap. viv. “‘dwarf’’ al. viv 6 segmented 6 segmented 5 segmented 6 segmented ant. II (mm) 0.25-—0.36 0.14—0.22 0.17-—0.35 0.25-0.37 ant.LV (mm) 0.11-—0.22 0.06—0.12 0.13-0.23 ant.V (mm) 0.14—0.18 0.06—0.14 0.07-0.13 0:13-0:21 ant. VIb (mm) 0.09—0.12 0.08-0.1 | 0.07—0.10 0.09—0.12 ant. VIpt (mm) (0.18) 0.21—0.26 0.14—0.20 0.14—0.20 0.20—-0.24 ant.1[1/ant. VI..pt. 1.21—-1.68 0.91—1.43 1.17-—1.75 1.19-1.64 ant. VIpt/ant. VIb (1.76) 2.00—2.35 (1.40) 1.70—2.00 1.87—2.18 1.82—2.33 u.r.s./ant. VIb 1.30-1.53 1.28—-1.53 1.35—1.60 1.16—1.50 VOLUME 101, NUMBER 2 with 8 to 12 setae, and in “‘dwarf”’ speci- mens 0.08 to 0.14 mm long, 0.86 to 1.10 (1.25) times its basal width and with 6 to 10 setae. Alate viviparous female (n = 63; 13 measured) (Fig. 2L, M).—Body 1.35 to 2.02 mm long. When alive, brown with head, thorax, antenna (2/10 basal part of ant.III, up to % basal part of ant.IV, and ant.V, dusky), legs (nearly all femur I, % base of femur III and %4 base of tibiae pale to smoky), abdominal bars, siphunculus and cauda dark brown to black. Abdomen with marginal presiphuncular and postsiphuncular sclerites, specially de- veloped intersegmental sclerites abd.seg.I1V— V, isolated sclerites or spinal bar on abd.seg.VI, a wide bar on ant.seg.VII, and 2 isolated sclerites or a bar on abd.seg. VIII. Setae similar to those of apterae (Table 2). Marginal papillae similar in size and shape to those of apterae; some specimens with- out one or both papillae on abd.seg.VII; 1 to 6 ones on abd.seg.II to abd.seg.IV. Antenna 6 segmented, 0.90 to 1.28 mm long and 0.58 to 0.76 times BL; measure- ments of antennal segments in Table 3. Ant.If with 7 to 13 secondary sensoria (sensoria per antenna (26 antennae exam- ined)—7—8: 54% antennae, 9-10: 31% and 11-13: 15%), ant.IV with O to 4 and ant.V very exceptionally with | or 2 (1 on one antenna of 2 specimens and one per antenna in one specimen). U.r.s.= 0.13 to 0.16 mm long, 1.87 to 3.00 its basal width and 1.12 tol 33 °times ht It. Siphunculus 0.13 to 0.27 mm long, 3.25 to 5.40 its width in the middle and 1.19 to 1.43 times the cauda. Subgenital plate with 2 or 3 (exceptionally 4) anterior setae and 8 to 15 posterior ones. Cauda similar in shape to those apterae, 0.10 to 0.19 mm long, 0.91 to 1.34 times its basal width and with 8—11 (15) setae. Type material.—Holotype: apterous vi- viparous female “‘big’’ (measured specimen number 2) collected on Grindelia chiloensis (Corn.) Cabrera at Malargtie (Mendoza province, Argentina, 35°00'S, 69°25’W, 435 1,400 m), 28-X-94, J. Ortego leg. in collec- tion Universidad de Leon (Departamento de Biologia Animal). Paratypes: 300 apterous and 63 alate viviparous females found (J. Ortego leg.) on the same host-plant at the same locality on 17-XI-93, 28-X-94, 6-XII- 94, 25-XI-95, 28-III-96, deposited in the author’s collections (Universidad de Le6én and INTA Malargiie) and in The Natural History Museum, London and Muséum Na- tionale d’ Histoire Naturelle, Paris. Etymology.—The specific name is a noun in the genitive case derived from the common (Spanish) name of the aphid’s host-plant: “‘melosa”’ (1.C.Z.N., article 11 (h) (4) (3), International Commision of Zoo- logical Nomenclature 1985). Biology and distribution.—Aphis(A.) me- losae 1S monoecious on the Asteraceae Grindelia spp., mainly G. chiloensis (Cor- nel.) Cabrera but also on G. tehuelches (Speg.) Cabrera (“‘La Cruz Negra’’, Tupun- gato, Mendoza, Argentina, 21-XI-97, J. Or- tego leg.) and perhaps on other related spe- cies of this genus. It forms dense colonies on the stems and the axil of leaves (“‘big”’ and ‘“‘dwarf’’) and on the underside of the leaves (“‘dwarf’’) of its host plant. The ‘big’ specimens are present from early spring (at the end of September) to the end of spring, and also in autumn (spec- imens found on 19-IV-96). The ‘‘dwarf”’ specimens appear in November and they have been found until March or April. It is evident that the “‘dwarf’’ form is a summer dwarf form (Miyazaki 1987), as in other Aphis spp., for example A. urticata Gmelin, 1790 or A. ruborum (Borner, 1932) in Eu- rope and A. gossypii Glover, 1877 around the World. The alate viviparous females coexist with the “‘big’’ apterous females, but we have not found alatae with the “‘dwarf’’ ones. We have not found sexual forms. It is possible that the species is holocyclic, but it is more probable that it is anholocyclic because the characteristics of its host-plant during the winter permit aphids to live protected. The new species is possibly distributed 436 in dry areas of the southern half of Argen- tina, because Grindelia chiloensis is distrib- uted on sandy or rocky dry areas of the cen- tral western part of Argentina and G. fe- huelches reaches south to Santa Cruz prov- ince. Discussion.—Aphis melosae is a very good example of the variability in aphids. There are several important differences be- tween the apterous viviparous females named by us as “‘big”’ and ‘“‘dwarf’’. These differences are so significant that it would be possible to think that two species are in- volved, but the coexistence of two kinds of apterous females in November, December and April and the characteristics of the alate females (Tables 2—3) allow us to affirm that only one species is involved. Aphis melosae belongs to the “‘cracci- vora’ or ‘“‘Pergandeida” species group, characterised by the presence of a more or less developed thoracico-abdominal or ab- dominal discal plate. Although A. cracci- vora is polyphagous and other species of this group are oligophagous, the majority of species in this group are strict monopha- gous (species living only on one host-plant species) Or non-strict monophagous (spe- cies living on few and related species). Aphis (A.) melosae and Aphis (A.) sp. un- published (Ortego 1998) are the only spe- cies of this group in South America living on Asteraceae (as well as A. craccivora). The apterous viviparous females of Aphis sp. are similar in size to the “‘big”’ speci- mens of A.(A.) melosae, but they do not have marginal papillae on abd.seg.II to VI. The “‘big”? specimens can be differentiated from the apterous viviparous females of the other species of this group having marginal papillae on abd.seg.II to VI, A.(A.) mulini and A.(A.) mulinicola, for the following characters: (1) the ratio ant.VIpt/ant.VIb, more or less 1 in mulinicola, 1.5—1.8 in mu- lini and 1.7—2.4 in melosae; (2) the number of caudal hairs, 8—10 in mulinicola, 10-14 in mulini and 7—12 in melosae; and (3) the host-plant, Mulinum (Apiaceae) for mutlini and mulinicola and Grindelia for melosae. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The “‘dwarf’’ apterous viviparous fe- males of A.(A.) melosae can be differenti- ated from the other Aphis spp. of the nom- inotypical subgenus recorded in South America by the combination of its small size and the host-plant. On the other hand, the metric character- istics of A.(A.) melosae and A.(A.) marthae Essig, 1953, which lives on Quilaja sapon- aria (Rosaceae) in Chile (Essig 1953, Re- maudiére 1994) are very close. Aphis mar- thae apterae and A. melosae “big” apterae are similar in size (BL). They can be dif- ferentiated by the general appearance; the front shape (deeply sinuate in marthae, and moderately sinuate in melosae “‘big’’); the cauda shape (without constriction, nearly triangular in marthae, with constriction, fin- gerlike, in melosae); the marginal papillae on abd.seg.II—VI (papillae exceptionally present on marthae and exceptionally ab- sent in melosae “‘big’’); the secondary sen- soria (always absent on marthae and com- monly present in melosae “‘big’’); the ratio u.r.s/h.t.IT 0,91—1.16 in marthae and 1.11— 1.32 in melosae “‘big’’); and the ratio BL/ rostrum (3.8—4.7 in marthae and 2.44—3.57 in melosae “‘big’’). The alatae females of A. marthae have segmental bands on the ab- domen and 11—21 secondary sensoria on ant.III; the alatae females of A. melosae have segmental bands only on abd.seg.VI- VIII and 7—13 secondary sensoria on ant. III. ACKNOWLEDGMENTS The authors express their gratitude to Prof. Georges Remaudiere (Muséum Na- tional d’Histoire Naturelle, Paris, France) and Prof. Juan M. Nieto Nafria (Universi- dad de Leon, Leén, Spain) for providing useful comments on the manuscript. We also thank Dr. Manuel Marcos, Professor of Latin Philology at the University of Leon, for his help with the names of the taxa de- scribed. This work was funded in part by grant LE20/99 from the Junta de Castilla y Leon, Educacion y Cultura (Spain). VOLUME 101, NUMBER 2 LITERATURE CITED Eastop, V. E 1966. A taxonomic study of Australian Aphidoidea (Homoptera). Australian Journal of Zoology 14: 399-592. Essig, E. O. 1953. Some new and noteworthy Aphidae from Western and Southern South America. Pro- ceedings of the California Academy of Sciences (Fourth Series) 28(3): 59-164. Ghosh, A. K. 1975. A list of aphids (Homoptera: Aphi- didae) from India and adjacent countries. Journal of the Bombay Natural History Society 71(2): 201-220. Ghosh, A. K. 1977. Additions to the list of aphids (Homoptera Aphididae) from India and adjacent countries. Journal of the Bombay Natural History Society 74(1): 29-44. Hille Ris Lambers, D. 1974. On American Aphids, with descriptions of a new genus and some new species (Homoptera, Aphididae). Tijdschrift voor Entomologie 117(4): 103-155. International Commission of Zoological Nomenclature 1985. Code International of Zoological Nomen- clature, Third edition, International trust for Zoo- logical Nomenclature, British Museum (Natural History) and University of California Press. Lon- don, Berkeley and Los Angeles, 338 pp. Millar, I. M. 1994. A catalogue of the aphids (Homop- tera: Aphidoidea) of sub-Saharan Africa. Agricul- ture Research Council (Plant Protection Research Institute Handbook 4). Pretoria, 130 pp. 437 Miyazaki, M. 1989. Forms and morphs of aphids, pp. 27-50. In Minks, A.K. and P. Harrewijin. World Crop Pests: Aphids, Their Biology, Natural Ene- mies and Control, Vol. 2A -vier. Amsterdam. Naumann-Etienne, K. and Remaudiére, G. 1995. A commented preliminary checklist of the aphids (Homoptera: Aphididae) of Pakistan and their host plants. Parasitica 51(1): 1-61. Ortego, J. 1998. Pulgones de la Patagonia Argentina con la descripci6n de Aphis intrusa sp. n. (Ho- moptera: Aphididae). Revista de la Facultad de Agronomia La Plata, 102(1): 59-80. Ortego, J. and Mier Durante, M.P. 1997. Les espéces sud-américaines d’Aphis intéodées au genre Ber- beris (Hemiptera: Aphididae). Annales de la So- cieté Entomologique de France (N.S.) 33(4): 411— 418. Raychaudhuri, D. N., Ghosh, M. R. and Basu, R. C. 1980. Subfamily Aphidinae, pp. 47-278. In Ray- chaudhuri, D. N. Aphids of North-east India and Bhutan. The Zoological Society. Calcutta. Remaudieére, G. 1994. Revue et clé des especes sud- américaines d’Aphidina et description d’un Aphis nouveau [Homoptera, Aphididae]. Revue Franga- ise d’Entomologie (N.S.) 16(3): 109-119. Remaudieére, G. and Remaudiere, M. 1997. Catalogue des Aphididae du Monde/Catalogue of the world’s Aphididae. Homoptera Aphidoidea. Institut Na- tional de la Recherche Agronomique. Paris, 475 PP- PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 438-448 TWO NEOTROPICAL HYPATOPA WALSINGHAM (GELECHIOIDEA: COLEOPHORIDAE: BLASTOBASINAE) WITH RETRACTILE LABIAL PALPI: A PREVIOUSLY UNKNOWN LEPIDOPTERAN FEATURE DAVID ADAMSKI Research Associate, Department of Entomology, Smithsonian Institution, Washington, DC 20560-0127, U.S.A. Abstract.—Hypatopa cryptopalpella, n. sp., is described and H. brevipalpella Wal- singham, 1897, is redescribed. Males of both species possess retractile labial palpi, a feature not known to occur elsewhere in Lepidoptera. Male sex scales on the distal segment of the labial palpi are associated with a deep invaginated pocket from the lower end of the frontoclypeus, extending within the head cavity to the area between the bases of the antennae. The invaginated pocket is filled with a copius brown mass that is believed to be a secretion deseminated by the male sex scales on the labial palpi. The location of the glandular cells cannot be identified until specimens suitable for histo- logical sectioning become available. A lectotype is designated for Auximobasis brevi- palpella Walsingham, 1897, and it is herein transferred to Hypatopa Walsingham, 1907 (new combination). Key Words: The Blastobasinae are small to medium- sized moths, with less than 150 species known world wide. This number, however, greatly underestimates the species richness of the group because many undescribed species, especially from North America and the Neotropics, are represented in museum collections. Although the blastobasine moths are probably one of the most com- monly collected groups of Gelechioidea in the New World, this subfamily may be one least known to science. Since Meyrick (1894) the Blastobasinae have been considered to be a monophyletic group. Recent studies (Adamski and Brown 1989; Hodges 1998) have corroborated this view and have rigorously established the monophyletic relationships among the gen- era and the phylogenetic relationships of the Blastobasinae within Gelechioidea. In retractile labial palpi, Blastobasini, Dominica, Grenada, West Indies this paper the Blastobasidae (sensu Adam- ski and Brown 1989) are treated as a sub- family within the Coleophoridae, following Hodges (1998), and Hypatopa Walsingham, 1907, to which the two species are assigned herein, are referred to the Blastobasin1. Nearly one-fourth of the 69 species of Neotropical Blastobasinae described are from the Lesser Antilles of the West Indies. All but one of these species, Pigritia troctis Meyrick, 1922, were described by Wal- singham (1892, 1897), and most of these species are from St. Thomas. The remain- ing few species are from Grenada, St. Vin- cent, and Barbados. Hypatopa includes 23 species from North American and several species from the Neotropics (new combinations of Wal- singham, Meyrick, and Zeller to be recog- nized in the future by the author). More- VOLUME 101, NUMBER 2 over, the genus contains dozens of unde- scribed species from both faunal regions known by the author. This paper not only describes a new spe- cies of Hypatopa from Dominica but in- cludes another previously described species and hypothesizes their relationship based upon the presence of retractile labial palpi, a feature not previously known to occur in Lepidoptera. METHODS Adults were examined with an incandes- cent light source (reflected light). The Me- thuen 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 mercurochrome and chlora- zol black were used as stains. Slide prepa- rations were examined with dissecting and compound microscopes. Measurements were made with a calibrated ocular microm- eter. Names of genitalic structures are de- scribed and follow Adamski and Brown (1989). The gross morphology of the head of Hy- patopa cryptopalpella Adamski was studied after scales were removed using a fine cam- el’s hair brush, cut about % length. Speci- mens were then placed in glycerine on a depression slide and illustrated using a camera-lucida. The ultrastructure of the head of Hypa- topa cryptopalpella was studied with an Hi- tachi HH-S-2R scanning electron micro- scope at an accelerating voltage of 20 kV. For SEM examination, heads and their ap- pendages were obtained from pinned spec- imens, mounted on stubs with silver paint and paste, and coated with gold-palladium in a Polaron E5100 sputter coater. Hypatopa cryptopalpella Adamski, new species (Figs. 1-17, 20) Diagnosis.—Male with retractile labial palpi, deep invaginated pocket from lower end of the frontoclypeus to the area be- 439 tween the bases of the antennal sockets: valva with dorsal articulation reflexed dis- tally, forming an arch; sacculus smooth, ex- cept outer margin with a cluster of bladelike setae between a dorsal obtuse spine and a ventral row of long marginal setae; base of upper part of valva deeply dentate; apical process of lower part of valva reduced; anellus with two overlapping plates, each with marginal setae; antrum of female with two spinelike projections. Description.—Male head (Figs. 2-3, 15): Vertex and frontoclypeus with gray- ish-brown scales tipped with pale grayish brown, or mostly with grayish-brown scales intermixed with pale-brown scales, or concolorous pale grayish-brown scales. One specimen near white, and few speci- mens with mostly orange-gray scales, in- termixed with pale orange-gray scales; scales of vertex intermixed with few iri- descent bluish-violet sex scales. Antennal scape and pedicel as above, except scape with iridescent bluish-violet squamiform sex scales on ventral surface and dorsodis- tal margins, and cylindrical sex scales on posterior margin near base (Figs. 12—14); squamiform sex scales of scape with nar- row scutes and many windows (Fig. 14), as conpared to adjacent unspecialized scales with broad scutes and few windows (Fig. 15); flagellum grayish brown, with many short sensory hairs, nearly twice di- ameter of female flagellum; retractile labial palpus recurved in front of frontoclypeus (Figs. 5—6) or inserted into a deep invagi- nated pocket from lower end of frontocly- peus, extending within head cavity to the area between bases of antennal sockets (Figs. 2—3); labial palpus with basal seg- ments pale grayish brown, terminal seg- ment with elongate iridescent bluish-violet sex scales on apical part (Figs. 5—11); sex scales with scutes broad and spatulate, with a raised distal part, windows appear absent; proboscis patterned as frontoclypeus. Female head (Fig. 4): As male, except sex scales and invaginated pocket absent; labial palpus longer than that of male (Fig. 440 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Holotype of Hypatopa cryptopalpella. 4), patterned as frontoclypeus, but paler on inner surface; second segment about twice length of terminal segment, basal segment short. Thorax: Tegula and mesoscutum grayish brown intermixed with pale grayish-brown scales; male with dark brown-scales at base. Some specimens with tegula paler than me- soscutum. Legs with outer surface scales brown tipped with white, or brown inter- mixed with white scales; legs mostly white near distal end of segment and tarsomeres, forming ring pattern; inner surface mostly white, intermixed with few brown and pale- brown scales. Forewing (Fig. 1): length 4.9-6.0 mm (n = 33), mostly with pale or- ange-gray scales, or orange-gray scales tipped with white; male with some brown scales along base of costa and base of wing. Some specimens with distal 4 wing darker than basal %. Discal cell with two brown spots near end of cell present or absent, midcell spot absent; undersurface uniform pale brown, except male with dark-brown scales at base extending to about % length of wing within cell; venation (Fig. 16) with four-branched cubitus; CuA, near right an- gle to base of M, and M,. Hindwing pale brownish gray; venation (Fig. 17) with three-branched cubitus. Abdomen: Pale grayish brown, except brown on ventrolateral surface. Male genitalia (Fig. 17): Uncus absent; gnathos medially reduced beneath tuba an- alis; tergal setae present; vinculum a thin band; juxta divided; valva with dorsal artic- ulation reflexed distally, forming an arch; sacculus smooth, except lower margin with a cluster of bladelike setae between a distal obtuse spine and a row of long proximo- VOLUME 101, NUMBER 2 44] basal setae along margin: base of fingerlike upper part of valva with outer margin deep- ly dentate; apical process of lower part of valva reduced; aedeagus s!:ghtly angled, anellus with two overlapping plates, each with marginal setae. Female genitalia (Fig. 20): Ovipositor in four membranous telescopic subdivi- sions; ostium within membrane slightly posterior to segment seven, antrum mem- branous with two large spinelike projec- tions, each pointed posteriorly, and con- vergent basally; inception of ductus sem- inalis within a slightly bulbous part of ductus bursae; corpus bursae subspheri- cal, with spinelike signum. Holotype—¢6, ““DOMINICA, Pont Cas- se, 2 mi[les] N[orth] W[est], V-1965, D. R. Davis’’. The holotype is not dissected and is deposited in the National Museum of Natural History [USNM], Smithsonian In- stitution, Washington, D.C., USA. Paratypes.—12 d, 11 ¢, same data as holotype except, ‘‘d Genitalia Slide by DA 3471, USNM 81624” [green label]; “d Genitalia Slide by DA 3472, USNM 81625” [green label]; ““d Wing Slide by DA 3475, USNM 81628” [green label]; “*? Genitalia Slide by DA 3473, USNM 81626” [green label]; ““? Genitalia Slide by DA 3474, USNM 81627” [green label]; “® Wing Slide by DA 3476, USNM 81629” [green label]. 3 6, 5 2, “DOM- INICA B{[ritish] West] I[ndies], Antrim 1000’ [feet], 11-III-1956, J. E G. Clarke’’, ‘Smithsonian Bredin Exped[ition]”; 5 6, 3 2, same data as above except, “18-III- 1956”; 1 2, same data as above except, ‘10-11-1956’; 1 2, same data as above ex- cept, “‘22-III-1956”. All paratypes are de- posited in the National Museum of Natural History [USNM], Smithsonian Institution, Washington, D.C. Figs. 2-3. Head of Hypatopa cryptopalpella. 2, Etomology.—The Greek prefix CTYPIO Head with labial palpi extended. 3, Head with labial and the suffix, palpelia together mean “hid- palpi within head capsule. Scale = 1.0 mm. den palpus’’, and refer to the retractile la- bial palpi of the male. Remarks.—Hypatopa cryptopalpella dif- fers from H. brevipalpella in having a gray- 442 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 4-9. Scanning electron micrographs of head of Hypatopa cryptopalpella. 4, Frontolateral view of female, 86%. 5, Frontolateral view of male, 102. 6, Inner surface of male right labial palpus, 200X. 7, Lateral view of male right labial palpus, 250. 8, Apical area of male right labial palpus, 650. 9, Apical area of male sex scales on apical portion of labial palpus, 15,000. ish-orange or pale grayish-orange ground ground color of the forewings, all discal color of the forewings, midcell spot of the spots present, and an incomplete median discal cell absent, and the median fascia ab- fascia. Male and female genitalia of both sent. H. brevipalpella has a grayish-brown _ species differ as figured. VOLUME 101, NUMBER 2 443 Figs. 10-15. Scanning electron micrographs of head scales of Hypatopa cryptopalpella. 10, Apical area of male sex scale on apical portion of labial palpus, 15,000. 11, Central area of male sex scale on apical portion of labial palpus, 15,000. 12, Undersurface of basal portion of male left antenna, note arrows pointing to cylindrical sex scales on posterior margin of scape, 175. 13, Cylindrical sex scales on posterior margin of scape, 8,000. 14, Squamiform scale on undersurface of male scape, 25,000. 15, Squamiform unspecialized scale of scape, 7,750. 444 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON S6f4; ty \, oN Aas cs TUR Pea RNG : ab i SS Ci Dae eS Si Figs. 16-17 Wing venation and male genitalia of Hypatopa cryptopalpella. 16, Wing venation. Scale = 1.0 17, Male genital capsule and aedeagus. Scale = 0.5 mm. Hypatopa brevipalpella (Walsingham Diagnosis.—Male with retractile labial 1897), new combination palpi, dorsal articulation of valva reflexed (Figs. 18-19, 21) toward vinculum, sacculus setose, with a small cluster of stout setae near distal mar- Auximobasis brevipalpella Walsingham — gin, juxta divided. Female with microtri- 1897:95; Becker 1984:41. chiate membrane surrounding ostium. VOLUME 101, NUMBER 2 445 Biss 18: Lactotype of Hypatopa brevipalpella. Description.—Head: Vertex and fronto- clypeus with scales mostly grayish brown, intermixed with grayish-brown scales tipped with pale grayish brown; distal por- tion of labial palpus inserted into a pocket in head capsule as in H. cryptopalpella, as far as could be ascertained without dissect- ing unique male specimen; visible basal portion grayish brown. Female labial palpus as in H. cryptopalpella. Antennal scape and pedicel as frontoclypeus, flagellum gray; proboscis pale brown. Thorax: Tegula and mesoscutum mostly with brown scales, intermixed with brown scales tipped with pale brown, and pale- brown scales tipped with white. Legs with outer surfaces mostly brown, intermixed with pale-brown scales, and pale-brown scales tipped with white, inner surface mostly with pale-brown scales, intermixed with white scales. Scales mostly white near distal end of segments and tarsomeres. Forewing (Fig. 18): length 5.0—5.2 mm (n = 3), mostly grayish-brown scales, inter- mixed with pale grayish-brown scales and white scales; costal and outer margin brown, fringe scales between R, and R, with alternating pale and dark patches forming an irregular marginal pattern; me- dian fascia incomplete, obliterated by pale- brown scales near midcell; two dark-brown spots near distal margin of cell, and one faint spot near midcell; a brown streak be- tween wing base and median fascia, poste- rior to CuP. One specimen with distal % darker than basal 4%. Undersurface brown. Hindwing pale brownish gray, gradually darkening to outer margin. Venation not studied. Abdomen: Pale brownish gray. Male genitalia (Fig. 19): Uncus short, apex rounded; gnathos forming a thin band around tuba analis; tergal setae present; vin- culum, a thin ventral support; juxta divided; dorsal articulation of valva reflexed toward vinculum; distal process of lower division of valva not freely articulated with basal part; basal part with marginal setae; saccu- lus setose (with mostly hairlike setae), a small circular cluster of stout setae on distal part; distal process of upper division of val- va with a sclerotized ridge fusing with sac- 446 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 19. culus; upper part of valva setose. Aedeagus broad at base narrowing to apex; anellus uniformly setose. Female genitalia (Fig. 21): Ovipositor as H. cryptopalpella; ostium within membrane slightly posterior to sternum seven; micro- trichiate membrane surrounding ostium; in- ception of ductus bursae and ductus sem- inalis demarcated by short, parallel-sided membranous antrum; inner surface of an- terior % of ductus bursae spinulate; corpus bursae with a spinelike signum. Type examined.—Lectotype here desig- nated, d, “Balthazar, Windward side, Gre- nada, W[est] I[ndies], H. H. Smith [Collec- tor]”. ““BM-d genitalia slide no. 26567”’. Male genital capsule and aedeagus of Hypatopa brevipalpella. Scale = 0.5 mm. In The Natural History Museum, London, England (BMNH). Paralectotypes.—1 ¢, ‘““Mount Gay, Est [action], (Leeward Side), Grenada, W[est] I[ndies], H. H. Smith, 25-30-VIII’’, ‘‘Wal- singham Collection, 1910-427, 65298”’, ‘““Auximobasis brevipalpella WLSM, Type 2”? [Specimen is missing abdomen]. | 2, same data as above except, “‘Paratype 2”’, “BM & genitalia slide no. BM 26568’’. 1 2, same data as above except, “‘I-5-X’’, “Walsingham Collection, 1910-427, 65305”, “‘Paratype 7,”’ [Specimen is not dissected]. All three paralectotypes are in BMNH. Remarks.—Because only one male is VOLUME 101, NUMBER 2 447 Bef fae as { : pes Aye ewer “2 Poss on ps eRe Figs. 20-21. Female genitalia of Hypatopa spp. 20, H. cryptopalpella. 21, H. brevipalpella. Scale = 1.0 448 known of this species, I did not dissect the head to confirm the presence of sex scales on the distal part of the labial palpi or the presence of the deep invaginated cranial pocket. DISCUSSION The invaginated pocket found in male H. cryptopalpella is filled with a copius mass that is probably a secretion deseminated by sex scales on the labial palpi. The location of the glandular cells responsible for the se- cretion of this material cannot be identified until suitable specimens are available for histological studies. The numerous genitalic differences be- tween Hypatopa cryptopalpella and H. brevipalpella are hypothesized to be elab- orations from different branches of the same evolutionary lineage. Consequently, the evolution of the retractile labial palpi in males and the deep invaginated cranial pocket are features that have evolved only once within this lineage. This hypothesis, however, can only be tested through phy- logenetic analysis of Hypatopa. ACKNOWLEDGMENTS I thank Klaus Sattler, Michael Shaffer, and Kevin Tuck of The Natural History Museum, London, England, for their help with examination and photography of type specimens; Niels P. Kristensen, Zoological Museum, University of Copenhagen, Co- penhagen, Denmark, for critical comments of the morphological description of the in- terior head of H. cryptopalpella; and Carl Hansen, of the Office of Imaging, Printing and Photographic Services for the photo- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON graph of the holotype of Hypatopa crypto- palpella. This research was supported in part by grants from NSF Grant BSR85- 01212 and Sigma Xi. LITERATURE CITED Adamski, D. and R. L. Brown. 1989. Morphology and Systematics of North American Blastobasidae (Lepidoptera: Gelechioidea). Mississippi Agricul- tural Forest Experiment Station Technical Bulletin 165. Mississippi Entomological Museum Publi- cation No. 1, 70 pp. Becker, V. O. 1984. Blastobasidae, pp. 41—42. Jn Hepp- ner J. B., ed. Atlas of Neotropical Lepidoptera, Checklist: Part I, Micropterigidae—Immoidea. Dr. W. Junk, The Hague, The Netherlands. xxvii + 112 pp. Clarke, J. E G. 1941. The preparation of slides of the genitalia of Lepidoptera. Bulletin of the Brooklyn Entomological Society 36: 149-161. Hodges, R. W. 1998. Gelechioidea. /n Kristensen, N. P., ed., Handbuch der Zoologie, Lepidoptera, part 1, Volume 35, Berlin, New York: Walter de Gruy- ter & Co. 494 pp. Kornerup, A. and J. H. Wanscher. 1978. Methuen Handbook of Colour, 2nd ed. Methuen and Co., Ltd., London. 243 pp. Meyrick, E. 1894. On a collection of Lepidoptera from upper Burma. Transactions of the Entomological Society of London 1894: 1—29. . 1916-1923. Exotic Microlepidoptera 2, pp. 385—480 (1922); Taylor and Francis, London. Re- printed by E. W. Classey, Ltd., 1969. 640 pp. Walsingham, Lord (Thomas de Grey). 1892. On the micro-Lepidoptera of the West Indies. Proceed- ings of the Zoological Society of London 1891: 492-548. . 1897. Revision of the West Indian micro-Lep- idoptera with descriptions of new species. Pro- ceedings of the Zoological Society of London 1897: 54-183. . 1907. Descriptions of new North American tineid months, with a generic table of the family Blastobasidae. Proceedings of the United States National Museum 33: 197-228. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 449—457 NATIVE BEES (HYMENOPTERA: APOIDEA) IN NATIVE TREES: NYSSA SYLVATICA MARSH. (CORNACEAE) SUZANNE W. T. BATRA Bee Research Laboratory, U.S. Department of Agriculture, Bldg. 476, BARC-East, Beltsville, MD 20705, U.S.A. Abstract.—Nyssa sylvatica (black gum, sour gum, pepperidge) is a functionally dioe- cious, insect-pollinated, ornamental, native tree that blooms from late May through early June in eastern Maryland. Its small, green flowers were visited for their abundant nectar and pollen by 46 species of native bees in 13 genera, and by other insects. Although honey bees were locally numerous, few (only 1.5% of all sampled bees), visited N. syl- vatica and there was no evidence that they displaced native bees on this host. The incon- spicuous flowers with vestigial green petals and sepals may attract pollinators by means of the lenslike, spherical droplets of nectar on glaucous floral discs that sparkle in the sunlight, and concentrate and reflect visible and ultraviolet light. Such flowers are here named ‘‘sparkle-flowers”’ (new coinage). Key Words: Nyssa, Cornaceae, gum, tupelo, Euphorbia, Hedera, insect pollination, bees, competition, reflective nectar, ultraviolet, ““sparkle-flowers”’ The original vegetation of eastern North America after glaciation and before agri- culture consisted of boreal, temperate and subtropical forests. The many native bee species of this region thus would be ex- pected to be best adapted to forage primar- ily on native flowering forbs in spring be- fore the forest canopy closes, and on the flowers of native trees and bushes. Our ex- otic agricultural crops, most ornamental plants, most weeds, and other plants that grow where the primeval forests have been cleared are visited by polylectic native bees, but these are not their original hosts. Sur- prisingly little is known about the bees and other insects that visit the flowers of our native North American trees, attention be- ing diverted to the remote rainforests of Latin America. In order to learn about the normal relationships of native bees and their hosts, I observed and collected the in- sects visiting flowers of several species of native trees. Host trees surveyed included red maple, Acer rubrum L. (Batra 1985); Allegheny chinkapin, Castanea pumila (L.) Mill.; flowering dogwood, Cornus florida L.; sassafras, Sassafras albidum (Nutt.) Nees (Batra, unpublished), and black gum, Nyssa sylvatica Marsh. (this publication). The results of this basic research are ex- pected to be useful for application in natural resource conservation, horticulture, plant breeding, wildlife management, and forest- ry. The possible competition with native bees for floral resources by introduced Eu- ropean honey bees has recently become a controversial issue among conservationists (reviewed by Paton 1996). Therefore, N. sylvatica trees near a large apiary were in- cluded in the survey, to try to find out if honey bees had depleted nectar and pollen, and had displaced native bees. Nyssa sylvatica is an abundant native tree 450 in the mesophytic forests of eastern North America. It ranges from southern Maine through southern Florida, and west through eastern Texas and Michigan, with a small population in the Mexican highlands (Eyde 1963, 1966). Popularly named black gum, black tupelo, sour gum, and pepperidge, it is cultivated as an ornamental tree because of the brilliant red and orange autumnal col- ors of its glossy leaves, and its attractively drooping, crooked branches. Birds eat its juicy, ovoid blue-black fruit (single-seeded drupes), disseminating this tree. The wood is used for veneer, containers, crossties and pallets. There are seven species of Nyssa world- wide, five of them native to eastern North America, one in eastern China, and one in southeast Asia (Tandon and Herr 1971, Wen and Stuessy 1993). Nyssa is closely related to Cornus, and it has been included in the Nyssaceae (Eyde 1963, 1966; Tandon and Herr 1971; Cipollini and Stiles 1991) or the Cornaceae (Cronquist 1988). The tupelo, water gum, or white gum, (N. aquatica L.), grows in the swamps of the southeastern United States (Eyde 1963). Tupelo nectar is the source of a delicately flavored, expen- sive honey, which contains 48% levulose and 24% dextrose, and does not granulate; hives of honey bees are taken on barges into the swamps when tupelo blooms (Wood 1958, Rahmlow 1960a,b). Staminate (male) and pistillate (female) flowers of N. sylvatica are borne on differ- ent trees; thus, this species is dioecious (Ci- pollini and Stiles 1991). A few pistillate flowers may have anthers with abortive pol- len (Eyde 1963). Male trees bear more nu- merous and heavier flowers than do female trees. The depletion of nutrients caused by fruiting results in less frequent and less re- liable flowering in female than in male trees (Cipollini and Stiles 1991). Flowers appear in the late spring, when the leaves are al- most fully expanded. They grow on 3 to 6 cm long pedicels, often below the foliage and largely shaded by leaves. The small (ca. 8 mm long), green, inconspicuous pistillate PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON flowers grow in groups of 2 to 4 on each pedicel. Clusters of staminate flowers are much more conspicuous, due to their larger size, their radiating yellow anthers on long filaments, and the glabrous discs of individ- ual flowers. Several of the small, green sta- minate flowers (each 3 to 5 mm in diame- ter) grow on each pedicel, forming spher- oid, umbel-like clusters of short racemes, up to 1.5 cm in diameter. Flowers of N. syl- vatica would appear to be usually pollinat- ed by the wind, due to their small size, green color, faint to no odor, and inconspic- uousness. However, Eyde (1963) mentioned unspecified bees and other insects visiting them, also wind-borne pollen, collected 160 m from a tree. Three species of solitary bees, Andrena hippotes Robertson, Perdita bradleyi Viereck, and P. townesi Timber- lake have been collected from flowers of N. sylvatica (Hurd 1979). FLOWERING OF N. SYLVATICA About twenty trees were studied from May 19 through June 9, 1997. They grew as scattered trees at the edge of a predom- inantly oak forest near the Bee Research Laboratory at Beltsville, MD, and in a sub- urban setting near my house in Greenbelt, MD, 4.8 km to the south. Insect visitors in the canopy were observed with the aid of 7 x 50 binoculars, and specimens on flowers were collected with a long-handled, aerial net. The weather was variable, being warm (to 36°C), dry, and sunny from May 19 through 23; cool and rainy from May 24 through 27; then mild and sunny from May 28 through June 9. Rain washed the ex- posed nectar from floral discs, but it was replenished during dry weather. Flowering began on May 22, when some anthers on the staminate flowers of male trees growing in sunny areas began to dehisce, and small, spherical droplets of nectar were simulta- neously secreted (Figs. 1,2). Pistillate trees in sunny areas began to bloom on May 23, also secreting nectar, which accumulated at the bases of their erect green styles and in the grooves surrounding their floral discs VOLUME 101, NUMBER 2 451 Figs. 1-3. “Sparkle-flowers” of N. sylvatica, the cushionlike floral discs are about 2 mm in diameter. I, Young staminate flower with vestigial green petals, viewed from above; one stamen has elongated and its yellow anther has dehisced; the glaucous floral disc bears reflective droplets of nectar (arrow). 2, Lateral view of the disc of a staminate flower, showing how the spherical, lenslike droplets stand above its surface. 3, Young pistillate flower (oblique view), with droplets of nectar; some droplets have coalesced around the base of the style, and nectar oozes from the floral disc onto the ovary (arrow). 452 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Bees caught and seen on flowers of N. sylvatica. F = female or worker bee; M = male bee; P = bee is collecting pollen; G = at Greenbelt; B = at Beltsville. Some of the unidentified male and female Andrena may be of the same species (the sexes are dissimilar; there are many species of Andrena, and many are difficult to identify). The female Andrena melanochroa on pistillate flowers was carrying 50% N. sylvatica pollen in her well-filled scopae, this shows that bees can pollinate this tree and they will carry pollen a considerable distance (the nearest male tree was over 100 m away). Bee species or genus, sex and number of individuals On Male Trees On Female Trees Apidae 2E Apis mellifera L. — B 1s Bombus bimaculatus Cresson PG — LE Bombus impatiens Cresson PaG — Anthophoridae IF, 1M Ceratina calcarata Robertson G = SF Nomada denticulata Robertson GB — 1F, 1M Xylocopa virginica (L.) G Megachilidae IF Chelostoma philadelphi Robertson — G 2M Chelostoma philadelphi Robertson B == Halictidae SF Augocholora pura (Say) 23P AG. IB — ive Dialictus cressoni (Robertson) B = 1F D. rohweri (Ellis) G — Ne D. tegularis (Robertson) B — 25 D. versatus (Robertson) B = 2F Dialictus sp. | GY B — ie Halictus (Seladonia) confusus Smith B — SE Lasioglossum sp. nr. coriaceum (Smith) Ph Nex 18} — IF Lasioglossum (Evylaeus) sp. | — G 1F Sphecodes confertus Say G — Andrenidae SE Andrena sp. nr. confederata Viereck lO (PG; B 2G 6M Andrena lamelliterga Ribble GB GiB ibe) Andrena sp. nr. lata Viereck syley (Gels) a IF Andrena melanochroa (Cockerell) — G IM Andrena perplexa vibernella Graenicher B — 2F Andrena perplexa vibernella Graenicher Dike. Gs — SF Andrena rugosa Robertson ZAR — a5 Andrena vicina Smith SP Gyb — IM Andrena (Micrandrena) sp. — G 2F Andrena sp. | IW 124s 18) — ie Andrena sp. 2 — B SF Andrena sp. 3 SP BG G 6F Andrena sp. 4 3pP BG G 2F Andrena sp. 5 23PB = 2F Andrena sp. 6 er as} — IF Andrena sp. 7 — G 1F Andrena sp. 8 B 3 Andrena sp. 9 | PiGoB — IF Andrena sp. 10 G == 1M Andrena sp. 11 G — 2M Andrena sp. 12 — G SM Andrena sp. 13 G G, B IM Andrena sp. 14 G = IM Andrena sp. 15 B = 3M Andrena sp. 16 —_— G, B 1M Andrena sp. 17 — G VOLUME 101, NUMBER 2 453 Table 1. Continued. Bee species or genus, sex and number of individuals On Male Trees n Female Trees Colletidae 8+ F Colletes thoracicus Smith ORG; B 34+ M Colletes thoracicus Smith G,B = IF Colletes willistoni Robertson PB - Ne Hylaeus modestus Say B — 3M Hylaeus modestus Say — G (Fig. 3). Flowering ended on June 9; the last trees to bloom being those growing in understory areas that were shaded by taller trees. During warm, sunny weather, individual pistillate flowers began flowering (secreting nectar) and ceased flowering (stopped nec- tar secretion, with their styles curled, shriv- eled, and brownish) within 24 hours. For example, some pistillate flowers on one tree began secreting nectar and attracted insects at noon on May 23. They had presumably been fertilized by noon the next day, be- cause they had ceased nectar production, and no longer were attracting insects. Other pistillate flowers on the same tree bloomed on other days, so that the total flowering period of this female tree lasted 10 days. The flowering period of individual stami- nate flowers lasted several days. During this time, nectar droplets were continuously se- creted, and the numerous anthers on each flower dehisced sequentially (Fig. 1). Sta- minate trees collectively and individually had a long flowering period. At mid-bloom, each tree had some branches with flowers not yet blooming, other branches with flow- ers in full bloom, and some branches were past bloom, with shriveled stamens falling from their flowers. Staminate flowers had a faint, pleasant, honeylike fragrance, but pistillate flowers seemed to me to be odorless. Both pistillate and staminate flowers produced copious nectar. When first secreted, the nectar ap- peared as small, discrete, transparent, shiny spheres on the glabrous floral discs (Fig. 2). These later enlarged (to 0.2—0.3 mm in di- ameter), then (at 0.5 mm) coalesced, to cov- er the disc with a sheet of glistening, dense, and sweet-tasting nectar. Excess nectar may drip from the discs of pistillate flowers (Fig. 3). Once the nectar has coalesced, the floral disc beneath it loses its pale, bluish gla- brous appearance, and becomes dark green. Nectar secretion by staminate flowers be- gins as their first anthers dehisce (Fig. 1), and ends when all anthers have shriveled and some have begun to fall off. In order to determine whether pollination by wind and insects occurs, several branch- es of a pistillate tree were securely bagged prior to bloom, and the bags were left in place for a month. Brown Kraft paper bags were used to exclude both wind-borne and insect-borne pollen. Fine-mesh gauze bags were used to exclude only insects. No fruit formed on any of the hundreds of bagged flowers, even though a few of them had vestigial anthers. This showed that cross pollination by insects is required. Several other native woody plants in the area bloomed at the same time as N. syl- vatica, potentially competing for visits by pollinating insects. These included huckle- berry, Gaylussacia frondosa (L.) T. and G.; American holly, //ex opaca Ait.; mountain laurel, Kalmia latifolia L.; tuliptree, Lirio- dendron tulipfera L.; staggerbush, Lyonia mariana (L.) D. Don; black cherry, Prunus serotina Ehrh.; black locust, Robinia pseu- doacacia L.; poison ivy, Rhus radicans L.; Rubus sp. and Viburnum sp. Most of these plants had conspicuous, fragrant flowers. Many of the bees that were caught on N. sylvatica flowers included pollen of other plants in their scopae. Allegheny chinkapin, which is very attractive to many species of 454 bees and other insects, began to bloom a day after N. sylvatica finished blooming. American chestnut, Castanea dentata (Marsh.) Borkh., was once a dominant tree of the eastern mesophytic forest; its almost total loss must have significantly altered the numbers and diversity of insects that de- pended on its flowering. BEES ON N. SYLVATICA Flowers Bees and other insects were collected from the pistillate and staminate flowers of about 20 trees (Table 1). Binoculars were used to observe bees high in the canopy, where they could not be collected; these in- cluded numerous foraging females of Col- letes thoracicus Smith, and both sexes of Xylocopa virginica (L.), including males patrolling and defending their territories. Both sexes of C. thoracicus were very abundant at mid-day on May 31 on a sta- minate tree in Greenbelt; some two females per cubic meter of canopy could be seen at a glance, while foraging for nectar and pol- len. This bee is also often abundant on Lir- iodendron tulipfera and Ilex spp. trees. In general, more bees were seen on flowers of N. sylvatica that were in sunlight (in full sun or in sun flecks) than on those in shade (the shady side of the tree, or beneath taller trees, or shaded by the host’s own canopy). During the course of a day, as individual N. sylvatica flowers became sunlit and shaded, bees usually visited them while they were in the sunshine. Most of the bees on N. sylvatica flowers were short-tongued species in the genera Andrena and Colletes; over 135 individuals in 46 species, 13 genera, and 6 families were collected or seen (Table 1). Although bees predominated, other insects also vis- ited N. sylvatica flowers (mostly staminate); they included thrips; syrphid, conopid, cal- liphorid, and other flies; Polistes, Vespula and solitary predaceous wasps; small para- sitic wasps; carpenter ants; cantharid and cerambycid beetles; and adult sawflies. Only two honey bees were collected or seen on flowers. They were on pistillate PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON flowers within 50 meters of the apiary at Beltsville. At the time, this apiary had 73 large colonies with thousands of foragers in hives with 2 supers, plus 8 small colonies, or “‘nucs”. Honey bees comprised only 3 percent of all bees collected from N. syl- vatica flowers near the apiary at Beltsville (2 of 53), and only 1.5 percent of all bees collected (2 of 135). This result was sur- prising, because nectar was always abun- dant on pistillate and staminate flowers, and honey bees produce a large amount of hon- ey for the commercial market from the nec- tar of tupelo trees (N. aquatica). However, N. aquatica and N. sylvatica may not be preferred hosts for honey bees. Perhaps they are forced to forage on tupelo due to the absence of other hosts in the southern swamps. Honey bees did not outcompete or displace native bees on N. sylvatica as ex- pected, including trees growing near nu- merous hives. The reasons for the attraction of bees and other insects to the small, green flowers of N. sylvatica, while more showy alternate hosts were available, were investigated. The staminate flowers have only a faint fra- grance, and the radiating yellow anthers may be attractive (Fig. 1). Nectar exposed on the floral discs is readily accessible to many insects. However, if the easily in- gested, noncrystallizing nectar sugars alone were the main attractant, N. sylvatica flow- ers would be expected to host a wide range of scavengers, and be teeming with these insects, as can be seen feeding on sap at slime fluxes. Instead, short-tongued bees predominated, accounting for about 70 per- cent of all insect visitors. A remarkable feature of the flowers of N. sylvatica is the appearance of the exposed nectar when it is in sunlight. The small, spherical droplets on young flowers glitter against their background of pale blue-green, glaucous floral discs. The sheet of nectar on older flowers also glistens in the sunlight. Even slight movements of the branches cause the sunlit nectar to sparkle. The smooth, glossy upper surface of the leaves VOLUME 101, NUMBER 2 also reflects sunlight, including ultraviolet (Fig. 7). It is probable that bees may be attracted at short range to the glittering of the nectar, which substitutes for the usual showy flowers with nectar guides and dis- tinct fragrances, found among most insect- pollinated plants. Bees and flies are attract- ed to small, shiny objects of metal, and to glass beads (Peis! 1997); sparkling, color- less, odorless nectar may be similarly at- tractive. The shiny leaves of N. sylvatica may also serve to attract bees to the trees, acting as if floral clusters, by assisting in their long-range orientation. Most bee-pol- linated flowers have distinctive shapes and colors that are attractive to bees, including patterns that reflect or absorb ultraviolet. They enhance bees’ learning and memory, thus aiding floral constancy and pollen transmission (Menzel et al. 1997, Lehrer 1997). According to Chittka et al. (1994), green leaves appear uncolored to bees (=bee-white, bee-gray, bee-black), and most flowers have contrasting hues, includ- ing the surprisingly few (4%) that reflect only in UV, which is readily seen and re- membered by bees. Most flowers contrast with surrounding foliage, which tends to absorb ultraviolet. The nectar and stamens of many flowers augment nectar-guide pat- terns on ultraviolet-reflecting petals, by ab- sorbing ultraviolet light; they may fluoresce (Thorp et al. 1975, Tanaka 1982). The nec- tar droplets and opened anthers of N. syl- vatica also fluoresce in ultraviolet. The flowers of cypress spurge, Euphor- bia cyparissias (L.), are also attractive to many bees and other insects (Batra, person- al observation). They resemble flowers of N. sylvatica in visible light, because they are mostly green to greenish-yellow, and they also have exposed droplets of nectar in shallow nectaries. This nectar glistens in direct sun. I photographed flowers of E. cy- parissias and N. sylvatica in sunlight, using a filter that transmits 350—390 nm ultravi- olet wavelengths (Wratten 18A filter, Nikon 55 mm f/3.5 glass lens and Tri-X film). Fo- liage, bracts, and other parts of both plants 455 that had appeared green to yellowish-green in daylight absorbed ultraviolet, appearing dark. The glistening nectar strongly reflect- ed ultraviolet, appearing white, contrasting with the plants’ flowers and foliage (Figs. 4-7). Evidently, insects are attracted by the nectar’s reflected ultraviolet visible light. Thus, like E. cyparissias, the nectar of N. sylvatica reflects ultraviolet, contrasting with surrounding UV-absorbent green fo- liage, pedicels, and other floral structures. The reflectance may be enhanced in young flowers by the spherical shape of the dis- crete droplets, which would act as convex lenses, concentrating both visible and ultra- violet light. The bluish color of the back- ground (the glabrous surface of the disc) may enhance the attractiveness of N. syl- vatica nectar to bees, which respond to blue colors. Floral discs of young staminate flowers that bear numerous spherical, lens- like nectar droplets (Figs. 1,2) resemble the multifaceted lenses that are used as reflec- tors for traffic signals. Honeydew that is secreted by aphids on conifers attracts bumble bees from long dis- tances (Batra 1993). It also glitters brilliant- ly in the sunshine, contrasting with the dark green needles of the conifers. This may be another example of attraction by nectar-re- flection; something worth reflecting about. Flowers that lack showy petals and sepals or a distinct perfume, and that rely on re- flectance from their exposed nectar to at- tract pollinators, represent a distinct cate- gory or pollination system, here termed ‘‘sparkle-flowers’? (new coinage). The small, yellowish-green flowers of ivy (Hed- era helix L.) have exposed nectar droplets borne on convex discs (Barth 1985), thus resembling Nyssa flowers. They attract their host-specific pollinators (Colletes hederae Schmidt and Westrich) only when they are in sunshine (Westrich 1996). Thus, the flowers of ivy probably are yet another ex- ample of “‘sparkle flowers.” In conclusion, this preliminary research demonstrates that we should not assume 456 46 ¥ Figs. 4-7. with a Wratten 18A filter, Nikon 55 mm f/3.5 glass lens and Tri-X film). Reflective nectar droplets are indicated by arrows. 4, Euphorbia cyparissias in sunlight. 5, The same flowers in ultraviolet; white dots are UV-reflecting Reflecting nectar of “‘sparkle-flowers,”’ PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON as seen in sunlight and in ultraviolet (as photographed nectar droplets. 6, Staminate flowers of N. sy/vatica in sunlight. 7, The same flowers in ultraviolet; note that the glossy tops of the leaves reflect. that honey bees (or other exotic bees) com- pete with, and displace, native bees, even when honey bees are abundant, and a host that is known to be visited by honey bees is being considered. It will be necessary to study the complex of bee species that visit each host species during several seasons, to allow for annual fluctuations in weather, in native and exotic bee populations, in flow- ering of the host, and in the presence of other simultaneously flowering host plants that may divert bees (especially wide-rang- ing honey bees) from the plants that are be- ing studied. Honey bees in this study evi- dently had been diverted from N. sylvatica to other, more attractive, resources. The variety of bees collected from N. syl- vatica during a short time, in a small area, suggests that more extensive surveys of this and other North American trees would yield many other insects, including undescribed species. Flowering trees often produce co- pious amounts of nectar and pollen, yet they are relatively unexamined resources that should have profound impacts on the population dynamics of our native bees. Beekeepers are aware of the value of sev- eral species of trees as resources for honey bees. The discovery of “‘sparkle-flowers,” a previously unrecognized type of inflores- cence, in a suburban habitat that has been investigated by many scientists, shows that new phenomena can still be revealed in VOLUME 101, NUMBER 2 such unpromising locations with the aid of simple, inexpensive equipment. The behav- ior of the bees as they approached flowers of N. sylvatica provided clues as to the at- tractant, much as did the behavior of insects at Vaccinium and Gaylussacia plants that had been infected by Monilinia fungi (Batra and Batra 1985). ‘‘Sparkle-flowers” may be a relatively common type of inflorescence, previously overlooked because they appear sO inconspicuous to us. ACKNOWLEDGMENTS I thank my botanist husband, Dr. Lekh R. Batra, for his help with the fieldwork and his review of the manuscript. Dr. S. E. Schlarbaum of the Department of Forestry, Wildlife and Fisheries, University of Ten- nessee, and Dr. B. B. Norden of the De- partment of Entomology, Smithsonian In- stitution, Washington, D.C. also improved the manuscript. LITERATURE CITED Barth, EF G. 1985. Insects and Flowers. George Allen and Unwin, London. 297 pp. Batra, L. R. and S. W. T. Batra. 1985. Floral mimicry induced by mummy-berry fungus exploits hosts’ pollinators as vectors. Science 228: 1011-1013. Batra, S. W. T. 1985. Red maple (Acer rubrum L.), an important early spring food resource for honey bees and other insects. Journal of the Kansas En- tomological Society 58: 169-172. . 1993. Opportunistic bumblebees congregate to feed at rare, distant, alpine honeydew bonanzas. Journal of the Kansas Entomological Society 66: 125-127. Chittka, L., A. Shmida, N. Troje, and R. Menzel. 1994. Ultraviolet as a component of flower reflections, and the color perception of Hymenoptera. Vision Research 34: 1489-1508. Cipollini, M. L. and E. W. Stiles. 1991. Costs of re- production in Nyssa sylvatica: sexual dimorphism in reproductive frequency and nutrient flux. Oec- ologia 86: 585-593. Cronquist, A. 1988. The evolution and classification of 457 flowering plants. New York Botanical Garden, N. YG555) pp: Eyde, R. H. 1963. Morphological and paleobotanical studies of the Nyssaceae, I. A survey of the mod- ern species and their fruits. Journal of the Arnold Arboretum 44: 1—54. 1966. They Nyssaceae in the Southeastern United States. Journal of the Arnoid 47: 117-125. Hurd, P. D., Jr. 1979. The Apoidea, pp. 1741-2209. In Krombein, K. V., P. D. Hurd, Jr., D. R. Smith, and B. D. Burks, eds. Catalog of Hymenoptera in America North of Mexico. Vol. 2. Institution Press, Washington, D.C. Lehrer, M. 1997. Honeybees’ use of spatial parameters for flower discrimination. Israel Journal of Plant Sciences 45: 157-167. Menzel, R., A. Gumbert, J. Kunze, A. Shmida, and M. Vorobyev. 1997. Pollinators’ strategies in finding flowers. Israel Journal of Plant Sciences 45: 141— 156. Paton, D. C. 1996. Overview of feral and managed honeybees in Australia. Report to the Australian Nature Conservation Agency, 71 pp. Peisl, P. 1997. Die Signalfunktionen von Bliiten. Bo- tanica Helvetica 107: 3-28. Rahmlow, H. J. 1960a. Tupelo honey production. Gleanings in Bee Culture 88: 457—461. 1960b. Our nation’s highest priced honey. Gleanings in Bee Culture 88: 532-534. Tanaka, H. 1982. Relationship between ultraviolet and visual spectral guidemarks of 93 species of flow- ers and the pollinators. Journal of Japanese Bot- any 57: 146-159. Tandon, S. R. and J. M. Herr, Jr. 1971. Embryological features of taxonomic significance in the genus Nyssa. Canadian Journal of Botany 49: 505-514. Thorp, R., D. L. Briggs, J. R. Estes, and E. H. Erick- son. 1975. Nectar fluorescence under ultraviolet irradiation. Science 189: 476—478. Wen, J. and T. F Stuessy. 1993. The phylogeny and biogeography of Nyssa (Cornaceae). Systematic Botany 18: 68-79. Westrich, P. 1996. Habitat requirements of central Eu- ropean bees and the problems of partial habitats, pp. 1-16. In A. Matheson, S. L. Buchmann, C. O’Toole, P. Westrich & I. H. Williams, eds. The Conservation of New York. Wood, L. B. 1958. White tupelo—rarest of honeys. Florida Grower and Rancher, March p. 17. \rboretum Smithsonian Bees. Academic Press, PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 458-459 NOTE Validation of Neohydatothrips samayunkur (Kudo) (Thysanoptera: Thripidae) for a Thrips Damaging Marigolds (Tagetes spp.) Mound and Marullo (1996, Memoirs on Entomology International 6: 171) upon examining a paratype of Neohydatothrips pseudoannulipes Johansen (1983, Anales del Instituto de Biologia Universidad Na- cional Aut6noma de Mexico 53: 108.) from Hidalgo, Mexico, concluded that the paratype and specimens collected on Tag- etes sp. plants in Costa Rica were the same species and redescribed the species based on material from Costa Rica. Mound et al. (1996. Australian Journal of Entomology 35: 201) subsequently re- ported N. pseudoannulipes from Queens- land and New South Wales, Australia, on Tagetes erecta L. and T. minuta L. More- over, they treated Hydatothrips (Neohy- datothrips) samayunkur Kudo (1995. Ap- plied Entomology and Zoology 30:169) as a junior synonym of N. pseudoannulipes. Kudo described H. samayunkur from specimens collected on Tagetes sp. in Shi- zuoka, Japan, and also examined speci- mens from Okinawa and Hawaii where it was first reported as N. variabilis (Beach) (Tsuda and Sakimura. 1988. Proceedings of the Hawaiian Entomological Society QSM6): After examining two paratypes of N. pseudoannulipes labeled with different col- lection data than the paratype examined by Mound and Marullo (1996) and specimens collected on marigolds from Hawaii and Australia, I conclude that the marigold specimens are not conspecific with N. pseu- doannulipes. Therefore, Hydatothrips sa- mayunkur Kudo is revalidated (revised sta- tus), and is further redesignated here as Neohydatothrips samayunkur (Kudo) (new combination). Moreover, N. pseudoannu- lipes is the correct name for the species de- scribed by Johansen (1983) and the descrip- tion of N. pseudoannulipes in Mound and Marullo (1996) represents N. samayunkur. A non-type specimen from Mexico with the same collection date and locality as the par- atype of N. pseudoannulipes examined by Mound and Marullo (1996) is identified here as N. samayunkur. Neohydatothrips samayunkur and N. pseudoannulipes are similar in coloration of the antenna and body, but the brown bands on the forewings and the coloration of tib- iae are different. Neohydatothrips sama- yunkur has the occipital apodeme on the head separated from the compound eyes, the forewing has a pale apical band and lacks setae on the hind vein, abdominal seg- ment X is as brown as are segments VII-— IX, and the tibiae are brown medially and otherwise yellow. In contrast, N. pseudoan- nulipes has the occipital apodeme touching the compound eyes, the forewing has a brown apical band and two setae on the hind vein within the brown band, abdomi- nal segment X is pale, and the tibiae are completely yellow. Neohydatothrips variabilis (Beach) is confused occasionally with N. samayunkur but is readily differentiated by the presence of two setae on the hind vein of the fore- wing, abdominal segments [X—X yellowish brown or paler than segments VII—VIII, and the hind tibiae completely yellow. The current distribution of N. samayunk- ur is Costa Rica, El] Salvador, Mexico (Hil- dago, Michoacan), United States (Florida, Hawaii), Australia (New South Wales, Queensland), Japan (Okinawa, Shizuoka), and Sri Lanka. Except for two countries, the reported hosts were marigolds (Tagetes er- ecta L., T. minuta’ Land) Tagetessp:): Specimens from El Salvador were inter- cepted at agricultural quarantine, Houston, VOLUME 101, NUMBER 2 TX, on unknown flowers. The slide label for a specimen from Hidalgo, Mexico, lists Bidens, Eupatorium, and Salvia. A speci- men from Michoacan, Mexico was inter- cepted on grass at agricultural quarantine, San Ysidro, CA, in 1965. 459 Sueo Nakahara, Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, 10300 Baltimore Avenue, Beltsville, MD 20705-2350, U.S.A. (e-mail: snakahar@ sel. barc.usda.gov) PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 459-460 NOTE Sialis vagans (Ross) (Megaloptera: Sialidae) and Amphinemura nigritta (Provancher) (Plecoptera: Nemouridae) Trapped by Mountain Laurel (Kalmia latifolia L.) (Ericaceae) Flowers While sampling a small impoundment in the Frederick City Municipal Forest along Little Fishing Creek (Frederick Co., Mary- land) on May 24, 1998, we observed moun- tain laurel (Kalmia latifolia L.) in full bloom around the pond. Closer examination of the floral clusters revealed large numbers of adult alderflies (Megaloptera: Sialidae). Some alderflies appeared to be stuck to the stems of the flowers by their tarsi; others were stuck to the corolla of the flowers by their wings. Although some of the speci- mens were dead, many were alive. As we removed several alderflies, we noted that the stalk, calyx, and corolla of the flowers were sticky to the touch. We determined the number of floral clus- ters on two bushes (approximately 1.3 m in height) within 1—2 meters of the pond mar- gin, and on a third bush along Little Fishing Creek exiting the pond. The two bushes along the margin of the pond had 25 and 97 floral clusters; the one along the stream had 18. We also counted the adult alderflies on each bush. Those along the pond margin contained 22 and 27 adult alderflies. Simi- lar numbers of alderflies were found on oth- er bushes along the pond margin. The bush along the stream had not trapped any al- derflies; however, we did recover 10 stone- flies (Plecoptera) from the floral clusters. Dr. Oliver S. Flint, Jr. (Department of En- tomology, Smithsonian Institution, Wash- ington, D.C.) identified the alderflies as Sialis vagans (Ross) (Megaloptera: Siali- dae). Ross (Ross. 1937. Bulletin of the I- linois National History Survey 21(3): 57— 78) first described this species from speci- mens collected from lakes and rivers. Dr. Charles Nelson (Department of Biological and Environmental Sciences, University of Tennessee at Chattanooga, Chattanooga, TN) determined the stoneflies to be Amphi- nemura nigritta (Provancher) (Plecoptera: Nemouridae). Mountain laurel is a common shrub dis- tributed from Maine to Mississippi and Al- abama. The inflorescence consists of ter- minal convex flower clusters. Richard A. Jaynes (Jaynes, 1997. Kalmia: Mountain Laurel and Related Species, Timber Press) indicates that the stalk, calyx, and corolla of K. latifolia flowers are covered with glandular, sticky hairs. This sticky secretion is believed to prevent access of crawling insects to pollen and nectar. The poisonous properties of mountain laurel sap are well documented (Jaynes, 1988. Kalmia: The Laurel Book II, Timber Press). The sap contains a group of related 460 grayanotoxins (Mancini and Edwards. 1979. Journal of Natural Products 42(5): 483—488), which occur in a number of spe- cies of Kalmia and other genera in the Er- icaceae. Nonetheless, it is not clear whether the alderflies and stoneflies stuck to the flowers actually died of poisoning or des- iccation. How both insect species moved to the mountain laurel flowers is not known. Sialis vagans and A. nigritta in flight may have selected the mountain laurel bushes along the pond and stream margins to rest or mate. They may also have emerged at or near the pond and stream margins, crawled up the bushes to expand their wings, and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON become stuck to the flowers as they climbed. This novel observation may prove useful to collectors, as more taxa may be collected from mountain laurel flowers. Kevin Stewart is a recipient of an under- graduate fellowship from the Environmen- tal Protection Agency (U-915452-01-0). Some of the optical equipment used for this study was purchased with funds provided by the Fund for Academic Excellence Grants Program, Howard University. K. D. Stewart and R. M. Duffield, De- partment of Biology, Howard University, Washington, DC 20059, U.S.A. (e-mail: rduffield @ fac.howard.edu) PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 460-461 NOTE New Synonymies in Australia Psix Kozlov and Lé (Hymenoptera: Scelionidae: Telenominae) Johnson and Masner (1985. Systematic Entomology 10: 33-58) revised the world species of the telenomine genus Psix Koz- lov and Lé, recognizing 18 species. The ge- nus is primarily found in the Old World tropics, with at least one probable acciden- tal introduction into the New World. In fact, several species had been described before Kozlov and Lé (1976. Zoologicheskii Zhur- nal 55: 143-145) first described the genus from Afghanistan. The varied placement of these early species reflected the confused taxonomy surrounding the large genera Te- lenomus Haliday and Trissolcus Ashmead. The work of Nixon (1935. Transactions of the Royal Entomological Society of Lon- don 83: 73—103, and subsequent papers) al- lowed us to recognize species of Psix among the Afrotropical and Oriental Tris- solcus (then known as Microphanurus Kieffer). However, the Australian species were a more difficult problem. Dodd, be- ginning in 1913, described 102 species of Telenominae, but because he had to rely solely upon the vague, insufficient, and sometimes inaccurate descriptions available at the time, it was difficult to equate his generic concepts with those that developed and were accepted in later years. Johnson and Masner (1985) were able to recognize that Telenomus olympus Dodd was, in fact, a species of Psix on the basis of a specimen identified by Dodd in the Australian Na- tional Insect Collection. Correct generic as- signment of the bulk of his species, how- ever, required first-hand examination of the remaining type material. Johnson (1988. Proceedings of the En- tomological Society of Washington 90: 229-243) reported on the types of Austra- lian species of telenomines described by Dodd and recognized that two further spe- VOLUME 101, NUMBER 2 cies should be placed in the genus Psix: Te- lenomus elpenor Dodd and Telenomus om- phale Dodd. However, the relationship of these two taxa to the species recognized in Johnson and Masner (1985) was not re- solved. Through the kindness of Dr. Gordon Gross of the South Australian Museum, I have been able to study the unique speci- mens of these two Australian Psix and place them in their proper context. Psix elpenor (Dodd) was described from two female specimens collected in Kuranda, northern Queensland (16°40'S, 145°38’E). Dodd’s description is based entirely on a comparison with the species Psix olympus (Dodd), the primary distinguishing charac- teristic being the difference in size between the two (P. elpenor ‘“‘much larger’’). I be- lieve these specimens belong to the same species, with the name Telenomus elpenor Dodd, 1914 becoming a junior synonym of Telenomus olympus Dodd, 1913 (new syn- onymy). Psix olympus is characterized by the acutely tridentate mandibles, the light- colored radicle, and the absence of notauli. The species is known only from the forests of eastern Queensland, from Mt. Tamborine in the south, north to Mossman. Psix omphale (Dodd) was described from a single female specimen collected in Nel- son (present day Gordonvale), Queensland Cig205°S,, 145-47 E). 1 earlier reported (Johnson 1988) that the radicles of the an- tennae are missing from the holotype. They 461 are absent from the slide mount of the an- tennae, but in fact are still attached to the head of the point-mounted specimen, hid- den in the glue that covers the lower part of the face and mouthparts. Most Psix spe- cies are noteworthy for the contrast in color between the radicle (dark brown to black) and the remainder of the scape (usually yel- low), a characteristic quite useful in sepa- rating some Australian species. The radicle of P. omphale is dark, and this specimen belongs to the complex of three closely re- lated species, P. fusus Johnson and Masner, P. metopa Johnson and Masner, and P. gla- briscrobus (Girault). The fore and mid cox- ae are clearly separated by the mesepister- na, thus eliminating P. fusus. The specimen is very similar to P. glabriscrobus, except for the near absence of any transverse mi- crosculpture on the second metasomatic ter- gite. Its body shape, however, matches the stout form of P. glabriscrobus and not the more elongate habitus of P. metopa. Thus, I conclude that Telenomus omphale Dodd, 1913, and Telenomus glabriscrobus Girault, 1916, are synonyms, with Dodd’s name now replacing that of Girault as the valid name for the species (new synonymy). Psix omphale appears to be widely distributed through eastern and central Australia. Norman FE Johnson, Department of En- tomology, The Ohio State University, Co- lumbus, OH 43212-1192, U.S.A. (e-mail: Johnson.2 @ osu.edu) PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 462-463 BooK REVIEW Revision der ‘‘Lestremiinae’’ (Diptera, Cecidomyiidae) der Holarktis. Mathias Jaschhof. Studia Dipterologia—Supple- ment 4. Ampyx-Verlag 1998. 552 pp. Softbound. ISBN 3-932795-03-2. E-mail: ampyxstark @ aol.com. Lestremiinae are the oldest subfamily of Cecidomyiidae, more comparable in their fungal diet, distribution, and presumed age to Mycetophilidae than to the more derived, plant-feeding members of their own family. While the subfamily lacks the direct eco- nomic importance of its plant-feeding cous- ins, it is nonetheless fascinating for its great diversity and for being the source of the rest of the family. The cover’s design of a remarkable as- sortment of 24 lestremiine gonostyli and a cladogram is a winning introduction to the book, showing at the same time some of the groups’s diversity and the establishment of order out of chaos. This study, the result of a thesis for the PhD at the University of Greifswald, Germany, is the first Holarctic revision of the subfamily. The Nearctic spe- cies were last revised in their entirety by A. E. Pritchard between 1947 to 1951. Since then, only a few papers have been pub- lished on the North American fauna, chiefly in response to need, most notably the re- vision of Anarete by K. C. Kim in 1967 in support of ecological studies by H. C. Chiang on swarming behavior. In the Pa- learctic Region, however, the study of this group has attracted a number of specialists, notably B. M. Mamaev, working from 1960 to the present mainly on adult taxonomy but also with larval biology. The Palearctic fauna was in particular need of revision be- cause of the large number of available names and the piecemeal method of its in- vestigation to date. In this work the 528 known species names in Lestremiinae are reduced to 318 valid species. Of the latter number, only 92 are Nearctic, reflecting the limited attention this subfamily has received in North Amer- ica, and 52 species are Holarctic in distri- bution. Anarete and Conarete are treated only as generic entities without species re- vision. These two genera are poorly repre- sented in Europe and the revisions by Kim and Pritchard, respectively, can still serve. Three monotypic genera, Groveriella, Yu- kawamyia, and Baeonotus, are not treated because Jaschhof did not see specimens. Jaschhof presents a fine modern revision. The book includes: overviews of adult Les- tremiinae; analyses of zoogeography and the place of the subfamily within the family and order; a fine section on adult morphol- ogy; a historical review of taxonomic work; and a summary of methods of collection and study. The book has also a key to tribes and genera and, for some large genera, sub- genera and species groups. Keys to species follow the separate generic treatments. There is no mention of larvae. This lack is forgivable in that larvae are known for few species and not all genera and their inclu- sion would have been too much to manage in the time available. The descriptions for the adult stage of genera and species are thorough, the meth- odology careful and consistent throughout, and the illustrations superb. Figures for each species include, as available, details of the gonostylus (often shown in different as- pects), tegmen, penis, and ninth tergite, the fourth flagellomere, and the palpus, and oc- casionally the spermathecae, variation in the genital apodeme, the head, the female postabdomen, and parts of the wing. De- scriptions of the 55 species recently de- scribed by Jaschhof elsewhere are not re- peated here to save space and cost, but il- lustrations for these species do appear here as well as discussions of relationships for each. Results of the author’s methodical anal- VOLUME 101, NUMBER 2 ysis of 41 characters are summarized effec- tively in two cladograms. The author found that the many character regressions and convergences made it difficult to interpret with confidence some of the supraspecific groups. The Lestremiinae are paraphyletic to the monophyletic remainder of the Ce- cidomyiidae, and the source of the latter ap- pears to be among the tribe Catochini. Jas- chhof is careful to treat his classification as tentative, holding out the hope that further collecting and the study of larvae and ex- traterritorial lestremiines will make possible a thoroughly monophyletic classification. Other than one junior generic synonym and two junior specific synonyms unac- countably missing, I found only a few slight errors. A single index to species and genera would have been easier to use than the sep- arate species and generic indices given here, and it would have been helpful to give 463 page references for junior synonyms instead of only referring them to their senior syn- onyms. The book is printed on g!ossy paper with excellent reproduction of the author’s fine drawings. As the reader wil! have noticed from the title, the book is in German, but included are an English summary and an English glossary/synopsis of the morpho- logical terminology used. This latter feature should allow a non German-reader the use of the keys and appreciation of the clado- grams. The figures are, of course, each worth a thousand words in any language. Raymond J. Gagné, Systematic Entomol- ogy Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, U.S.A. (e-mail: rgagne@sel.barc. usda. gov) PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 464-465 REPORTS OF OFFICERS EDITOR Volume 100 of the Proceedings totaled 844 pages. Eighty articles, 5 notes, | book review, | obituary, and the minutes of So- ciety meetings were published. Two Memoirs were published, Memoir 20 “The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae)”’ by Steven W. Lingafelter, and Memoir 21 “New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae)”’ by David G. Furth. Memoir 22, ““System- atics of the North America species of Tri- chogramma Westwood (Hymenoptera: Tri- chogrammatidae)’’ by John D. Pinto, is at the press and will be out at the end of the year on the first of 1999. I extend my appreciation to Gary Miller for his continuing efforts to acquire infor- mative book reviews, and to Tom Henry and Wayne Mathis, members of the Publi- cations Committee, for their continued ad- vice and support. I thank Marie Blair and Cathy Anderson for their much needed as- sistance in handling correspondence, rout- ing manuscripts, and preparation of manu- scripts and plates for the printer. Without their help, my job would have been much more difficult. I am also grateful to the many reviewers for the time-consuming efforts and con- structive reviews. Their contributions are essential to help increase the quality of pa- pers published in the Proceedings. Respectfully submitted, David R. Smith, Editor TREASURER SUMMARY FINANCIAL STATEMENT FOR 1998 Special General Publications Total Fund Fund Assets Assets: November 1, 1997 $14,836.30 $107,833.41 $122,669.71 Total Receipts for 1998 624637211 17,210.80 79,847.91 Total Disbursements for 1998 66,842.67— 13,97330— 80,815.97 Assets: October 31, 1998 10,630.74 111,070.91 121,701.65 Net Changes in Funds $ (4,205.56) S322 30850 $ (986.06) Audited by the Auditing Committee, January 11, 1999, consisting of Donald M. Anderson, Steven W. Lingafelter, and Norman E. Woodley, Chair. Presented to the membership at the meeting of December 3, 1998. Respectfully submitted, Michael G. Pogue, Treasurer VOLUME 101, NUMBER 2 465 MEMBERSHIP In 1998 the Society received applications for new membership from {9 people as follows: Tevis Baier Jeng Ming-Luen Douglas C. Currie Bradley A. Mullens Bonny Dodson Ronald Ochoa David A. Etnier J. H. Pedrosa-Macedo George A. Foster J. Marc Revol Volker Hollman-Schirrmacher Leopold M. Rueda Nayeem Hoq Vera Cristina Silva Matthew D. Kane John Swann Louis M. LaPierre Stephen J. Taft John Wilterding Each applicant was sent a letter acknowledging receipt of his/her application, and his/ her name was read at a regular meeting of the Society and repeated in the minutes of the following meeting. The number of applications for new membership showed a slight decrease from 1997. Respectfully submitted, John W. Brown Membership Chair PROC. ENTOMOL. SOC. WASH. 101(2), 1999, p. 465 New Society PUBLICATIONS (see inside back cover for ordering information) New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae) by David G. Furth Memoir No. 21, 109 pp., 1998 Thirty-eight New World species of leaf beetles are treated. A key is provided and each species is illustrated. Systematics of the North American Species of Trichogramma Westood (Hymenoptera: Trichogrammatidae) by John D. Pinto Memoir No. 22, 287 pp., 1999 This is the first revision of this beneficial group which is important in biological control. A total of 68 species are included, with a key, distribution maps, and illustrations. The scope includes all of North America, including Central America. PROC. ENTOMOL. SOC. WASH. 101(2), 1999, pp. 466-468 SOCIETY MEETINGS 1,032nd Regular Meeting—October 1, 1998 The 1,032nd regular meeting of the Ento- mological Society of Washington (ESW) was called to order at 7:39 pm by President War- ren Steiner, in the Waldo Schmitt Room of the National Museum of Natural History, Washington, D.C. In the absence of a gavel, President Steiner used an electric plug to bring the angry mob to order. Fourteen mem- bers and 8 guests were present. Minutes of the 1,029th and 1,030th meetings were read by pinch-hitting secretary John Brown, and were accepted with minor modification. President Steiner’s request for reports from the committees was answered by Brown, Membership Chair, who read the names of 11 new applicants for member- ship: Louis LaPierre, J. Marc Revol, David Etnier, Leopold Rueda, Bonny Dodson, Douglas Currie, John Swann, Vera Cristina Silva, Matthew Kane, Tevis Baier, and George Foster. One new member was pres- ent—George Foster. Guests in attendance were introduced. President Steiner announced that time was approaching to compile a slate of can- didates for Society officers for next year. He solicited volunteers either to be on the nom- inating committee or to serve as officers, and asked for nominations for honorary membership. In Furth’s absence the show-and-tell por- tion of the program was brief. Ralph Eck- erlin presented a few comments on a recent meeting of the Washington Academy of Sciences, at which he represented ESW; and Jil Swearinger shared with the group her lovely new dung beetle earrings. President Steiner introduced the evening’s speaker, Dr. Charles Triplehorn of Ohio State University, whose football team currently is ranked number | in the nation. Dr. Triple- horn’s talk, entitled ““The rest of the story: a darkling beetle chronicle,’ was an interesting and entertaining look at the joyful field work, interesting personal associations, and gor- geous scenery “behind the scenes” of most of the mundane, though useful, taxonomic works that so many of us produce. The talk was illustrated by title pages of publications, photographs and drawings of tenebrionid beetles, and scenic natural landscapes. Triple- horn’s astute summation of the fact that when you’re sick and tired of a paper, its probably done, struck a responsive chord with many of those present. The meeting was adjourned at 8:40 pm. Refreshments were provided by Jil Swear- ingen. Respectfully submitted, John W. Brown, acting Recording Secretary 1,033rd Regular Meeting—November 5, 1998 The 1,033rd regular meeting of the En- tomological Society of Washington (ESW) was called to order at 7:30 pm by President Warren Steiner, in the Waldo Schmidt Room of the National Museum of Natural History, Washington, D.C. Sixteen mem- bers and eleven guests attended. John Brown’s minutes from the 1,032nd Regular Meeting were read and approved with mi- nor modification. President Steiner asked for reports from the committees. John Brown, Membership Chair, read the names of two new members in ESW: John Wilterding, a graduate stu- dent at Michigan State University, and Dr. J. H. Pedrosa-Macedo, at the Labortorio de Protegao Florestal, Curitiba, Brazil. Chris Thompson listed the ESW officer candidates for next year, noting that, with few exceptions, officers from this year of- ferred to continue in their positions. As al- ways, additional candidates can be taken from the floor before the vote. Ed Barrows presented a collection of sawflies taken in a survey of Malaise traps VOLUME 101, NUMBER 2 suspended high over water. Gabriela Cha- varria displayed the handsome T-shirts still available from the Pollinator Conservation event last June, noting that all profits from T-shirt sales will fund on-the-ground con- servation projects. Ralph Eckerlin dis- played a collection of staphylinids found on Central American mice; whether the beetles are parasitic or phoretic is unknown. Ed Saugstad passed around a 1/2 Penny token from 1793 featuring an image of a bee hive. David Furth introduced Dr. Martha Weiss, an ecological entomologist at Georgetown University. She quickly set the tone of the evening by paraphrasing her presentation’s scholarly title “‘Defecation Ecology: Why do Silver-Spotted Skippers Eject Their Frass?”’ as *“‘Why do the cater- pillars shoot their poop?” The silver-spot- ted skipper is a legume specialist that, as a caterpillar, builds shelters and ejects its frass, building different kinds of shelters as it grows. She described the amazingly reg- ular ontogenetic sequence of shelter-type and proposed mechanisms by which the lar- vae achieve an accuracy of 0.1 mm in leaf- cut distances. Dr. Weiss also described like- ly mechanisms of how caterpillars shoot their poop, which appears to be from changing homeostatic pressure rather than a simple flick. She generously revealed to the audience her unpublished datum of the world’s record holder for frass ejection dis- tance, which is not included in these min- utes so as to avoid scooping her poop shooter news. Her lab is currently exploring three evolutionary hypotheses of why the silver-spotted skipper shoots its poop: pred- ator/parasitoid evasion; hygiene; and space conservation. She closed by asking the au- dience about frass ejection in their own or- ganisms—their study organisms, that is. The meeting was implicitly adjourned around 8:30 pm as the rowdy crowd broke up. Respectfully submitted, Stuart H. McKamey, Recording Secretary 467 1,034th Regular Meeting—December 3, 1998 The 1,034th regular meeting of the En- tomological Society of Washington (ESW) was called to order at 7:35 pin by President Warren Steiner, in the Waldo Schmidt Room of the National Museum of Natural History, Washington, D.C. Twenty mem- bers and twenty seven guests attended. The minutes from the 1,033rd Regular Meeting were read and approved with minor modi- fication. President Steiner asked for reports from the committees. John Brown, Membership Chair, reported that there were no new members in ESW, and none was present at the meeting. Fourteen guests from the USA, Russia, Spain, and Germany were intro- duced. President Steiner thanked the ESW offi- cers, past president, and president-elect, for helping make a good year of Society activ- ities. Dave Smith (Editor) read his annual report. The Treasurer’s annual report and the proposed amendments to the bylaws were passed around the room. Our accounts dropped by over $1,000 but we are still in the black. The ESW gained 19 new mem- bers during the year. Chris Thompson listed the new ESW of- ficer candidates for next year: President, Mike Schauff; Recording Secretary, Stu McKamey; Corresponding Secretary, Holly Williams; Custodian, Andy Jensen; Trea- surer, Mike Pogue; Program Chair, John Brown; Membership Chair, Steve Lingafel- ter; Editor, Dave Smith; and President-elect, Dave Adamski and Dave Furth (a new can- didate nominated from the floor). At Chris Thompson’s suggestion, the uncontested seats were unanimously accepted. A written ballot was made for president-elect, with Furth winning the position. The proposed amendments to the ESW bylaws were discussed: that the Program Chair serve a ““Program Year” rather than a calendar year (Art. V.13, from Dave Furth); that the number of honorary mem- 468 bers be increased from 3 to 6 (and from 4 to 7, if one of the members is elected as honorary president) (Art. III.7, from Stein- er); and that the recommended order of business be changed such that visitors be introduced after new members are intro- duced (Art. VIII.2—3, from Steiner). The vote will be taken at the next meeting if a quorum is present. For exhibits, Ed Barrows (Georgetown University) showed slides of Dyke Marsh, Va., where he is conducting an insect sur- vey, including floating Malaise traps. Dave Furth showed two beetle T-shirts available; Terry Lachman countered with a T-shirt from John May Museum of Boulder, Colo., covered with exquisitely realistic illustra- tions of many insects. Stu McKamey an- nounced the upcoming new on-line re- source ‘“‘New Entomological Taxa,’ and Chris Thompson correctly pointed out that they have not yet announced the subscrip- tion rates. Alma Solis announced that she will be gone all of calendar year 1999. Dave Furth introduced the evening’s speaker, David L. Wagner, from the Uni- versity of Connecticut. The title of his pre- sentation was ‘Natural History and De- fense Strategies of Caterpillars.’’ Dr. Wag- ner began his talk with a synopsis of mor- tality factors of immature Lepidoptera. He PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON then focused in on the importance of avian predators, arguing that birds affect not only what a caterpillar looks like, but also how it eats, when it eats, and even, perhaps, what it eats. Using images from his recently published caterpillar book, Wagner re- viewed many of the strategies that palatable caterpillars use to avoid or deceive birds, from the obvious background matching to those that appear to resemble bird poop, plant debris, snakes, and even the cast skins of tarantulas. Switching gears he displayed an equally varied set of images that illus- trated the diverse defensive strategies of aposematic or warningly colored caterpil- lars. Here we learned of caterpillar toxins, stings, and the ability of some caterpillars to forcibly eject defensive secretions over several centimeters. One of the most note- worthy aspects of the talk was that most of the 80 or more species that were used to illustrate his presentation are to be found within the city limits of Washington, D.C. Don Davis wrapped up the meeting with slides of lichen-feeding tineid moths. The meeting was adjourned at 9:00 pm. Delicious refreshments were generously and abundantly provided by Ralph Ecker- lin. Respectfully submitted, Stuart H. McKamey, Recording Secretary 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 __ : atts ohh Sh Ls a eee eee soe Eo LAMY te EE De $10.00 A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. Eric Gresell. and sMichaelth.“schauits 67ypp O97 t= ae a ee IS ese 15.00 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 _- oS $15.00 No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 _---__ SES 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 TITUS PRISIV TSIEN OD eyo RM EET C49) fie SNA BURSA ee DET SRE CMs SVL NS Se Pal 2 ee ln Ros te PRE eC 15.00 No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato- pogonidac); by W. L. Grogan, Jr: and W.W: Wirth. 125 pp. 1979 2-2 12.00 No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. ETI G By wt RSENS ok es RE ES BRE AD A ae 2 ST eal cic ek 5.00 No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 jB] OY) SSSI Be ON eA SAR DRS 9 Sere Ps 8 de aE oe Eee LL Se ene oon ae Sh ae 25.00 No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent Zusvaly god Da Cra yea ok Fes MUS Fos 0) 0} es! S)2 1>) Rea tah A SG BS ek Bale eRe eee ee a ee Ae aero eae eee 25.00 No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 _ 25.00 No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited byob sNordentand Aj s-. Menke: 2.6: pps 19965 22s aes Le eee 25.00 No. 18. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and Wile WS (reas ts ee AS 0) 0)5, SLL H/o Bee eek ena ge ce POND Pe hae UY a VC BRU roel ee Se RB 25.00 No. 19. Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by AOA AST Sep EAC TAR AOS WIS pero kee a ee Es a a Se he ee 18.00 No. 20. The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- I PeM MSc MMe haf ot “ISIS KS) cee sate heed Ns ea SEMEL Leer 25 EW ge AEGON T UBS ENE ORCL eel le Deora 12.00 No. 21. New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. IPN mem OM oy ot: AS Le be 5 Selene Lie ee RM ea cee Ie Se RUT Rh a es Des OCW Parte Rete 12.00 No. 22. Systematics of the North American Species of Trichogramma Westwood (Hymenoptera: Machorraminaudae) by sohneD. Pinto! 2570pp. 990 nese ee ee ee 28.00 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 except on prepaid orders. Dealers are allowed a discount of 10 per cent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Custodian, Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. CONTENTS (Continued from front cover) MANLEY, DONALD G.—Dasymutilla tomberlini, a new species of velvet ant (Hymenoptera: Mutiliidae); fromiNe ws Mexico)», 25) 3 Jaco as eae Reasearch os ene rete Nyaa Ss en oc era 412 MARSHALL, S. A.—New North American species and records in the genus Xenolimosina Robacek (Diptera: Sphaeroceridae: Limosiminae)e a2. - see ea. ieee oe eee east eee 270 MATHIS, WAYNE N.—A review of the beach-fly genus /socanace Mathis (Diptera: Cana- 73 (0-1) | OBE RARE Hee SAE REM R smmene eti cn a Ee Ma RA Ea AN is te memo e ea gabe S00 BO. 347 MIER DURANTE, M. PILAR and JAIME ORTEGO—Two new species of Aphis L. (Hemiptera: Aphididae)! from :Argentina living on) Asteraceae tga. aes sees ee fee eee ae dee eee eee 428 MOULTON, STEPHEN R., II, STEVEN C. HARRIS, and JOSEPH P. SLUSARK—The mi- crocaddisfly genus /thytrichia Eaton (Trichoptera: Hydroptilidae) in North America ........ 233) NAKAHARA, SUEO and RENATA C. MONTEIRO—Frankliniella zucchini (Thysanoptera: Thripidae), a new species and vector of Tospovirus in Brazil ................2..:202eeeeeee ees 290 O’ NEILL, KEVIN M. and HOWARD E. EVANS—Observations on the prey and nest clusters ot Podalonia. valida (Cresson) (Hymenoptera; Sphecidae) i. cen ee nsee sae eee eee Si, POGUE, MICHAEL G. and ANNETTE AIELLO—Description of the immature stages of three species of Eulepidotis Guenée (Lepidoptera: Noctuidae) with notes on their natural history ...._ 300 PUJOL-LUZ, JOSE R. and ROBERTO pE XEREZ—The larva of Chalcidomorphina aurata En- derlein 1914 (Diptera: Stratiomyidae) from ‘Ilha de Marambaia,’’ Rio de Janeiro, Brazil .... 295 RIBEIRO-COSTA, CIBELE S.—Description of two new species of Amblycerus Thunberg (Co- leoptera: Bruchidae) with a probable stridulatory mechanism ......................2..0020eee0e: 337 SMITH, DAVID R.—Identity of Syrista speciosa Mocsary and notes on the genus Urosyrista Maa (iiymenoptera:|Cephidae) i. 5 i. done serene Meta ieee traleidiag Stat a meh es oie Re ae age a ap SET 285 TEERINK, JEFFREY A. and RICHARD D. GOEDEN—Description of immature stages of irupanea imperfecta (Coquillett) (Diptera: Tephritidae) ai. 2 s.eeessereates see tee ee eee aes 225 WHEELER, A. G., JR.—Craspedolepta eas (McAtee) and Trialeurodes phlogis Russell (He- ‘miptera: Sternorrhyncha: Psyllidae and Aleyrodidae): New distributional and host-plant rec- ordssoh twolittle-knownr Phlox: Specialists) Haupt Bocydium Latreille (9 spp.) Stylocentrus Stal (3 spp.) Umbelligerus, New Genus (1 sp.) Oeda Amyot and Serville (4 spp.) Lycoderes Germar (22 spp.) Stegaspis Germar (10 spp.) Flexocentrus Goding (2 spp.) Lirania Stal (1 sp.) Cryan and Deitz (present) Subfamily Stegaspidinae Haupt Tribe Stegaspidini Haupt Bocydium Latreille (15 spp.) Stylocentrus Stal (3 spp.)* Umbelligerus Deitz (3 spp.)" Oeda Amyot and Serville (4 spp.)* Lycoderes Germar (36 spp.)* Stegaspis Germar (2 spp.)* Flexocentrus Goding (1 sp.)* Lirania Stal (1 sp.) Smerdalea Fowler (4 spp.) (Euwalkeria removed from Stegaspidini by Deitz and Dietrich [1993a]) Euwalkeria Goding (1 sp.) | Species counts based on Metcalf and Wade (1965a). > As Stegaspinae. * As Stegaspini. + Discussed in Cryan and Deitz (in preparation). Finally, independent phylogenetic ana- lyses of morphological characters (Cryan and Deitz, in preparation; Dietrich et al., in preparation) and DNA nucleotide sequence data (Cryan and Wiegmann, in preparation) indicate that the genus Deiroderes Ramos is included in the subfamily Stegaspidinae. The tribal affiliation of Deiroderes, how- ever, is unclear, and may even warrant a new tribe within the subfamily. Cryan and Deitz (in preparation) will present a revi- sion of Deiroderes based on morphology. ACKNOWLEDGMENTS We are grateful to R. L. Blinn, W. M. Brooks, S. C. Mozley, H. H. Neunzig, and B. M. Wiegmann, who offered many help- ful suggestions for improving the manu- script. S. H. McKamey and C. H. Dietrich both supplied invaluable information, ad- vice, and specimens. We also thank: V. M. Knowlton, for tech- nical assistance with scanning electron mi- croscopy; T. J. Henry, for lending a custom SEM specimen stub; J. W. Hardin, for as- sistance with botanical nomenclature; U. K. Cryan, for helping to validate distribution records and assemble SEM plates. For lend- ing specimens, we are indebted to: R. T. Schuh (AMNH); M. D. Webb and J. Mar- gerison-Knight (BMNH); S. E. Miller and K. Arakaki (BPBM); J. A. Chemsack (CISC); K. G. A. Hamilton (CNCID); G. Moragues (GMPC); M. A. Gaiani (IZAV); R. Danielsson. CUMZLU); -R. L. Blmn GNCSU)s -R. OW; “Brooks, .(SEMCG):.S; -H. McKamey (SHMC); T. K. Wood (TKWC); R. C. Froeshner, G. E Hevel, and C. H. Die- trich (USNM); H. Striimpel (ZMUH). This research was funded in part by the North Carolina Agricultural Research Ser- vice, North Carolina State University (Ra- leigh, North Carolina). Funds for electron microscopy were provided by a minigrant from the Center for Electron Microscopy, College of Agriculture and Life Sciences, North Carolina State University. This work is based on a portion of the thesis submitted by J. R. C. in partial fulfillment of his M.S. degree in entomology, North Carolina State University. LITERATURE CITED For consistency within membracid liter- ature, we conform to the letter designations for publications listed in bibliographies by Metcalf and Wade 1963a, Deitz and Kopp 1987a, Deitz 1989a, and McKamey 1998a. Amyot, C. J. B. and A. Serville. 1843a. Deuxiéme par- tie. Homopteres. Homoptera Latr., pp. 455—676 In 488 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Histoire Naturelle des Insects. Hémipteres. Li- braire Encyclopédique de Roret, Paris, France, 1— Ixxvi +675) pp: + lips vatlass (Sapp a 2 plates]. Arnett, R. H. Jr, G. A. Samuelson, and G. M. Nishida. 1993a. The Insect and Spider Collections of the World. 2nd edit. 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Deitz, L. L. 1975a. Classification of the higher cate- gories of the New World treehoppers (Homoptera: Membracidae). North Carolina Experiment Sta- tion Technical Bulletin 225: [i-iv], 1-177. 1983b. Name changes in the Membracidae (Homoptera). Proceedings of the Entomological Society of Washington 85: 856-857. 1989a. Bibliography of the Membracoidea (Homoptera: Aetalionidae, Biturritiidae, Membra- cidae, and Nicomiidae) 1981-1987. North Caro- lina Agricultural Research Service Technical Bul- letin 290: 1-31. Deitz, L. L. and C. H. Dietrich. 1993a. Superfamily Membracoidea (Homoptera: Auchenorrhyncha). I. Introduction and revised classification with new family-group taxa. Systematic Entomology 18: 287-296. Deitz, L. L. and D. D. Kopp. 1987a. Bibliography of the Membracoidea (Homoptera: Aetlionidae, Bi- turritiidae, Membracidae, and Nicomiidae) 1956— 1980. North Carolina Agricultural Research Ser- vice Technical Bulletin 284, [ii] + 39 pp. Dietrich, C. H. and L. L. Deitz. 1993a. Superfamily Membracoidea (Homoptera: Auchenorrhyncha). II. Cladistic analysis and conclusions. Systematic Entomology 18: 297-311. Fabricius, J. C. 1803a. Systema Rhyngotorum secun- dum ordines, genera, species, adiectis synonymis, locis, observationibus, descriptionibus. C. Rei- chard, Brunsvigae, x + 21 [index] + 314 pp. Fairmaire, L. M. H. 1846b. Revue de la tribu des Membracides. (Suite et fin.). Annales de la Sociéte Entomologique de France (série 2) 4: 479-531. Fowler, W. W. 1896e. Order Rhynchota. Suborder He- miptera-Homoptera. (Continued). Biologia Cen- trali-Americana 2: 161-168. Funkhouser, W. D. 1927f. Membracidae. General Cat- alogue of the Hemiptera. Fascicle 1: 1-581. . 1951a. Homoptera Fam. Membracidae. Gen- era Insectorum 208: 1—383; plates 1—xiv. Germar, E. E 1835a. Species Membracidum Musae E. FE Germari. Revue Entomologique (G. Silber- mann, Strasbourg) 3: 223-262. Goding, F W. 1926e. Classification of the Membraci- dae of America. 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Entomologicheskoe Obozrénie 61(3): 518-536. [In Russian; English translation (1982b) in Entomological Review (Washington, D:C.) 61): 70-78.] Stal, C. 1862e. Bidrag till Rio Janeiro-traktens Hem- ipter-fauna. II Ofversigt af Kongliga [Svenska] Vetenskaps Akademiens Forhandlingar 3: 1—75. Strimpel, H. 1972a. Beitrag zur Phylogenie der Mem- bracidae Rafinesque. Zoologische Jahrbticher, Ab- teilung fiir Systematik Okologie und Geographie der Tiere 99(3): 313-407. 1983a. Homoptera (Pflanzensauger). Hand- buch der Zoologie 4(28): i—vi, 1-222, vii—xi. Suchantke, A. 1983a. Die Buckelzirpen (Membraci- dae) und die Formensprache der Insekten. Geo- theanistische Naturwiss (Stuttgart) 3: 74—90. Vignon, P. 1930a. Introduction a la Biologie Expéri- mentale. Les étres organisés activités, instincts, structures. Encyclopédie Biologique (P. Lecheva- lier, Paris) 8: 388-389, 392-393, 403-413. Walker, FE 1851a. List of the Specimens of Homopter- ous Insects in the Collection of the British Mu- seum (Trustees [of the British Museum], London) 2: 261-636. Wood, T. K. 1984a. Life history patterns of tropical membracids (Homoptera: Membracidae). Socio- biology 8: 299-344. PROC. ENTOMOL. SOC. WASH. 101(3), 1999, pp. 490-495 PLATYSCYTISCA BERGMANNAE, A NEW GENUS AND SPECIES OF NEOTROPICAL PLANT BUG RESEMBLING SPECIES OF PLATYSCYTUS REUTER (HETEROPTERA: MIRIDAE: PHYLINAE) Luiz A. A. COSTA AND THOMAS J. HENRY (LAAC) Departamento de Entomologia, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta Boa Vista, Rio de Janeiro, RJ, Brazil 20942; (TJH) Systematic Ento- mology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, Washington, DC 20560-0168 U.S.A. (e-mail: thenry @sel.barc.usda.gov) Abstract.—The new genus Platyscytisca is described to accommodate the new species P. bergmannae, collected in Sao Paulo, Brazil, on Ficus sp. A dorsal and ventral habitus, male genitalia, male genital capsule, and male and female antennae are illustrated to help with recognition. Amazonophilus Carvalho and Costa is resurrected from synonymy under Platyscytus Reuter, and its relationship to Platyscytisca is discussed. Key Words: mannae, new species, Brazil During cooperative work on New World Miridae, we discovered a peculiar new phy- line that was taken on Ficus sp. in Sao Pau- lo, Brazil. Externally, this new species re- sembles some taxa now included in the Neotropical genus Platyscytus Reuter (Car- valho 1958, Carvalho and Costa 1994, Schuh 1995) or Amazonophilus (Carvalho and Costa 1993), a genus recently synony- mized by Kerzhner and Schuh (1995). Herein, we describe the new genus Pla- tyscytisca to accommodate the new species Platyscytisca bergmannae and provide a dorsal and ventral habitus and illustrations of the pretarsus, male genitalia, male genital capsule, and male and female antennae. Amazonophilus is resurrected from synon- ymy under Platyscytus, and the relationship to Platyscytisca is discussed. Platyscytisca Costa and Henry, new genus Type species.—Platyscytisca bergman- nae, new species. Insecta, Heteroptera, Miridae, Phylinae, Platyscytisca, new genus, berg- Diagnosis.—This new genus is distin- guished from other phyline mirids by the combination of the overall pale coloration, banded second antennal segment, broad head with the concave vertex, pale hemel- ytra with a small, round, dark spot on the cuneus and another on the membrane just distal to the large areole, cluster of four spines on the male genital capsule, and by the long, slender vesica, with a very slen- der, sharply bent, apical process. Description.—Small, delicate phyline, overall coloration pale or whitish. Head im- punctate, much broader than long, convex anteriorly in dorsal aspect, strongly pro- duced ventrally below eyes a distance slightly greater than the lateral height of an eye, vertex wide, concave, wider than com- bined dorsal widths of eyes. Rostrum slen- der, extending to metacoxae or beyond. An- tenna relatively slender; segments I and II subequal in diameter; segments HII and IV more slender; segment II longest, with two VOLUME 101, NUMBER 3 Figs. 1-3. claw. dark bands. Pronotum impunctate, much wider than long, posterior width wider than anterior width, lateral margins rounded, basal margin distinctly emarginate; calli weakly delimited laterally and posteriorly by a shallow impressed line. Mesoscutum distinctly swollen transversely and raised well above surface of pronotum; scutellum subequilateral, slightly wider than long, middle of base raised to level of mesoscu- tum, then gradually sloping to level of hem- elytra. Hemelytron impunctate, translucent; cuneus longer than wide with a small dark 49] 50 um 05 mm Playtscytisca bergmannae. 1, Dorsal adult habitus. 2, Antenna (a, male; b, female). 3, Pretarsal spot on basal half; membrane translucent, with two areoles and a small dark spot just beyond large areole and a slender fuscous streak near apex of cuneus. Ventral surface pallid. Legs slender, unmarked; tibial spines slender, pale; claws typically phyline, arolia large, fleshy, extending nearly to apex of each claw. Genital capsule typically round- ed, with a cluster or field of four spines (Figs. 4—6) ventrolaterally on left side, two lateral spines shorter and two, sometimes branched, inner ones longer. Vesica (Fig. 7a) long and slender, apical third more slen- 492 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0,25 mm 50 um Figs. 4—6. Platyscytisca bergmannae. 4, Ventral adult habitus. 5, Male genital capsule showing position of spine cluster. 6, Cluster of spines on male genital capsule enlarged. der, sharply bent, apex with an even more cess having ventral arm of crescent bifid; slender, sharply bent, weakly serrated pro- right paramere simple, rounded (Fig. 9); cess (Fig. 7b); left paramere (Figs. 8a, b) phallotheca (Fig. 10). with a distinct crescent-shaped lateral pro- Etymology.—Platyscytisca is a noun de- VOLUME 101, NUMBER 3 rived from the generic name Platyscytus and the suffix “‘isca,”’ taken from the Anglo Saxon “‘isc,”’ denoting “origin or pertaining to,” to draw attention to the overall simi- larity of it to Platyscytus. The gender is feminine. Remarks.—Platyscytisca appears similar to Amazonophilus Carvalho and Costa and some species of Platyscytus Reuter based on the structure of the head, overall pale coloration, banded antennae, and dark spots on the cuneus and membrane, but very dif- ferent genitalia suggest that this resem- blance simply reflects convergence. The pe- culiar vesica having a slender, abruptly nar- rowed apical process, the cluster of four distinct, apically acute spines on the left la- teroventral area of the male genital capsule, and the crescent-shaped lateral process on the left paramere appear quite unique in the Neotropical mirid fauna. We note that Kerzhner and Schuh (1995) synonymized Amazonophilus under Playts- cytus by stating “‘Judging from the habitus figure and illustrations of male genitalia, bi- punctatus is a species of Playtscytus, and we are so treating it.”’ Although we have not studied the genitalia of the type of the genus, P. binotatus Reuter (nor have the genitalia been illustrated in the literature), we have examined the similar P. blantoni Carvalho (1955), and find that the extreme- ly long, nearly filamentous vesica of Ama- zonophilus bipunctatus Carvalho and Costa, having multiple coils, is quite unlike the relatively short, stout vesica of P. blantoni, having only a single coil. In addition, Car- valho (1955) considered the short, singly coiled vesica of P. tucumanus (Carvalho 1953) of the same generic type as P. bi- notatus. Our observations also indicate that Platyscytus is likely not monophyletic and seems to be made up of at least three spe- cies groups, each of which probably repre- sents a separate genus. Based on this infor- mation, we feel it is premature to consider Amazonophilus a junior synonym of Pla- tyscytus and, therefore, resurrect Amazono- philus, revised status, recognizing that 493 much more work on these seemingly simi- lar taxa is needed. Platyscytisca bergmannae Costa and Henry, new species (Figs. 1-10) Diagnosis.—Platyscytisca bergmannae is best distinguished by the generic characters, particularly by the cluster of four spines on the male genital capsule and the structure of the left paramere and vesica. The com- bination of a dark first antennal segment, two bands on the second antennal segment, and the small round dark spot on each cu- neus and one on the membrane just beyond the large areole (and a narrow fuscous streak just beyond apex of cuneus) will dis- tinguish this species from similar appearing species of Platyscytus. Description.—Male (n = 5): Length 2.80—-3.04 mm, width 0.98-1.16 mm. Head: Dorsal length 0.30—0.32 mm, width 0.62—0.66 mm, vertex 0.32—0.34 mm; uni- formly pale or whitish. Rostrum: Length 0.80—0.84 mm, extending to about meta- coxae. Antenna (Figs. 2a, b): Segment I, length 0.24 mm, dark brown to fuscous, paler at apex; II, 0.88—1.00 mm, pale or whitish, with basal % and a broad band on apical % fuscous; II, 0.30—0.34 mm, pale or white, with basal % fuscous; IV, 0.30— 0.32 mm, pale or white, with basal % fus- cous. Pronotum: Length 0.34—0.36 mm; basal width 0.86—0.92 mm; uniformly pale or whitish. Hemelytron: Uniformly pale or whitish, large portion of clavus, corium, and membrane translucent; a small fuscous spot on basal % of cuneus, and on mem- brane a round fuscous spot just distal to large areole and a fuscous streak just be- yond apex of cuneus. Ventral surface: Uni- formly pale or whitish. Legs: Uniformly pale or whitish; tibial spines small, pale; claws with large fleshy arolia (Fig. 3). Male genitalia: Genital capsule evenly rounded, with field of four prominent spines (Figs. 5, 6); vesica (Figs. 7a, b); left paramere (Figs. 8a, b); right paramere (Fig. 9); phallotheca (Fig.10). 494 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 100 um = <> — Figs. 7-10. Platyscytisca bergmannae. 7, Vesica (a, entire structure, including phallobase; b, Apex showing slender apical process). 8, Left paramere (a, lateral aspect; b, lateral aspect, rotated 180° from Fig. 8a). 9, Right paramere. 10, Phallotheca. Female (n = 7): Length 2.72—0.288 mm, mm; II, 0.78—0.84 mm; III, 0.34—0.36 mm; width 1.12—1.14 mm. Head: Length 0.32— IV, 0.26—-0.30 mm. Pronotum: Length 0.34 mm, width 0.64—0.66 mm, vertex 0.34—0.36 mm, basal width 0.84—0.92 mm. 0.32—0.34 mm. Rostrum: Length 0.82—0.86 Etymology.—This species is named in mm. Antenna: Segment I, length 0.22—0.24 honor of its collector, Dr. Eliana Cherubini VOLUME 101, NUMBER 3 Bergmann (Instituto Biol6gico, Sao Paulo, Brazil). Type specimens.—Holotype <6, Brasil, S. P., Sao Paulo, Instituto Biol6gico, May 1997, E. C. Bergmann coll., taken on Ficus sp. (Museu Nacional, Rio de Janeiro, Bra- sil). Paratypes: 16 6, 32 9, May 1997 & 19 March 1998, same locality and collector as for holotype (Museu Nacional; National Museum of Natural History, Smithsonian Institution, Washington, DC, USA). Remarks.—All specimens have been in alcohol, so the quality of many is poor. ACKNOWLEDGMENTS We are grateful to Eliana Cherubini Bergmann (Instituto Biol6gico, Sao Paulo, Brazil) for sending the specimens used to describe P. bergmannae. We also thank R. C. Froeschner (Department of Entomology, National Museum of Natural History [NMNH], Smithsonian Institution, Wash- ington, DC), D. R. Miller (Systematic En- tomology Laboratory, ARS, USDA [SEL], Beltsville, MD), and D. R. Smith (SEL, % 495 NMNH, Washington, DC) for kindly re- viewing the manuscript. LITERATURE CITED Carvalho, J. C. M. 1953. Neotropical ‘‘Miridae,”’ LVIII: A new genus and species from South America (Hemiptera). Revista Brasileira Biologia 13(1): 33-40. Carvalho, J. C. M. 1955. Neotropical Miridae, LXXVI: Genus *‘Platyscytus”’ Reuter, with key and de- scription of new species (Hemiptera). Revista Brasileira Biologia 15(2): 137-140. Carvalho, J. C. M. 1958. Catalogue of the Miridae of the world. Part III. Orthotylinae. Arquivos Museu Nacional, Rio de Janeiro. 47(3): 1-161. Carvalho, J.C. M. and L. A. A. Costa. 1993. Mirideos neotropicais, CCCLXXVII: Dois géneros e espé- cies de Phylinae (Hemiptera). Anais Academia Brasileira Ciencias 65(2): 203-207. Carvalho, J. C. M. and L. A. A. Costa 1994. Mirideos neotropicais, CCCLXXIV: Géneros e espécies no- vos de Rondonia, Brasil (Hemipera). Revista Brasileira Biologia 54(2): 229-245. Kerzhner, I. M. and R. T. Schuh. 1995. Homonymy, Synonymy, and New Combinations in the Miridae (Heteroptera). American Museum Novitates, Number 3137, 11 pp. Schuh, R. T. 1995. Plant Bugs of the World (Insecta: Heteroptera: Miridae). Systematic Catalog, Distri- butions, Host List, and Bibliography. New York Entomological Society, New York, 1329 pp. PROC. ENTOMOL. SOC. WASH. 101(3), 1999, pp. 496-502 A NEW SPECIES OF DASYHELEA KIEFFER (DIPTERA: CERATOPOGONIDAE) AND NEW RECORDS OF BITING MIDGES FROM THE STATE OF SAN LUIS POTOSI, MEXICO HERON HUERTA AND SERGIO IBANEZ-BERNAL Departamento de Entomologia, Instituto Nacional de Diagnostico y Referencia Epide- miolégicos, Secretaria de Salud. Carpio 470, Col. Santo Tomas, 11340, México, D.EF, Mexico (e-mail: hjh@minervaux.fciencias.unam.mx) Abstract.—Four previously described species of Ceratopogonidae: Lasiohelea anitae (Huerta and Ibanez-Bernal), Forcipomyia (Thyridomyia) nodosa Saunders, Dasyhelea scissurae Macfie, and Culicoides (Haematomyidium) paraensis (Goeldi), are recorded for the first time in the State of San Luis Potosi, Mexico. In addition, descriptions and illus- trations of Dasyhelea huasteca, new species, and the female of Lasiohelea anitae (Huerta and Ibanez-Bernal) are presented. Resumen.—Se registran por primera vez para el estado de San Luis Potostf, México, cuatro especies de Ceratopogonidae previamente descritas: Lasiohelea anitae (Huerta e Ibanez-Bernal), Forcipomyia (Thyridomyia) nodosa Saunders, Dasyhelea scissurae Macfie y Culicoides (Haematomyidium) paraensis (Goeldi). Adicionalmente, se presentan las descripciones e ilustraciones de Dasyhelea huasteca nueva especie y de la hembra de Lasiohelea anitae (Huerta e Ibanez-Bernal). Key Words: Diptera, Ceratopogonidae, Lasiohelea, Forcipomyia, Dasyhelea, Culicoides, Mexico, San Luis Potosi, new species The Ceratopogonidae remain poorly studied in many areas of Mexico, for ex- ample, the Mexican Plateau is a region of current interest for faunistic studies. Some states in this area, such as the states of Mex- ico and Aguascalientes, have no ceratopo- gonid species recorded at present. Near these states is San Luis Potosi, with nine species of biting midges previously record- ed: Forcipomyia incubans (Macfie), F. mexicana Wirth, Culicoides blantoni Var- gas and Wirth, C. eadsi Wirth and Blanton, C. neopulicaris Wirth, Stilobezzia coquil- letti Kieffer, Paryphoconus anomalicornis Kieffer, P. maya Spinelli and Wirth, and Stenoxenus johnsoni Coquillett. We recently studied some specimens of Lasiohelea Kieffer, Forcipomyia Meigen, Dasyhelea Kieffer, and Culicoides Latreille, collected near the towns of San Antonio and San Martin Totolteo in San Luis Potosi, and found new geographical records of four species, as well as the an undescribed spe- cies of Dasyhelea and the previously un- known female of Lasiohelea anitae (Huerta and Ibanez-Bernal). We follow Yu and Wirth (1997) and con- sider Lasiohelea as a genus instead of a subgenus of Forcipomyia. We used _ the slide mounting method suggested by Bor- kent and Bissett (1990), and the morpho- logical terms of Downes and Wirth (1981). All the specimens are deposited in the Col- lection of Arthropods with Medical Impor- VOLUME 101, NUMBER 3 tance of the Instituto Nacional de Diagnos- tico y Referencia Epidemiologicos (IN- DRE), Secretaria de Salud, Mexico. Lasiohelea anitae (Huerta and Ibanez- Bernal) (Figs. 1—8) Forcipomyia (Lasiohelea) anitae Huerta and Ibafez-Bernal 1996: 350, figs. (d, Mexico, Chiapas). Female description.—Head: Eyes bare, mesally with ocular margins in contact, but with marginal facets separated at narrowest distance by one to one and a half facet di- ameters (Fig. 3). Flagellum brown (Fig. 1); lengths of flagellomeres (wm): 30-20-22- 22-22-23-24-27-63-68-69-70-98; antennal ratio (AR): 1.76 (1.70-—1.82; n = 3); basal flagellomeres semispherical, flagellomere 8 with 6—7 basiconica sensilla. Palpus (Fig. 2) with lengths of segments (wm): 0.3-42-22- 35; palpal ratio (PR): 1.1 (1.0-1.3, n = 3); third segment swollen at midlength, with a large oval pit, containing several small, ir- regularly arranged sensilla capitata. Man- dible (Fig. 4) with 30—32 small teeth; man- dible length (um): 108.5, width (wm): 17.6. Cibarial armature with 15 or 16 teeth in sin- gle row (Fig. 5); each tooth with basal apo- deme, each about double tooth length. Thorax: Scutum, scutellum brown; legs uniformly pale yellowish. Tarsal ratios of foreleg (1), midleg (II), hindleg (III) (TR): ee So 362.42), MM: 2.02.0), TM: 2:0 (2.0—2.1) (n = 3); hind tibial comb with 7 spines, one near spur longest. Wing (Fig. 6) with radial cells coalesced, slitlike; wing length: 0.83 mm (0.82—0.85; n = 3), width: 0.34 mm (0.32—0.35; n = 3), costal ratio (CR): 0.59 (0.59—0.60; n = 3). Halter pale. Abdomen: Brown. Genitalia (Fig. 8) with 3 small lobes arising at different level from posterior base of genital fork; sper- matheca (Fig. 7) partially collapsed in ex- amined material. Distribution.—Mexico (Chiapas, San Luis Potosi). Specimens examined.—3 @, 1 6. Mex- 497 ico: San Luis Potosi, San Antonio, El Pu- ente, April 23, 1997, Malaise trap, Paz- Rodriguez, R. & Pérez-Renteria, C., cols. Comments.—The female of this species can be associated with the male by the number and arrangement of the cibarial teeth, the form of the palpus and the frontal sclerite, the body coloration, and the ar- rangement and form of the sensilla on the flagellomeres. The single male collected from San Luis Potosi was compared with the type specimens previously described by Huerta and Ibaénez-Bernal (1996). Unfortunately, the descriptions of fe- males of most American species of Lasioh- elea do not include many important char- acteristics useful for their separation, such as the number of cibarial and mandibular teeth, and certain ratios, thereby making species determination very difficult. We be- lieve it is necessary to redescribe most of the American species in order to recognize the important characteristics useful for sep- arating these species (Yu and Wirth 1997). Forcipomyia (Thyridomyia) nodosa Saunders (Fig. 9) Forcipomyia (Thyridomyia) nodosa Saun- ders 1959: 43 (all stages, Costa Rica, figs.). Several specimens of this species were collected in San Luis Potosi. An illustration of the male genitalia is presented in Fig. 9. Dow and Wirth (1972) gave detailed rede- scriptions of both sexes. Distribution.—U.S.A.; Mexico (Baja California, Sonora, Sinaloa, San Luis Po- tosi); Costa Rica; Colombia. Specimens examined.—8 ¢, 6 2. Mex- ico: San Luis Potosi, San Antonio, El Pu- ente, April 23, 1997, Malaise trap, Paz- Rodriguez, R. & Pérez-Renteria, C., cols. Dasyhelea scissurae Macfie (Fig. 10) Dasyhelea scissurae Macfie 1937: 15 (6, Trinidad, figs.). 498 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON aE waar 9d) 11 G4} a Figs. 1-8. Lasiohelea anitae, female. 1, Flagellomeres. 2, Palpus. 3, Head. 4, Mandible. 5, Cibarial armature. 6, Anterior veins of wing. 7, Spermatheca. 8, Abdomen, distal segments, in ventral view. Scale lines in milli- meters. VOLUME 101, NUMBER 3 499 0.05 Figs. 9-10. Male genitalia, in ventral view. 9, Forcipomyia (Thyridomyia) nodosa. 10, Dasyhelea scissurae. Scale line in millimeters. his as the first record? of this species from the state of San Luis Potosi, and only the second record from Mexico (Ibaénez- Bernal et al. 1996). We provide an illustra- tion of the male genitalia (Fig. 10). Distribution.—Mexico (San Luis Potosi, Guerrero); Costa Rica; Bermuda; Trinidad: Argentina. Specimens examined.—17 6, 11 @. Mexico: San Luis Potosi, San Martin To- tolteo, April 24, 1997, Malaise trap, Paz- Rodriguez, R., Pérez-Renteria, C., cols. Seven males mounted on slides; the re- maining specimens preserved in ethanol. Dasyhelea huasteca Huerta and Ibanez- Bernal, new species (Figs. 11—16) Diagnosis.—Small brown species, with the fifth tarsomeres dark and female cerci spatula-like. Female.—Head (Fig. 13): Dark brown. Eyes with short pubescence, separated at narrowest distance by 5 pm. Frontal sclerite with two small, inclined, sclerotized bars. Flagellum (Fig. 11) brown; antennal ratio (AR): 0.72; with more or less cylindrical non-reticulate flagellomeres, with lengths (um): 37-30-30-32-32-35-35-35-35-35-37- 37-46. Palpus (Fig. 12) yellowish; lengths of segments (jm): 27-42-22-35; palpal ratio (PR): 2.2; third segment with a few sensilla capitata on mid portion of mesal surface. Thorax: Scutum and postscutellum dark brown; scutellum yellowish. Legs pale yel- lowish, apex of femora, base of tibia dark- ish, fifth tarsomeres dark; hind tibial comb with four spines, one nearest spur, short; hind tarsal ratio 1.9. Wing (Fig. 14) length: 0.74 mm; width: 0.31 mm. Wing membrane with macrotrichia extending from near base of vein r-m to wing margin; cell r, narrow, nearly twice length of cell r, closed; vein r- m oblique. Costal ratio (CR): 0.50. Halter with brown stem, white knob. Abdomen (Fig. 15): Brown. Genitalia (Fig. 16) lightly sclerotized arched subgenital plate; cerci very long, with spatula-like cover on mesal sides of ventral surfaces, cercus length 98.4 wm. Spermatheca subspherical (Fig. 15) measuring 0.04 mm by 0.031 mm, and a second rudimentary spermatheca pre- sent. Spermathecal/cercus length: 0.44. Male.—Unknown. 500 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 11-16. Dasyhelea huasteca, female. 11, Flagellomeres. 12, Palpus. 13, Head. 14, Wing. 15, Abdomen showing the two spermathecae, in ventral view. 16, Abdomen, distal segments, in ventral view. Scale lines in millimeters. Types.—Holotype 2, 1 ¢ Paratype: San Antonio, El Puente, April 23, 1997, Malaise Trap, Paz-Rodriguez, R., and Pérez-Renteria, C., cols. Both specimens deposited in the Collection of Arthropods with Medical Im- portance of the Instituto Nacional of Diag- nostico and Referencia Epidemiologicos (IN- DRE). VOLUME 101, NUMBER 3 Etymology.—The name huasteca refers to the natural region of Mexico that extends from the maritime slope of the Sierra Madre Oriental to the Gulf of Mexico, between the river Cazones and the Tamesi basin. The region includes the southern part of the state of Tamaulipas, the northern area of the state of Veracruz, the oriental strip of the state of San Luis Potosi, in which the spec- imens were collected, and a small sector of the states of Puebla and Hidalgo. The re- gion was inhabited in the past by the huas- tecas, a pre-Columbian Mexican people of the Maya-quiche group, now restricted to two small areas inside the region. Comments.—Only two species of Dasy- helea have modified spatula-like cerci: D. spathicerca Wirth and D. huasteca. How- ever, D. spathicerca differs from this new species because it has the two well devel- oped spermathecae, wing is larger (wing length 1.11 mm), has a greater antennal ra- tio (0.68), the cerci are thinner and shorter, and the ratio of spermathecal length/cercus length is only 0.36. It is very difficult to place this species in the subgenera proposed by Remm (1962, 1979) or in the species-groups proposed by Wirth (1952) and Waugh and Wirth (1976) because of the considerably modified fe- male cerci and subgenital plate. The flagel- lomeres are more or less similar to other species in the Leptobranchia group of Waugh and Wirth (1976), but the third seg- ment is shorter than the combination of fourth and fifth palpal segments. Therefore, we can not accurately place D. huasteca in any of the species-groups or subgenera pro- posed to date, and its true affinities are un- known. Culicoides (Haematomyidium) eadsi Wirth and Blanton Culicoides eadsi Wirth and Blanton 1971: S(O. ©. Lexas): This species was previously reported for this region by Wirth and Blanton (1971). Distribution.—U.S.A. (Texas, Florida); 501 Mexico (Sonora, Nayarit, San Luis Potosi, Yucatan); Cuba. Specimens examined.—1 2. Mexico, San Luis Potosi, San Antonio, El Puente, April 23, 1997, Malaise trap, Paz-Rodri- guez, R., Pérez-Renteria, C., cols. Culicoides (Haematomyidium) paraensis (Goeldi) Haematomyidium paraensis Goeldi 1905: 137 ¢2.2Brazil), Culicoides undecimpunctatus Kieffer 1917: 307 (2, Argentina). This is the first record of this species from the state of San Luis Potosi. Distribution.—Eastern U.S.A.; Mexico; Central and South America south to Argen- tina; Barbados, Grenada and Trinidad in the West Indies. Specimen examined.—1 ¢. Mexico, San Luis Potosi, San Antonio, El Puente, April 23, 1997, Malaise trap, Paz-Rodriguez, R.., Pérez-Renteria, C., cols. ACKNOWLEDGMENTS We gratefully acknowledge Rafael Paz- Rodriguez and Crescencio Pérez-Renteria for the collection of the specimens studied. We appreciate the revision and useful cor- rections made to the manuscript first by our friend Larry Hribar, and later by Art Bor- kent and William L. Grogan, which strik- ingly contribute to its improvement. LITERATURE CITED Borkent, A. and B. Bissett. 1990. A revision of the Holarctic species of Serromyia Meigen (Diptera: Ceratopogonidae). Systematic Entomology 15: 153-217. Cavalieri, EF 1962. Notas sobre Ceratopogonidae (Dipt. Nematocera) III. Sobre un nuevo diptero hema- tofago para Argentina, Lasiohelea saltensis n. sp. y notas sobre las especies neotropicales de La- siohelea. Acta Zoologica Lilloana 18: 359-365. Dow, M. I. and W. W. Wirth. 1972. Studies on the genus Forcipomyia, 2. The Nearctic species of the subgenera Thyridomyia and Synthyridomyia (Dip- tera: Ceratopogonidae). Annals of the Entomolog- ical Society of America 65: 177-201. Downes, J. A. and W. W. Wirth. 1981. Chapter 28, Ceratopogonidae, pp. 393—421 Jn Manual of Ne- 502 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON arctic Diptera, Vol. 1. Agriculture Canada Re- search Branch Monograph No. 27. 674 pp. Goeldi, E. 1905. Os Mosquitos no Para. Memorias do Museu Goeldi (Museo Paraense) de Historia Nat- ural e Ethnographie 4: 1—154. Kieffer, J. J. 1917. Chironomides d’ Amérique con- servés au Musée National Hongrois de Budapest. Annales Historico-Naturales Musei Nationalis Hungarici 15: 292-364. Huerta, H. and Ibanez-Bernal, S. 1996. Especie nueva de Forcipomyia (Lasiohelea) de Chiapas, México (Diptera:Ceratopogonidae). Anales del Instituto de Biologia Universidad. Nacional Autonoma de México, Serie Zoologia 67: 349-355. Ibanez-Bernal, S., W. W. Wirth, and H. Huerta. 1996. Chapter 36, Ceratopogonidae (Diptera), pp. 567— 577 In Llorente-Bousquets, J., A. N. Garcta-Al- drete, and E. Gonzalez-Soriano, eds., Biodiversi- dad, taxonomia y biogeografia de artro6podos de México: Hacia una sintesis de su conocimiento. CONABIO-IBUNAM, México, 660 pp. Macfie, J. W. S. 1937. Ceratopogonidae from Trinidad. Annals and Magazine of Natural History, Series 10, 20: 1-18. Remm, H. 1962. Genus Dasyhelea Kieffer v Estonii (Diptera: Ceratopogonidae). Tartu Riikliku Uli- kodi Toimetised 120: 108-133. 1979. Eesti NSV_habesaasklaste (Diptera, Ceratopogonidae) fauna kataloog [in Estonian, with Russian and English summary] pp. 40—60 Jn Dipteroloogilisi Uurimusi (Tartu). Eesti NSV Teaduste Akadeemia Eesti Looduseuurijate selts. Saunders, L. G. 1959. Methods for studying Forcipo- myia midges, with special reference to cacao-pol- linating species (Diptera, Ceratopogonidae). Ca- nadian Journal of Zoology 37: 33-51. Waugh, W. T. and W. W. Wirth. 1976. A revision of the genus Dasyhelea Kieffer of the eastern United States North of Florida (Diptera: Ceratopogoni- dae). Annals of the Entomological Society of America 69: 219-247. Wirth, W. W. 1952. The Heleidae of California. Uni- versity of California Publications in Entomology 9: 95-266. 1956. New species and records of biting midges ectoparasitic on insects (Diptera, Helei- dae). Annals of the Entomological Society of America 49: 356-364. Wirth, W. W. and E S. Blanton. 1971. New Neotropical sandflies of the Culicoides debilipalpis group (Diptera: Ceratopogonidae). Proceedings of the Entomological Society of Washington 73: 34—43. Yu Y. X. and W. W. Wirth. 1997. Lasiohelea of South- east Asia (Diptera: Ceratopogonidae). Military Medical Science Press, XIV+ 89 pp. PROC. ENTOMOL. SOC. WASH. 101(3), 1999, pp. 503-513 WHAT IS THE REAL I[SWAROIDES (HYMENOPTERA: TIPHIIDAE: THYNNINAE)? LYNN S. KIMSEY Bohart Museum of Entomology, Department of Entomology, University of California, Davis, CA 95616, U.S.A. (e-mail: Iskimsey @ucdavis.edu). Abstract.—Phylogenetic relationships among /swaroides Ashmead, Thynnoturneria Rohwer, Acanthothynnus Turner, Aspidothynnus Turner, and Epactiothynnus Turner are reexamined. As a result of this analysis, synonymy of Thynnoturneria under [swaroides is confirmed. /swaroides is redescribed and species placements corrected for this revised generic grouping. Twenty-three species are placed in /swaroides, and the new species Iswaroides robusta is described. Key Words: Tiphiidae, Hymenoptera, /swaroides, Thynnoturneria, Australia, Acanthoth- ynnus, Aspidothynnus, Epactiothynnus Several generic groupings of Australian thynnine tiphiids are part of the confusion over the identity of Aeolothynnus of Ash- mead (1903) versus the concept of Aeo- lothynnus used by Turner (1908). As dis- cussed in detail by Kimsey (1999) Turn- er’s concept of this genus actually was not congeneric with Ashmead’s, and his des- ignation of A. cerceroides as the type of Aeolothynnus included quite a different group of species. Unfortunately, some time earlier Ashmead (1899) had de- scribed the genus /swaroides, based on the species J. koebelei. Turner never saw the type of /swaroides and simply re-used Ashmead’s description in the Genera In- sectorum review of the subfamily Thyn- ninae (Turner 1910c). Although these two type species are very different one from the other, there are a number of seemingly intermediate species, including one de- scribed below, and the distinctions be- tween the two groups have never been clearly resolved. The difficulty with these generic group- ings has several origins. A large part of the confusion comes from the nomenclatural chaos created by Turner, Ashmead, and Rohwer. This confusion is in turn com- pounded by our incomplete knowledge of the species diversity. A six-week collecting trip to Western Australia yielded twenty species of Iswaroides/Thynnoturneria, yet only five species have been described from this part of Australia. Based on examination of museum collections there are very few widespread species in this group. ““Thyn- noturneria’ cerceroides appears to occur in Western Australia, South Australia, and Northern Territory. However, this entity is actually a cluster of at least three species, all closely resembling one another in col- oration, gross morphology, and a peculiarly indented forefemur. However, each of these appears to have a discrete allopatric distri- bution, with one in SA, one in NT and one in WA. Thorough phylogenetic analysis of all available species in these groups has re- vealed that there is no robust support for two genera as represented by /swaroides Ashmead and Thynnoturneria Rohwer. 504 MATERIALS AND METHODS Specimens were studied in situ or were borrowed from the following: The Natural History Museum, London, S. Lewis; Hope Museum, Oxford University, C. O’Toole; Australian National Insect Collection, CSI- RO, Canberra, ACT, I. Naumann and J. Cardale (CANBERRA); Bohart Museum of Entomology, University of California, Da- vis, S. L. Heydon (DAVIS), Naturhistoris- che Museum, Vienna, Austria, M. Fisher (VIENNA), and Western Australian Muse- um, Perth, Terry Houston (PERTH). Type repositories for the new species described below are indicated by the city name in capital letters in parentheses. Males of twenty-five putative species were analyzed for this study, including the new species /. robustus described below, and representatives of three other genera, Acanthothynnus sannae Turner, Aspidoth- ynnus fossulatus Turner, and Epactiothyn- nus opaceiventris Turner. An asterisk (*) in- dicates primary types examined for this study. Fifteen of the species given in the data matrix are morphospecies assigned by a letter name, because there does not appear to be an available species name. Nearly all of the types of species placed in these genera are in The Natural History Museum, London. It has not been possible to restudy these types in light of the re- evaluation of /swaroides and Thynnoturner- ia. Ordinarily this paper would not have been published without correct names or new species names being assigned to these taxa. However, there is considerable need to stabilized the generic framework in this speciose subfamily of wasps because of their roles both in the biodiversity of the Australian continent and their probable im- portance in population control of soil- dwelling scarab beetles, including imported dung beetles. Phylogenetic analyses were conducted using the Hennig86 software (Ferris 1988), using branch swapping (mhennig and bb commands), followed by successive weight- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ing and generation of a Nelson consensus tree (Fig. 1): PHYLOGENETIC ANALYSIS Character States 1. Labrum small, apical plate foreshort- ened, with row of subapical setae and narrow basal attachment (Fig. 4)(0); la- brum large highly sclerotized and ase- tose except apical fringe, broadly at- tached, apex broadly rounded (Figs. 2, 3) (CD): 2. Clypeal apex as broad or broader than medial eye width, apical margin broad- ly truncate (Figs. 2, 3)(0); apex narrow, often medially rounded, narrower than eye width (Fig. 4)(1). 3. Stipal fringe of setae long and dense, continuous along most of inner margin (Figs. 5, 6)(0); stipal fringe reduced to less than half of inner margin (1). 4. Stipes unmodified, stipal fringe occu- pying more than half of stipes (Fig. 5)(O); stipes shortened and posteriorly twisted, posterior half strongly cupped (Fig. 6)(1). 5. Prementum broad and _parallel-sided (Figs. 6, 9)(0); prementum elongate (Fig. 5), often narrowed anteriorly (1). 6. Face flat or slightly concave between antennal socket and eye margin (0); area between antennal socket and eye longitudinally sunken and often pol- ished and impunctate (1). 7. Occipital fossa and oral fossa narrowly separated by a ridge (Figs. 5, 7, 9)(O); broadly separated by flat or concave polished area (Fig. 6)(1). 8. Occipital carina meeting oral carina at an acute angle (Fig. 6)(0); reaching oral carina at an obtuse or right angle (Figs. ie 9) oy 9. Vertex with red spot between hindocelli and upper eye margin (Fig. 8)(0); with- out red spot (1). 10. Pronotum strongly narrowed medially, depressed subapically (0); pronotum elongate medially and flat without transverse subapical depression (1). VOLUME 101, NUMBER 3 505 Table 1. Character matrix for genera and species related to /swaroides. Taxon Character | 2 3 4 ) 6 7 8 9 10 11 12 13 14 15 16 17 18 Epactiothynnus! Oe Oe Oe One OM OR s One Om OpneOw 0) O%.:O% 40: 207) 108 7 O49 10 Acanthothynnus? ik "OO | OF VOF O00 | Os OrO ae (0) § 10) l Aspidothynnus* OPO) +0 l I OR On ON Oe OFS 0. 0" OF OLaO Iswaroides baccata Oo @ @© @ l 0) | | | O | | 0 | 0) I. cerceroides | OF OM OF 0 0s 0 | 0) | 0) | Oe) ( | 0) I. illustris O | 0) ] (0) (0) | O | OO x0) | oO 0 Ome (0) I. koebelei O | 0) | O O l O | l O | 0) | | | OnNesO I. robustus 0) l OP) | OOO ] OO | O O | | | 0) I. sanguinulentus OF FO) Oe a l 0 l 0 l 1 l 0) l l 0 | 0) I. xerophila O I 0) l O O | O | | O | ( 3@ | | OF 0 I. species A 0 | | 1 OF 0) | 0) | 0) | I | | | | 0-0 J. species B O | 0) | l 0) | O | OF 0 | 0) | | | 0 0 I. species C | OF, OF 20 | 0) | 0) | O | | | | | 0) I I. species D 0) | | | O O l OF 0) 10), 30 I | 0) | | OF 0 I. species E O l 0) | Ov 0 | Oe sO 0) | OFF 0 | l oO” 0 Especiessk | O05 0 0 | 0) | O | O | I | | l | 0) I. species G ] OR OR OREO. | 0) | O I 0) l | | | | | 0) I. species H | ORTLOP 320 | | O | 0) | | | O | | | O ] I. species I On LOre nO | | O | O | 0) | O | | O | 0 I. species J O | | | Oe) | O | O O l On | One O'e 0 I. species K 0) | 0) | OF 0 | 0) | Oh | O O | | OF SiO I. species L l OO} .@ 1 | 0) | 0) | 0) | l | | | 0) | I. species M | Onn OF | l 0) l 0) | 0) | OF 0 | | 0) | I. species N 0) | 0) | OF 20 | Op LOROF BO | OP 10) | | OFF0 I. species P 0) | | | ORO | De Oe 0) 0) l O O | | O10) ! Epactiothynnus opaceiventris Turner. 2 Acanthothynnus sannae Turner. ’ Aspidothynnus fossulatus Turner. . Mesopleuron anteriorly rounded (0); anterior declivity margined by a verti- cal omaulus (1). . Mesopleuron with transverse scrobal groove extending anteriorly from scrobe (0); mesopleuron evenly convex without trace of scrobal groove (1). . Gastral tergum V evenly rounded sub- apically (Fig. 16)(0); subapical margin produced into an acute shelf-like ridge (Eiger) Cl): . Gastral sternum V evenly rounded lat- erally (OQ); with lateral tooth (1). . Gastral sternum VI evenly rounded lat- erally (O); with lateral tooth (Figs. 15, 16)(1). . Volsella convex (0); concave at least on apical half and U-shaped in cross-sec- tion with digitus obscured (Fig. 22)(1). . Gonocoxal dorsal apex deeply emar- ginate medially appearing strongly bi- lobate (Figs. 18-20) or narrowly uni- lobate (Fig. 24)(0); apex convex me- dially, appearing trilobate (Fig. 17)(1). 18. Gonocoxal dorsal apex deeply emar- ginate medially appearing strongly bi- lobate or narrowly unilobate (QO); apex broadly truncate (Fig. 23)(1). RESULTS Representatives of the closely related genera Acanthothynnus Turner, Aspidoth- ynnus Turner, and Epactiothynnus Turner were used in the analysis as outgroups to polarize the character states. These taxa all belong to the Jswaroides group of genera characterized in the males by having a par- tial or complete transverse carina or ridge at or near the apical margin of the epipy- gium. Males also have well-developed pe- 506 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1—5—-6 151018 LJ Synapomorphy C) Reversals 13 Homoplasy Pisaele dothynnus, and Acanthothynnus. nis valves and a tridentate hypopygium. Fe- males are characterized by the apical gastral tergum having a smooth, broadly or nar- rowly ovoid medial plate enclosed at least laterally by a carina and subtended medially by a long dense tuft of setae. The character matrix, given in Table 1, was analyzed using intensive branch swap- ping and successive weighting, which yielded ten trees. A Nelson Consensus Tree generated from this last set of trees yielded the tree given in Fig. 1. The ci of the con- sensus tree was 74, with a retention index of 94. Most of the trees generated came from instability within the apical clades. The intent of this study was not to resolve species level relationships but rather to ex- amine the stability of basal clades and to Epactiothynnus Aspidothynnus Acanthothynnus 14—— species-G a bs species-F baccata — cerceroides Act: a a sanguinolentus A= 6-[8]-1 9) - species-l ——_—_————_- species-H species-M {16} 5-18 species-C species-L robustus species-B illustris species-E species-K species-N 14—— keobelei ll Mead ae Agy— species-J (Q)y— species-P 11-14- species-A ‘onlays species-D* xerophila Cladogram showing phylogenetic relationships among species of /swaroides, Epactiothynnus, Aspi- see if currently accepted generic groupings could be confirmed or repudiated. The results of this study are twofold. First, there is litthe support for retaining Thynnoturneria and Iswaroides as separate entities. Although the types of each genus sort out consistently into two clades the basal rooting of these clades is weak, and they are not sufficiently well supported to justify their division into two genera. These two groups are referred to below as the koe- belei and cerceroides clades. Second, the genus Epactiothynnus exhibited the ances- tral condition in all of the characters ex- amined for this study. This genus must be reexamined to determine if it is character- ized by any apomorphies. If not, then the stability of all of the genera characterized VOLUME 101, NUMBER 3 by the traits discussed above is in question. Given the results of this analysis, the genus Thynnoturneria Rohwer is synonymized under Jswaroides Ashmead as the junior name. The entire group, as /swaroides, is rediagnosed below. The cerceroides clade contains the type species of Thynnoturneria (T. cerceroides) as well as, 7. baccata, T. sanguinolentus andspecies C, EG, H, I, L, and M. The apomorphy supporting this clade is the oc- cipital carina reaching the oral carina at an acute angle. Other characters that distin- guish most members of this clade include the large shield-like labrum, polished and sunken area between the eye and the anten- nal socket and the dorsally flat and medially elongate pronotum. These latter features are also shared with Aspidothynnus and Acan- thothynnus species. The second clade is distinguished by one apomorphy—the strongly narrowed clypeal apex, as well as the loss of a red spot on the vertex in most, but not all species. Clade 2 contains [. koebelei (the type spe- cies of [swaroides) I. robustus, T. illustris, i *