ie if MIN ie iy UN AM ae ne we a it i any : UN 2 n Ae, Oh iy A ‘hi ie ; OLE | (Omar) ¢ ie ant a Y ) A ri , eb iit i i , | . hy rs a i VOL. 108 JANUARY 2006 NO. 1 p (ISSN 0013-8797) (aN 46) e64X PROCEEDINGS SA ied of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED QUARTERLY ascm > apr a ly JAN :1.8 2006 3 LIRRARIES BICHA, WESLEY—New scorpionflies (Mecoptera: Panorpidae) from Jalisco, Michoacan;~and OAC WV ICKT CO ee cas te INE. , SOs Haas SE Ev ad renee Meteo n toy AM Ws e mat Os fied aot danse aa bats tees aon ap 24 CONTENTS BOUCHER, STEPHANIE and PAUL HANSON—A new Costa Rican species of Japanagromyza (Diptera: Agromyzidae) forming galls on Lonchocarpus (Fabaceae) ............ 9 CALMASUR, ONDER and HIKMET OZBEK—A willow sawfly, Nematus salicis (Linnaeus) (Hymenoptera: Tenthredinidae), a new record and new pest of Salix spp. in Turkey .............. 139 CARROLL, J. FE, V. B. SOLBERG, and T. L. CYR—Development of a bioassay for evaluating tick (Acari xocidas)rattrackants i db stite lela tite tere seek et ae eee raceei nha easdessoae Schade ceeaet scat stacns 82 CERVANTES PEREDO, LUIS and SAGRARIO GAMEZ VIRUES—Lethaeini (Hemiptera: Lygaeoidea: Rhyparochromidae) associated with figs in Mexico, with the description of a BOWE SPECIES Ol CUSEMIIA aoe OF veclas tesdatts. ok N ak ad bares Buenas copoly eal cs goenngesns spec Met Mnobte taal ee Mesto tae sa 101 CHOI, WON-YOUNG and JAMES B. WHITFIELD—Cuwneogaster, a new genus of the subfam- ily Microgastrinae (Hymenoptera: Braconidae) From the Neotropical Region ................:045 119 DARSIE, RICHARD F.,, JR—Description of the pupae of three species of the genus Luizia, a com- parison of New and Old World pupae, and a key to pupae and larvae of the genus ( Diptera: ROTI IGI IE) ereeee ce ete ee ek EL et VEAL RNIN St yelp WOLSEY CIE ls Ngan dad os Mesa a Nant sa aR Acs nega dei 145 FERRO, MICHAEL L. and ROBERT W. SITES—Description of the larva of Gomphidictinus perakensis (Laidlaw) (Odonata: Gomphidae), with distributional notes ...........:::::::ceeeeeeeee 76 HARBACH, RALPH E., EMMA PARKIN, BIN CHEN, and ROGER K. BUTLIN—Anopheles (Cellia) minimus Theobald (Diptera: Culicidae): Neotype designation, characterization, and SV SECITIALIGS hiteee secrets ee fae P nC 2 UG TENET URL Ate gal vauis Soca totes eter rvs ctetedtnssncges dees seeits=cuawpesgeanaeatas 198 HOLMAN, JAROSLAV, SEUNGHWAN LEE, and JAN HAVELKA—A revision of the genus Macrosiphoniella del Guercio (Hemiptera: Aphididae) from the Korean Peninsula, Part I: Subgenera Asterobium, Chosoniella, Papillomyzus, Phalangomyzus, Sinosiphoniella ............ 174 HOLSTON, K. C. and C. R. NELSON—A quantitative evaluation of intraspecific and interspe- cific variation in eye morphology for Efferia Coquillett (Diptera: Asilidae) species .............. 210 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON e ORGANIZED Marcu 12, 1884 OFFICERS FOR 2006 STEVEN W. LINGAFELTER, President Jon A. Lewis, Custodian MIcHAEL W. Gates, President-Elect MIcHAEL G. PoGur, Treasurer Stuart H. McKamey, Recording Secretary | JOHN W. BRowN AND Davip G. Furtu, Program Chairs Ho tuts B. WILLIAMS, Membership Secretary JASON P. W. HALL, Past President Davip R. Smitn, Editor Publications Committee RAYMOND J. GAGNE Tuomas J. HENRY Wayne N. Martuis Honorary President DoNALD M. ANDERSON Honorary Members WILLIAM E. BICKLEY Davip R. SMITH RONALD W. HopGEs Manya B. STOETZEL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168. MEETINGS.— Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu- tion, on the first Thursday of each month from October to June, inclusive, at 7:00 P.M. Minutes of meetings are published regularly in the Proceedings. 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. 739-740 OF THE JULY ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 5 January 2006 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 1-8 GAMPSOCORIS DECORUS (UHLER) AND METACANTHUS TENELLUS STAL (HEMIPTERA: BERYTIDAE): NEOTROPICAL STILT BUGS AS COLONISTS OF AN AFRICAN GRASS, UROCHLOA MUTICA (POACEAE), IN FLORIDA, WITH A REVIEW OF BERYTID-GRASS ASSOCIATIONS A. G. WHEELER, JR. AND THOMAS J. HENRY (AGW) Department of Entomology, Soils, and Plant Sciences, Clemson University, Clemson, SC 29634-0315, U.S.A. (e-mail: awhlr@clemson.edu); (TJH) Systematic En- tomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agricul- ture, % National Museum of Natural History, Smithsonian Institution, PO. Box 37012, MRC-168, Washington, DC 20013-7012 (e-mail: thenry @sel.barc.usda.gov) Abstract.—Host plants of the Neotropical stilt bugs Gampsocoris decorus (Uhler) (sub- family Gampsocorinae) and Metacanthus tenellus Stal (subfamily Metacanthinae), whose U.S. distributions are limited to southern Texas and peninsular Florida, have been un- known or little known. In Florida, both berytids have colonized Para grass (Urochloa mutica (Forssk.) T. Q. Nguyen; Poaceae), an invasive plant native to Africa. We collected G. decorus on Para grass in 20 counties and M. tenellus in 17 counties. Para grass is the first host plant recorded for G. decorus and first grass documented as a host for M. tenellus. Grass-feeding habits have evolved in three berytid clades and have been demonstrated for certain gampsocorines and metacanthines; grass feeding needs to be verified for bery- tine stilt bugs. Key Words: Lygaeoidea, Berytidae, Gampsocorinae, Metacanthinae, Para grass, host plants, new records, grass feeding Gampsocoris decorus (Uhler) and Meta- canthus tenellus Stal are mainly Neotropi- cal berytids recorded previously in the United States from Florida and Texas (Hen- ry 1997a, Henry and Froeschner 1998). Host plants previously were unknown for G. decorus, and nymphs of M. tenellus were reported only from a euphorbiaceous plant in Bermuda (Henry and Hilburn 1990). Here we give additional Florida re- cords of both stilt bugs and discuss their colonization of Para grass, Urochloa mutica (Forssk.) T. Q. Nguyen, which is native to Africa. We also review the worldwide use of grasses as hosts of the Berytidae. METHODS After we first found G. decorus and M. tenellus on Para grass in Florida in May 2003, we surveyed the plant to determine the prevalence of this host association and better understand the Florida range of both berytids. Para grass was sampled mainly along highways but also along canals and rivers and in marshes. We collected stilt bugs by beating the stems of Para grass over a shallow, short-handled net or white enamel pan and aspirating or hand picking with plastic vials the dislodged adults. When fifth instars, but not adults, were ob- served, we collected one or more nymphs and held them on a sprig of Para grass in a small plastic box to obtain adults. Speci- mens from each locality are deposited in the National Museum of Natural History, Smithsonian Institution, Washington, DC. Specimens from May 2003 were collect- 2, Va — % f " .— y iS ; " o a a * a % 4 > . 8 4 y 4 f a fc — ae a QZ Fig. 5. HAMILTON w=. = fnassa m \ pr + 2 \ ¢ Distribution of stilt bugs in Florida: @ = records of Gampsocoris decorus only; O = records of Metacanthus tenellus only; Hf = records of G. decorus and M. tenellus collected together. clude Parad grass in his manual of south- eastern plants, he listed it from Florida un- der the synonymic name Panicum barbi- node Trin. Apparently before the plant had become invasive in Florida, it was recom- mended for pasturage or hay (Thompson 1919). It was planted to camouflage certain military installations in southeastern Florida during World War II, and might have been introduced later (Austin 1978). Urochloa mutica is the first host plant known for G. decorus and the first grass VOLUME 108, NUMBER 1 shown to serve as a host for M. tenellus. Nymphs of M. tenellus previously were known only from a euphorbiaceous plant; adults collected from grasses, including U. mutica (as Panicum barbinode) in Puerto Rico, therefore, were assumed to represent accidental or resting records (Henry 1997a). Metacanthus tenellus can be added to the extensive list of insects associated with Para grass in Puerto Rico (Martorell 1976). Both berytids were first reported from Florida after Para grass was introduced into the state. Either or both species, however, could have been present in Florida prior to the introduction of the grass, living on one or more native grasses. Alternatively, the bugs could have dispersed to southern Flor- ida from the West Indies (both are known from Cuba) and colonized an already-nat- uralized U. mutica. In either case, natural- ization of U. mutica, a pubescent grass (Langeland and Craddock Burks 1998) sim- ilar to many other hosts favored by bery- tids-that is, hairy and/or viscid (Wheeler and Schaefer1982)—would have provided a suitable host and might have facilitated northward spread of the bugs in Florida. Neither berytid has been observed to de- velop on grasses native to Florida. Phy- tophagous insects sometimes become so specialized on novel plants that they no lon- ger can be found on their original hosts (e.g., Valley and Wheeler 1976, Thompson 1994). Grass feeding has developed in at least three clades of the Berytidae (Henry 1997b, fig. 3). Certain other narrow-elongate het- eropterans also feed on grasses, for exam- ple, leptocorisine and micrelytrine alydids, chorosomine rhopalids, and stenodemine mirids (Wheeler and Schaefer 1982, Wheel- er 2001). The only berytid subfamily for which grasses might not be used for nymphal development by at least one spe- cies is the Berytinae. Although several ber- ytinine species have been collected from grasses, the use of grasses as host plants by Berytinus hirticornis (Brullé) and several other species of the genus requires confir- mation. ACKNOWLEDGMENTS We thank Debbie Dixon for allowing us to collect insects at the Crowley Museum and Nature Center, Sarasota, FL, and Thomas Dobbs (APHIS-PPQ, USDA, Mi- ami, FL) for companionship in the field and facilitating fieldwork in Miami-Dade Co. Peter Adler (Division of Entomology, Clemson University), J. W. Brown (System- atic Entomology Laboratory [SEL], ARS, USDA, % National Museum of Natural History [NMNH], Washington, DC), and N. E. Woodley (SEL, % NMNH) kindly of- fered suggestions for improving an earlier draft of the manuscript. LITERATURE CITED Austin, D. FE 1978. Exotic plants and their effects in southeastern Florida. Environmental Conservation 5: 25-34. Harris, H. M. 1941. Concerning Neididae, with new species and new records for North America. Bul- letin of the Brooklyn Entomological Society 36: 105-109. Henry, T. J. 1997a. Monograph of the stilt bugs, or Berytidae (Heteroptera), of the Western Hemi- sphere. Memoirs of the Entomological Society of Washington No. 19, 149 pp. . 1997b. Cladistic analysis and revision of the stilt bug genera of the world (Heteroptera: Bery- tidae). Contributions of the American Entomolog- ical Institute 30(1): 1-100. . 2002a. Review of the stilt bug genus Hopli- nus, with the description of a new species and notes on other Hoplinini (Heteroptera: Berytidae: Gampsocorinae). Journal of the New York Ento- mological Society 110: 182-191. . 2002b. Review of the stilt bug genus Yem- matropis (Insecta: Hemiptera: Heteroptera: Bery- tidae), with the description of a new species from Viet Nam. Species Diversity 7: 165-172. Henry, T. J. and R. C. Froeschner. 1998. Catalog of the stilt bugs, or Berytidae, of the world (Insecta: Hemiptera: Heteroptera). Contributions of the American Entomological Institute 30(4): 1-72. Henry, T. J. and D. J. Hilburn. 1990. An annotated list of the true bugs (Heteroptera) of Bermuda. Pro- ceedings of the Entomological Society of Wash- ington 92: 675—684. Hickman, V. V. 1976. The biology of Neides tasman- iensis Gross (Hemiptera: Berytidae). Journal of 8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the Entomological Society of Australia (N.S.W.) 9: 3-10. Institute for Systematic Botany. 2002. Atlas of Florida vascular plants. http://www.plantatlas.usf.edu/ source.asp?plantID = 127 (accessed 20 October 2004). Langeland, K. A. and K. Craddock Burks, eds. 1998. Identification and Biology of Non-native Plants in Florida’s Natural Areas. University Press of Flor- ida, Gainesville, 165 pp. Martorell, L. E 1976. Annotated Food Plant Catalog of the Insects of Puerto Rico. Agricultural Exper- iment Station, University of Puerto Rico [Rio Pie- dras], Department of Entomology, 303 pp. Oman, P. W. 1949. The Nearctic leafhoppers (Homop- tera: Cicadellidae): A generic classification and check list. Memoirs of the Entomological Society of Washington No. 3, 253 pp. Small, J. K. 1933. Manual of the Southeastern Flora; Being Descriptions of the Seed Plants Growing Naturally in Florida, Alabama, Mississippi, East- ern Louisiana, Tennessee, North Carolina, South Carolina and Georgia. The Author, 1554 pp. Stal, C. 1860. Hemiptera. Species novas descripsit. Konglika svenska fregattens Eugenies resa om- kring jorden under befal af C. A. Virgin aren 1851-1853. 2, Zoologi 1. Insecta. Pp. 219-298. Thompson, J. 1919. Para grass. University of Florida Agricultural Experiment Station Press Bulletin 308, 2 pp. Thompson, J. N. 1994. The Coevolutionary Process. University of Chicago Press, Chicago, 376 pp. Uhler, P. R. 1893. A list of the Hemiptera-Heteroptera collected in the island of St. Vincent by Mr. Her- bert H. Smith; with descriptions of new genera and species. Proceedings of the Zoological Soci- ety of London 1893: 705-719. Valley, K and A. G. Wheeler, Jr. 1976. Biology and immature stages of Stomopteryx palpilineela (Lepidoptera: Gelechiidae, a leafminer and leaftier of crownvetch. Annals of the Entomological So- ciety of America 69: 317-324. Van Duzee, E. P. 1909. Observations on some Hemip- tera taken in Florida in the spring of 1908. Bul- letin of the Buffalo Society of Natural Sciences 9: 149-230. Wheeler, A. G., Jr. 1978. Neides muticus (Hemiptera: Berytidae): Life history and description of the fifth instar. Annals of the Entomological Society of America 71: 733-736. . 1986. A new host association for the stilt bug Jalysus spinosus (Heteroptera: Berytidae). Ento- mological News 97: 63-65. . 1994. A new host for Jalysus spinosus (Het- eroptera: Berytidae) and new host family (Com- melinaceae) for stilt bugs. Entomological News 105: 201—203. . 2001. Biology of the Plant Bugs (Hemiptera: Miridae): Pests, Predators, Opportunists. Cornell University Press, Ithaca, New York, 507 pp. Wheeler, A. G., Jr. and T. J. Henry. 1981. Jalysus spi- nosus and J. wickhami: Taxonomic clarification, review of host plants and distribution, and keys to adults and 5th instars. Annals of the Entomolog- ical Society of America 74: 606-615. Wheeler, A. G., Jr. and C. W. Schaefer. 1982. Review of stilt bug (Hemiptera: Berytidae) host plants. Annals of the Entomological Society of America 75: 498-506. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 9-13 A NEW COSTA RICAN SPECIES OF JAPANAGROMYZA (DIPTERA: AGROMYZIDAE) FORMING GALLS ON LONCHOCARPUS (FABACEAE) STEPHANIE BOUCHER AND PAUL HANSON (SB) Department of Natural Resource Sciences, McGill University, Macdonald Cam- pus, Ste-Anne-de-Bellevue, Quebec, H9X 3V9, Canada (e-mail: Stephanie.boucher@ megill.ca); (PH) Escuela de Biologia, Universidad de Costa Rica, San Pedro, San José, Costa Rica Abstract.—Japanagromyza lonchocarpi Boucher, n. sp., is described from specimens reared from petiole-rachis galls on Lonchocarpus oliganthus FJ. Herm. (Fabaceae) in Costa Rica. This is the first record of a Japanagromyza species forming petiole or rachis galls and also the first record of an agromyzid feeding on Lonchocarpus. Key Words: Neotropical Japanagromyza Sasakawa is a small ge- nus of agromyzid flies with 71 species found worldwide. There are 26 species known from the Neotropical Region (Mar- tinez and Etienne 2002). Species with known life history are almost all leaf min- ers. They form large blotch mines on the leaves of their host plant, and subsequently pupate in the soil. However, gall induction by members of this genus has been ob- served on rare occasions. Spencer (1963) reared specimens of Japanagromyza frosti (Frick) from stem-galls on an unidentified plant in Costa Rica and Etienne and Mar- tinez (2003) obtained specimens of Japan- agromyza inferna Spencer from leaf galls on Centrosema virginianum (L.) (Fabaceae) in Guadeloupe. Here we record a new spe- cies of Japanagromyza forming a petiole and rachis gall on Lonchocarpus oliganthus E J. Herm. (Fabaceae) in Costa Rica. MATERIALS AND METHODS Galls were collected from a sapling in a regenerating, former coffee plantation (the Leonel Oviedo reserve) on the University of Costa Rica campus. Both gall collection and Agromyzidae, Japanagromyza, galls, Lonchocarpus, Fabaceae, systematics, adult emergence occurred during October, 2000. Galls were reared in transparent plas- tic bags and emerging adults were preserved in 75% alcohol, then later dried with HMDS (hexamethyldisilazane). Dissection of male genitalia follows procedures in Boucher (2002). Morphological terminology follows McAlpine (1981), except that the orbital and frontal setae are referred to as ors and ori, respectively, in accordance with common usage in agromyzid literature. Type specimens are deposited in the fol- lowing collections (acronyms used in the text are in parentheses): Instituto Nacional de Biodiversidad, Santo Domingo de He- redia, Costa Rica (INBio); Lyman Ento- mological Museum, McGill University, Ste- Anne-de-Bellevue, QC, Canada (LEM); National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (USNM). Japanagromyza lonchocarpi Boucher, new species (Figs. 1—7) Diagnosis.—This species can be distin- guished from other Neotropical species of 10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs.1-6. Male genitalia of Japanagromyza lonchocarpi. 1, Lateral view. 2, Ventral view. 3, Lateral view (variation). 4, Ventrolateral view of phallus (variation). 5, Ejaculatory apodeme. 6, Epandrium with surstylus and cercus (ventral, left). Abbreviations: c, cercus; e, inner corner of epandrium; h, hypandrium; p, phallus; s, surstylus. Scale bars = 0.1 mm VOLUME 108, NUMBER 1 Japanagromyza by the combination of the following characters: prescutellar bristles absent, two strong ors present, fringe of ca- lypter white, male cerci without strong spines, and the shape of the phallus. Description.—Head including antenna, mat brown. Posterior and ventral margin of gena a little darker, brownish black. Thorax with a greenish metallic sheen; calypter in- cluding margin and fringe white; halter completely white. Legs brown. Abdomen with a bronze and/or green metallic sheen. Frons narrow, width 0.31—0.35 mm; ratio of frons width to eye width 0.95—1.1; orbit 0.18 times width of frons at midpoint; frons not projecting above or in front of eye in profile; lunule with silvery pubescence best seen in posterodorsal view; 2 strong recli- nate ori and 2 strong reclinate ors; orbital setulae upright or reclinate, reaching level of posterior ors; in some specimens orbital setulae in two rows anterior to lower ors; first flagellomere small, rounded, slightly pubescent; arista as long as eye height with short pubescence; gena deepest at rear; gena height at midpoint: 0.13—0.15 times eye height. Two strong postsutural dorsocen- trals; acrostichals numerous, 8—9 rows; prescutellar acrostichal bristles absent. Foretibia with one (rarely two) posterior bristle, near midpoint, midtibia with two posterodorsal bristles near midpoint. Wing length 2.4—2.6 mm; R,,; ending close to wing tip; costa extending to M,,,; second costal sector 3.8—4.0 times length of fourth; last section of CuA,: 0.70—0.77 times length of penultimate. Cross-vein r-m lo- cated at 0.4 length of cell dm. Male genitalia: Phallus with variable long coiled distal tubules (Fig. 1—4), sursty- lus with 5—6 stout spines at apex (Fig. 6). Surstylus elongated in side view and fused with epandrium (Fig. 1) . Cercus greatly en- larged without strong bristles or spines, only weak hairs present (Fig. 6). Epandrium with many stout spines on inner corner (Fig. 6). Ejaculatory apodeme narrow and elongated (Fig. 5). Female ovipositor short and retracted. 1] Type material.—Holotype ¢: COSTA RICA, San José, San Pedro de Montes de Oca, (9°56’N, 84°03’W), 1200 m, ex Lon- chocarpus oliganthus, petiole gall, x.2000, P. Hanson (INBio). Paratypes: 3 6, 3 2°, same data as holotype (INBio); 6 6, 4 @, same data as holotype (LEM); 2 6, 2 9, same data as holotype (USNM). Etymology.—The species name is de- rived from the generic name of the host plant. Host plant and life history.—This species forms galls on the petiole and rachis of Lonchocarpus oliganthus (Fig. 7). All 21 specimens were reared from a single gall and the puparia were found in a pile in the basal part of the gall chamber. Because this gall was partially dissected, natural emer- gence was not observed. Although the lar- vae were not observed, they presumably feed gregariously in a single gall. An un- identified microlepidopteran (possibly Cos- mopterigidae; K. Nishida, personal com- munication) larva that bores into the young stems, petiole and rachis is found on the same plant, causing a similar gall. Empty galls of both J. lonchocarpi and the lepi- dopteran are frequently colonized by vari- ous species of ants, some of which tend mealybugs within the empty gall chamber. DISCUSSION Host plants in nine different families have been recorded for Japanagromyza species but members of this genus are best represented on the Fabaceae, subfamily Papilionoideae (Spencer 1990). The host of Japanagromyza lonchocarpi, Lonchocar- pus oliganthus, is in the subfamily Papi- lionoideae and this represents the first rec- ord of an agromyzid species feeding on Lonchocarpus. Although Lonchocarpus oli- ganthus 1s restricted to Central America, the genus Lonchocarpus is widespread in Cen- tral and South America. It is possible that J. lonchocarpi may feed on other species of Lonchocarpus and might be more widely distributed than presently known. There are published records of leaf-min- Fig. 7. ing species of Agromyzidae in which mul- tiple larvae develop in a single mine. For example, up to five larvae of J. etiennei Martinez (Martinez 1994), and up to 20 lar- vae of Chromatomyia alpigenae (Hendel) (Griffiths 1974) can develop in the same mine. Very few gall-inducing agromyzid species are known, but according to pub- lished data, larvae of gall forming species are often solitary (e.g., Hexomyza cecido- gena (Hering) (Spencer 1976), H. simpli- coides (Hendel) (Yamazaki 2001)) or de- velop in a small group of less than five lar- vae (e.g., Japanagromyza inferna Spencer (Etienne and Martinez 2003)). Thus, having as many as 21 specimens of J. lonchocarpi developing in a single gall seems quite un- usual. There are still many species of Japana- gromyza whose biology is unknown. Three species, all from the Neotropical region, are now known to induce galls. It is to be ex- pected that more described and undescribed species are also gall inducers. 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ee) Gall of Japanagromyza lonchocarpi on Lonchocarpus oliganthus. ACKNOWLEDGMENTS We thank the Instituto Nacional de Bio- diversidad for inviting S. Boucher to visit the institute to study their Agromyzidae collection, and Manuel Zumbado (INBio) for assistance during her visit. We also thank JuLin Weng for bringing this gall to our attention, Terry Wheeler for reviewing the manuscript, and Kenji Nishida for iden- tification of the lepidopteran larva. LITERATURE CITED Boucher, S. 2002. Revision of Nearctic species of Cer- odontha (Cerodontha) (Diptera: Agromyzidae). The Canadian Entomologist 134: 577—603. Etienne, J. and M. Martinez. 2003. Les Agromyzidae de Guadeloupe: Espéces nouvelles et notes additionnelles (Diptera). Nouvelle Revue d’Entomologie 19: 249-272. Griffiths, G. C. D. 1974. Studies on boreal Agromy- zidae (Diptera). V. On the genus Chromatomyia Hardy, with revision of Caprifoliaceae-mining species. Quaestiones Entomologicae 10: 35—69. Martinez, M. 1994. Japanagromyza etiennei n.sp. (Diptera: Agromyzidae) ravageur potential des Phaseolus spp. [Leguminosae] dans les caraibes. VOLUME 108, NUMBER 1 Revue Frangaise d’entomologie (nouvelle serie) 16: 81-85. Martinez, M. and J. Etienne. 2002. Liste systématique et biogéographique des Agromyzidae (Diptera) de la région néotropicale. Bollettino di zoologia agraria e di Bachicoltura, Ser.II, 34: 25-52. McAlpine, J. EF 1981. Morphological terminology— adults, pp. 9-63. In McAlpine, J. E et al., eds. Manual of Nearctic Diptera, Vol. 1. Monograph No. 27, Research Branch, Agriculture Canada. 674 pp. Spencer, K. A. 1963. A synopsis of the Neotropical 13 Agromyzidae (Diptera). Transactions of the Royal Entomological Society of London 115: 291-389. . 1976. The Agromyzidae (Diptera) of Fennos- candia and Denmark. Fauna Entomologica Scan- dinavica, Vol. 5, Part 1, 304 pp. . 1990. Host specialization in the World Agro- myzidae (Diptera). Series Entomologica 45. Klu- wer Academic Publishers, Dordrecht, 444 pp. Yamazaki, K. 2001. Preference-performance linkage in the willow twig-galling agromyzid fly, Hexomyza simplicoides (Diptera: Agromyzidae) on the wil- low Salix chaenomeloides. Entomological Science 4: 301-306. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 14-23 PALEARCTIC SYMPIESIS ACALLE AND SYMPIESIS GORDIUS (HYMENOPTERA: EULOPHIDAE) IN NORTH AMERICA: TAXONOMIC CHANGES AND A REVIEW OF NEARCTIC HOST RECORDS CHRIS T. MAIER AND CHRISTER HANSSON (CTM) Department of Entomology, Connecticut Agricultural Experiment Station, P.O. Box 1106, New Haven, CT 06504, U.S.A. (e-mail: Chris.Maier@po.state.ct.us); (CH) Department of COB, Zoology, Helgonavagen 3, SE-223 62 Lund, Sweden (e-mail: Christer. Hansson @cob.lu.se) Abstract.—The Palearctic eulophids, Sympiesis acalle (Walker) and S. gordius (Walker), are widespread in North America where they have been known as S. bimaculatipennis Girault and S. marylandensis Girault, respectively. To update their taxonomy, we have established seven new synonyms: Astichus bimaculatipennis Girault, Sympiesis bimacu- lata Crawford, and S. meteori Girault = S. acalle (Walker); and, S. marylandensis Girault, S. rex Girault, S. miltoni Girault, and S. lexingtonensis Girault = S. gordius (Walker). We also have designated six lectotypes: Eulophus acalle Walker, E. gordius Walker, E. ala- parus Walker, E. pisenor Walker, Sympiesis rex Girault, and S. lexingtonensis Girault. Finally, we summarize the North American host records of Sympiesis acalle and S. gor- dius, which frequently are cited as biocontrol agents of pestiferous leafminers. Key Words: Sympiesis marylandensis Many eulophid wasps (Chalcidoidea: Eu- lophidae) have a large distribution, with some species occurring on two or more continents (Noyes 2002). Numerous species occur in both Europe and North America (e.g., Hansson 1987), a fact often over- looked because certain species have a dif- ferent name in Europe and North America. Such nomenclatural mistakes make it diffi- cult to understand fully the natural history of a species and even might interfere with biocontrol efforts. Here we reveal two ad- ditional cases of mistaken identity, both in the genus Sympiesis Forster. In the process, we establish seven new synonyms and des- ignate six lectotypes. Bellostas et al. (1998) have published color photographs of parasitoids of Euro- pean leafminers, Phyllonorcyter spp. (Lep- Eulophidae, hosts, leafminer, Phyllonorycter, Sympiesis bimaculatipennis, idoptera: Gracillariidae), of cultivated ap- ple, Malus domestica Borkhausen. Among their images were two eulophid species, Sympiesis acalle (Walker) and S. gordius (Walker) that appeared remarkably similar to the North American species, S. bimacu- latipennis (Girault) and S. marylandensis Girault, respectively. In North America, Miller (1970), Maier (1988a, b), and others have recorded hosts of these two parasitoids to be species of Phyllonorycter and other Lepidoptera whose larvae live in mines and other concealed situations. In Canada and the United States, S. marylandensis is an important natural enemy of leafminers, es- pecially the apple pests, Phyllonorycter blancardella (Fabricius), P. crataegella (Clemens), and P. mespilella (Hiibner) (e.g., Maier 1994, 2002). In this paper, we VOLUME 108, NUMBER 1 note the synonymy and summarize the North American records of hosts of the two Sympiesis spp. This information hopefully will assist economic entomologists in rec- ognizing that the biocontrol previously at- tributed to two “‘native”’’ species is, in fact, caused by two European species, or possi- bly holarctic species, that have long been associated with apple leafminers in Euro- pean orchards. LECTOTYPE DESIGNATIONS AND NEw SYNONYMS Type specimens discussed below are de- posited in the following museums: BMNH, The Natural History Museum, London, England; INHS, Illinois Natural History Survey, Champaign, IL, USA; USNM, Smithsonian Institution, National Museum of Natural History, Washington, DC, USA. Sympiesis acalle (Walker) Eulophus acalle Walker 1848: 234. Lecto- type male (new designation) in BMNH, examined. Entedon nubeculatus Ratzeburg 1848: 158. Synonymized by Bouéek (1959). Eulophus bifasciatus Thomson 1878: 230. Synonymized by Bouéek (1959). Eulophus punctifrons Thomson 1878: 231. Synonymized by Bouéek (1959). Astichus bimaculatipennis Girault 1912: 8. Holotype female in INHS, examined. New synonymy. Sympiesis bimaculata Crawford 1913: 259. Holotype female in USNM, examined. New synonymy. Sympiesis meteori Girault 1916: 37. Holo- type female in USNM, examined. New synonymy. Sympiesis acalle: Graham 1959: 182. A single male with the name ‘“Eulophus acalle”’ is in the collection at BMNH (with the label “Eulophus acalle Walker’? and our lectotype label). The gaster has been gnawed on left side, but otherwise the spec- imen is intact. The original description was based on one or more male specimens. The 15 single remaining specimen agrees with the original description. Walker (1848) did not indicate on how many specimens his de- scription was based; therefore, to keep the nomenclature stable, the name hereby is fixed to the male specimen which is here designated the lectotype. The female holotypes of A. bimaculati- pennis, S. bimaculata, and S. meteori agree well with our interpretation of S. acalle (i1.e., with females linked to males that are conspecific with the lectotype male of S. acalle); and, S. acalle, S$. bimaculatipennis, S. bimaculata, and S. meteori undoubtedly belong to the same species. The holotype of A. bimaculatipennis has the mesosoma, in- cluding only front left leg, glued to a point (type no. 45.012); the head (squashed) and the antennae are on a slide (type no. 45.013); wings, most of the legs, and the gaster are missing. The holotype of S. bi- maculata is glued to a point (type no. 15094); the entire left antenna, right anten- na beyond the scape, and right wing pair are missing. The holotype of S. meteori is glued to a point (type no. 19995) and is broken in two parts, gaster and mesosoma; the head, pair of right wings and left hind wing, right fore tarsus, right mid tibia and tarsus, and left and right hind tibiae and tar- Si are missing. Symptesis gordius (Walker) Eulophus gordius Walker 1839: 129. Lec- totype male (new designation) in BMNH, examined. Eulophus alaparus Walker 1839: 163. Lec- totype female (new designation) in BMNH, examined. Synonymized by Bouéek (1959). Eulophus pisenor Walker 1839: 153. Lec- totype male (new designation) in BMNH, examined. Synonymized by Bouéek and Askew (1968). Eulophus cervicornis Forster 1841: 43. Synonymized by Bouéek (1959). Entedon padellae Ratzeburg 1844: 166. Synonymized by Bouéek (1959). Eulophus bulmerincqii Ratzeburg 1848: 16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 155. Synonymized by Boucek and Askew (1968). Eulophus laevissimus Ratzeburg 1848: 157. Synonymized by Bouéek (1959). Eulophus stramineipes Thomson 1878: 232. Synonymized by Bouéek (1959). Sympiesis rex Girault 1917a: 2-3. Lecto- type female (new designation) in USNM, examined. New synonymy. Sympiesis miltoni Girault 1917b: 7. Lecto- type female, designated by Miller (1970), in USNM, examined. New synonymy. Sympiesis marylandensis Girault 1917c: 37. Holotype female in USNM, examined. New synonymy. Sympiesis lexingtonensis Girault 1917d: 6. Lectotype female (new designation) in USNM, examined. New synonymy. Eulophus albiscapus Erdos 1954: 327. Syn- onymized by Bouéek (1959). Sympiesis gordius: Graham 1959: 183. Single specimens under the names “Eu- lophus gordius” (a male), ““Eulophus ala- parus” (a female), and “‘Eulophus pisenor” (a male) are in the collection at BMNH. Walker’s (1839) descriptions of these three species were based on an unknown number of specimens. Therefore, to stabilize the no- menclature, we chose these three specimens as the lectotypes for each species. The spec- imens are labelled ‘“‘Eulophus gordius Walker,” ““Eulophus alaparus Walker,” and **Eulophus pisenor Walker’; each also has our lectotype label. A single specimen under the name Sym- piesis rex is in the collection at USNM. Gi- rault (1917a) did not indicate on how many specimens he based his description. There- fore, to stabilize the nomenclature, we chose this specimen as the lectotype for S. The specimen carries the labels ‘“‘Quaintance No. 11468,” “Salem, Ore, vi.15,” ““Reared Phyllonorycter crataegel- la,” “*E. J. Newcomer collector,’ “20218,” “Sympiesis rex Girault female type,’ and our lectotype label. A single specimen stands under the name Sympiesis lexingtonensis in the collection at rex. USNM. Girault (1917d) did not indicate on how many specimens he based his descrip- tion. Therefore, to stabilize the nomencla- ture, we chose this specimen as the lecto- type for S. lexingtonensis. The specimen carries the labels “1866,” “20196,” “‘Sym- piesis lexingtonensis Gir. female type,” and our lectotype label. The female holotype of S$. marylandensis, and the lectotypes of S. rex, S. miltoni, and S. lexingtonensis agree well with our inter- pretation of S. gordius (1.e., with females linked to males that are conspecific with the lectotype male of S. gordius); and, S. gor- dius, S. rex, S. miltoni, S. marylandensis, and S. /exingtonensis undoubtedly belong to the same species. The lectotype of S. rex is glued to a point (type no. 20218); the head and right wing pair are missing. The lec- totype of S. miltoni is glued to a point (type no. 21399), and the type is intact. The ho- lotype of S. marylandensis is glued to a point (type no. 20137); the head is missing (according to the original description the head is on a slide, not examined by us). The lectotype of S. lexingtonensis is glued to a point (type no. 20196); the head, right legs (all three), and pair of right wings are miss- ing. BIOLOGY Hosts of both S. acalle and S. gordius that were discovered during original North American studies (Table 1) are repeated in the catalogs of Peck (1951, 1963) and Burks (1979) and, to some extent, in the revision of Miller (1970). Green (1979) and Maier (1988b) have since reported many new North American hosts. Sympiesis acal- le, previously known as S. bimaculatipennis in North America, is both a primary and secondary ectoparasitoid of lepidopteran larvae and pupae (Table 1) that usually live in concealed places, such as blotch mines or rolled leaves. Sympiesis acalle has at least 15 hosts in North America. It is a pri- mary parasitoid of Tischeriidae (1 species), Gracillariidae (5), Oecophoridae (1), and Tortricidae (6). In addition, it parasitizes the VOLUME 108, NUMBER 1 7 Table 1. Hosts of Sympiesis acalle and S. gordius that were recorded as S. bimaculatipennis and S. mary- landensis, respectively, in the North American literature. Due to taxonomic revisions or changes in generic use and species concepts, some of the hosts cannot be placed in a genus and are excluded here. Hosts given only to genus are followed by the genus of their host plant in brackets; if these hosts could be one of the fully identified hosts, they are not included in the total number of hosts given in the text. Generic records without an associated host plant are excluded from the main part of the table. To the best of our knowledge, all of the scientific names are the ones currently in use. Order, Family, and Species of Host S. acalle — S. gordius Literature Sources LEPIDOPTERA Nepticulidae Nepticula variella (Braun) x Green (1979) Tischeriidae Tischeria malifoliella Clemens* x xX Dunnam (1924), Peck (1951, 1963), Miller (1970), Burks (1979), Oatman (1985) Tischeria sp. [Solidago] x Maier (1994, unpublished data) Heliozelidae Coptodisca powellella Opler x Green (1979) Lyonetidae Bacculatrix albertiella Busck x Green (1979) Gracillariidae Caloptilia agrifoliella Opler xX Green (1979) Caloptilia sp. [Betula] x Miller (1970), Burks (1979) Cameraria agrifoliella (Braun) x Green (1979) C. caryaefoliella (Clemens) xX Heyerdahl and Dutcher (1985) C. wislizeniella Opler xX Green (1979) Cameraria spp. [Ostrya] xX Maier (1988b) Cremastobombycia solidaginis x Maier (1988b) (Frey and Boll) Neurobathra bohartiella (Opler) x Green (1979) Parectopa robiniella Clemens XxX Weaver and Dorsey (1965) Parornix geminatella (Packard)* x XxX Haseman (1916), Girault (1916), Copenhafer and Parker (1938), Peck (1951, 1963), Mill- er (1970), Burks (1979), Oatman (1985), Maier (1988b) Parornix sp. [Betula] xX Maier (1988b) Parornix sp. [Lyonia] x Maier (1988b) Phyllocnistis populiella Chambers Xx Miller (1970), Burks (1979) Phyllonorycter aeriferella (Clemens) xX Maier (1988b) P. albanotella (Chambers) x Maier (1988b) P. argentifrimbriella (Clemens) x Maier (1988b) P. argentinotella (Clemens) xX Maier (1988b) P. antiochella (Opler) xX Green (1979) P. auronitens (Frey and Boll) x Maier (1988b) P. basistrigella (Clemens) xX Miller (1970), Burks (1979), Maier (1988b) P. blancardella (Fabricius) x x Miller (1970), Pottinger and LeRoux (1971), Johnson et al. (1976), Dutcher and Howitt (1978), Burks (1979), Weires et al. (1980), Van Driesche and Taub (1983), Maier (1984, 1988b, 1992, 1993, 1994, 2002), Hagley (1985), Ridgway and Mahr (1985, 1989, 1990), Biggs and Hagley (1988), Hagley and Barber (1991), Gagné and Bar- rett (1994), Bishop et al. (2001) P. caryaealbella (Chambers) x Heyerdahl and Dutcher (1985) 18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Order, Family, and Species of Host S. acalle S. gordius Literature Sources P. celtisella (Chambers) x Miller (1970), Burks (1979) P. crataegella (Clemens) xX x Beckham et al. (1950), Peck (1951, 1963), Par- ent (1962), Miller (1970), Burks (1979), Wei- res et al. (1980), Maier (1982, 1988b, 1992, 1993, 1994), Gambino and Sullivan (1982), Van Driesche (1983), Van Driesche and Taub (1983), Drummond et al. (1985), Oatman (1985), Van Driesche et al. (1985), Gagné and Barrett (1994), Maier and Weseloh (1995) P. emberizaepenella (Bouché) x Maier (1988b) P. felinelle Heinrich Xx Miller (1970), Burks (1979) P. fitchella (Clemens) x Maier (1988b) P. inusitatella (Braun) xX Green (1979) P. kearfottella (Braun) xX Maier (1988b) P. lucetiella (Clemens) Xx Maier (1988b) P. lucidicostella (Clemens)* x Miller (1970), Burks (1979), Maier (1988b) P. mespilella (Hiibner) x x Doganlar and Beirne (1980), Barrett and Jor- gensen (1986), Varela dn Welter (1992), Cossentine and Jensen (1994), McGregor (1996) P. morrisella (Fitch) xX Maier (1988b) P. obscuricostella (Clemens) Xx Maier (1988b) P. propinquinella (Braun) x Miller (1970), Burks (1979), Maier (1988a, b, 1994) P. restrictella (Braun) Xx Maier (1988b) P. robiniella (Clemens) xX Weaver and Dorsey (1965), Miller (1970), Burks (1979), Maier (1988b) P. salicifoliella (Chambers)! xX Miller (1970), Burks (1979), Maier (1988b) P. sandraella (Opler) x Green (1979) P. scudderella (Frey and Boll) xX Maier (1988b) P. tremuloidiella (Braun) Xx Maier (1988b) P. trinotella (Braun)! xX Maier (1988b) P. tritaenianella (Chambers) Xx Maier (1988b) Phyllonorycter sp. near crataegella x xX Maier (1988a, b, 1994) (Clemens) Phyllonorycter sp. [Alnus] Xx Miller (1970) Phyllonorycter sp. [Amphicarpa| x Miller (1970) Phyllonorycter sp. [Fagus]' xX Miller (1970) Phyllonorycter sp. |Lonicera] xX Miller (1970) Phyllonorycter sp. [Malus] x Girault (1917a), Peck (1951, 1963), Newcom- ber (1958), Miller (1970) Phyllonorycter sp. [Populus] xX Miller (1970) Phyllonorycter sp. [Prunus] xX Miller (1970) Phyllonorycter sp. [Tilia] XxX Miller (1970) Oecophoridae Agonopteryx posticella (Wal- xX Miller (1970), Burks (1979) singham) Cosmopterygidae Stilbosis dulcedo (Hodges) Xx Green (1979) Gelechiidae Coleotechnites thujaella (Kearfott) xX Miller (1970), Burks (1979) Tortricidae Acleris chalybeana (Fernald) xX Miller (1970), Burks (1979) VOLUME 108, NUMBER 1 Table 1. Continued. IS) Order, Family, and Species of Host S. acalle S. gordius Literature Sources Ancylis comptana (Frolich) x Choristoneura rosaceana (Harris) x Epinotia laracana (Kearfott) xX Rhopobota naevana (Hibner) x Spilonota laricana (Heinemann) xX Spilonota ocellana (Denis and xX Schiffermiiller) HYMENOPTERA Tenthredinidae Heterarthrus nemoratus (Fallén) Braconidae" Pholetesor ornigis (Weed)' x Chalcididae Spilochalcis sp.' x Miller (1970), Burks (1979) Schuh and Mote (1948), Peck (1963) Miller (1970) Fitzpatrick et al. (1994) Burks (1979) Sadava and Miller (1967), Miller (1970), Burks (1979) Miller (1970), Burks (1970) Pottinger and LeRoux (1971), Maier (1982, 1984), Bishop et al. (2001) Bishop et al. (2001) *Dunnam’s (1924) record apparently is repeated by Peck (1951, 1963), Miller (1970), and Burks (1979). > Both Miller (1970) and Burks (1979) gave the Caloptila host as C. elongella (Linnaeus), but this species is not known to occur in North America. © Haseman’s (1916) study provided the record of S. acalle given in Girault (1916) and Peck (1951, 1963), and Copenhagen and Parker’s (1938) record apparently also was repeated for one of the junior synonyms of S. acalle in Peck (1963); the record in Burks (1979) probably was repeated from an earlier catalog. 4 Girault (1917d) listed a host as a blotch miner on maple, which may have been a Phyllonorycter sp. or possibly a Cameraria sp. © This species on Populus sp. mentioned in Miller (1970) and probably repeated by Burks (1979) is more likely to be Phyllonorycter tremuloidiella (Braun) or another Phyllonorycter sp. on Populus; P. salicifoliella is a willow-feeder. ‘Procter (1938, 1946) mentioned S. gordius as a parasite of blotch miners on Fagus (probably Phyllonorycter restrictella) and Salix (probably Phyllonorycter scudderella or P. salicifoliella), and the record of the blotch miner on Fagus is repeated in Peck (1951, 1963); the record of the blotch miner on Fagus is duplicated in Peck (1951, 1963) and Miller (1970). = Newcomer (1958) did not report his record from an apple-feeding Phyllonorycter sp. (misidentified as P. crataegella) until 1958; this record is the one mentioned in Girault (1917a). " Webster (1909) retrieved S. acalle from an insectary and reported that is parasitized an unidentified internal primary parasitoid of the tortricid moth, Acleris minuta (Robinson), a record that was repeated, sometimes in altered form, by Girault (1912), Frison (1927), Peck (1951, 1963), and Burks (1979). ‘Primary hosts of S. acalle on Pholetesor ornigis were Phyllonorycter blancardella and P. crataegella; that from Spilochalcis sp. was P. blancardella. braconid Pholetesor ornigis (Weed) and the chalcid Spilochalcis sp., which both attack apple leafminers, Phyllonorycter spp. The casual rearing methods used in some stud- ies (e.g., Oatman 1985) may have resulted in overlooking hyperparasitic associations or in identifying the wrong primary host. In northeastern North America, S. acalle is a facultative hyperparasitoid of the gracillari- id leafminers Phyllonorycter blancardella and P. crataegella, attacking especially their braconid parasitoid, Pholetesor ornigis (e.g., Maier 1982, 1984). Under the name of S$. marylandensis in the North American literature, S. gordius is known to be a primary ectoparasitoid of lar- vae and sometimes pupae of miners, skel- etonizers, and rollers of leaves, and of other insects (Table 1) that live in sheltered situ- ations. Most of the original host records are from Miller (1970), Green (1979), and Maier (1988b). In North America, S. gor- dius parasitizes at least 53 species in the families Nepticulidae (1), Tischeriidae (2), Heliozelidae (1), Lyonetidae (1), Gracillar- lidae (44), Cosmopterygidae (1), Gelechi- 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON idae (1), Tortricidae (1), and Tenthredinidae (1). Maier (1988b) has concluded that in the northeastern United States S. gordius pre- fers gracillariid larvae inhabiting mines on the underside of leaves. Some of the west- ern host records may refer to parasitization by S. yuekseli Doganlar (1979), which is similar to S. gordius and was described af- ter some of the records in Table 1 were pub- lished. Based on host records given by Bou- cek and Askew (1968), Storozheva (1982), and others, both S. acalle and S. gordius have similar hosts in the Palearctic and Ne- arctic regions. Delucchi (1958), Boucek and Askew (1968), Storozheva (1982), and oth- ers have reported that S. gordius sometimes is a secondary parasitoid of braconids and eulophids, but this type of association is not yet known in North America. Both S. acalle and S. gordius are men- tioned in the biocontrol literature about gra- cillariids in apple orchards in Europe (e.g., Baggiolini 1960, Casas and Baumgartner 1990, Bellostas et al. 1998) and in North America (Maier 1994, 2002, and references therein). Especially S$. gordius has been touted as an important natural enemy of the pestiferous apple leafminers, Phyllonoryc- ter blancardella, P. crataegella, and P. mespilella (see references in Table 1). Maier (1984, 1992) and Gagné and Barrett (1994) have examined the seasonal flight activity of S. gordius; Maier (1992) sug- gested that it could have six or seven gen- erations per year while its host P. cratae- gella has only three. Also, Maier and We- seloh (1995) have shown that S. gordius tended to have higher abundance near the edge than the center of apple orchards. We consider it to be highly likely that the two European eulophids were introduced accidentally into North America with one or both of their European hosts, P. blan- cardella and P. mespilella, which are leaf- miners of non-native apple. The European facultative hyperparasitoid, S$. sericeicornis (Nees), which for many years masqueraded under the junior synonym S. conica (Pro- vancher), probably had a similar route of entry into North America. Although it is less likely, these Sympiesis spp., known so long by their junior synonyms in North America, actually could be Holarctic or could have spread from North America to Europe. Comparative studies of genetic di- versity of populations in Europe and North America might clarify the continent(s) of origin of these Sympiesis spp. ACKNOWLEDGMENTS We are grateful to C. Favret (INHS), M. W. Gates (USNM), and S. Lewis (BMNH) for the loan of specimens. LITERATURE CITED Baggiolini, M. 1960. 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Inoculative release and establishment of Holcothorax testaceipes (Hymenoptera: Encyrti- dae), a Palaearctic agent for the biological control of Phyllonorycter spp. (Lepidoptera: Gracillari- idae) in Connecticut apple orchards. Journal of Economic Entomology 86: 1069—1077. . 1994. Biology and impact of parasitoids of Phyllonorycter blancardella and P. crataegella (Lepidoptera: Gracillariidae) in northeastern North American apple orchards, pp. 6—24. Jn Maier, C. T., ed. Integrated Management of Tentiform Leaf- miners, Phyllonorycter spp. (Lepidoptera: Gracil- lariidae), in North American Apple Orchards. Thomas Say Publications in Entomology, Ento- mological Society of America, Lanham, Mary- land, 82 pp. . 2002 (2001). Exotic lepidopteran leafminers in North American apple orchards: Rise to prom- inence, management, and future threats. Biologi- cal Invasions 3: 283-293. Maier, C. T. and R. M. Weseloh. 1995. Spatial distri- bution of Sympiesis marylandensis Girault (Hy- menoptera: Eulophidae) in apple orchards infested by its host, the apple blotch leafminer, Phyllono- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rycter crataegella (Clemens) (Lepidoptera: Gra- cillariidae). The Canadian Entomologist 127: 235-243. McGregor, R. 1996. Phenotypic selection by parasit- oids on the timing of life history in a leafmining moth. Evolution 50: 1579-1584. Miller, C. D. F 1970. The Nearctic species of Pnigalio and Sympiesis (Hymenoptera: Eulophidae). Mem- oirs of the Entomological Society of Canada 68: 1-121. Newcomer, E. J. 1958. Some parasites and predators of fruit pests in the Pacific Northwest. Pan-Pacific Entomologist 34: 87-91. Noyes, J. S. 2002. Interactive Catalogue of World Chalcidoidea. Electronic publication (CD-ROM). Taxapad 2002, Vancouver, British Columbia. Oatman, E. R. 1985. Parasites associated with lepidop- terous leaf mines on apple in northwestern Wis- consin. Journal of Economic Entomology 78: 1063-1066. Parent, B. 1962. Observations sur trois nouveaux in- sectes nuisibles aux pommiers dans le Québec: Thamnosphecia scitula scitula Harris (Lépidop- tere: Aegeriidae), Lithocolletis crataegella Clem. (Lépidoptéres: Lithocolletidae), et Quadraspidi- otus ostreaeformis Curtis (Homopteéres: Diaspidi- dae). Annales de la Société Entomologique du Québec 7: 19—24. Peck, O. 1951. Superfamily Chalcidoidea, pp. 410— 594. In Muesebeck, C. FE W., K. V. Krombein, and H. K. Townes, eds. Hymenoptera of America North of Mexico, Synoptic Catalog. U.S. Depart- ment of Agriculture Monograph No. 2, 1420 pp. . 1963. A Catalogue of the Nearctic Chalcidoi- dea (Insecta: Hymenoptera). The Canadian Ento- mologist, Supplement 30, 1092 pp. Pottinger, R. P. and E. J. LeRoux. 1971. The biology and dynamics of Lithocolletis blancardella (Lep- idoptera: Gracillariidae) on apple in Quebec. Memoirs of the Entomological Society of Canada 77: 1-437. Procter, W. 1938. Biological Survey of the Mount De- sert Region. Part VI. The Insect Fauna. Wistar In- stitute of Anatomy and Biology, Philadelphia, Pennyslvania, 496 pp. . 1946. Biological Survey of the Mount Desert Region. Part VII. The Insect Fauna. Wistar Insti- tute of Anatomy and Biology, Philadelphia, Penn- sylvania, 566 pp. Ratzeburg, J. T. C. 1844. Die Ichneumonen der For- stinsekten in forstlicher und entomologischer Be- ziehung. 1. Berlin, 224 pp. . 1848. Die Ichneumonen der Forstinsekten in forstlicher und entomologischer Beziehung. 2. Berlin, 238 pp. Ridgway, N. M. and D. L. Mahr. 1985. Natural ene- mies of the spotted tentiform leafminer, Phyllon- orycter blancardella (Lepidoptera: Gracillariidae), VOLUME 108, NUMBER 1 in sprayed and unsprayed apple orchards in Wis- consin. Environmental Entomology 14: 459—463. . 1989. Laboratory methods for rearing spotted tentiform leafminer (Lepidoptera: Gracillariidae) and two of its parasitoids. Journal of Economic Entomology 82: 319-321. . 1990. Reproduction, development, longevity, and host mortality of Sympiesis marylandensis (Hymenoptera: Eulophidae), a parasitoid of spot- ted tentiform leafminer (Lepidoptera: Gracillari- idae), in the laboratory. Annals of the Entomolog- ical Society of America 83: 795-799. Sadava, D. and C. D. FE Miller. 1967. Taxonomy of last-instar larval remains of parasites reared from Spilonota ocellana. The Canadian Entomologist 99: 436-442. Schuh, J. and D. C. Mote. 1948. The oblique-banded leaf roller on red raspberries. Oregon (Corvallis) Agricultural Experiment Station Technical Bulle- tin 13: 1—43. Storozheva, N. A. 1982. A key to Palearctic species of parasitic wasps of the genus Sympiesis Forster, 1856 (Hymenoptera, Eulophidae). Entomologi- cheskoye Obozreniye 61(1): 164-176 (English translation, Entomological Review 61(1): 156— 168). Thomson, C. G. 1878. Hymenoptera Scandinaviae, 5. Lund, 307 pp. Van Driesche, R. G. 1983. Meaning of “percent par- asitism” in studies of insect parasitoids. Environ- mental Entomology 12: 1611—1622. Van Driesche, R. G. and G. Taub. 1983. Impact of i) eS) parasitoids on Phyllonorycter leafminers infesting apple in Massachusetts, U.S.A. Protection Ecolo- gy 5: 303-317. Van Driesche, R. G., J. M. Clark, M. W. Brooks, and FE J. Drummond. 1985. Comparative toxicity of orchard insecticides to the apple blotch leafminer, Phyllonorycter crataegella (Lepidoptera: Gracil- lariidae), and its eulophid parasitoid, Sympiesis marylandensis (Hymenoptera: Eulophidae). Jour- nal of Economic Entomology 78: 926—932. Varela, L. G. and S. C. Welter. 1992. Parasitoids of the leafminer, Phyllonorycter nr. elmaella (Lepidop- tera: Gracillariidae), on apple in California: Abun- dance, impact on leafminer, and insecticide-in- duced mortality. Biological Control 2: 124—130. Walker, F 1839. Monographia Chalciditum. I. London, 333 5pp: . 1848. List of the specimens of hymenopterous insects in the collection of the British Museum, part 2, additional species. London, pp. 99-237. Weaver, J. E. and C. K. Dorsey. 1965. Parasites and predators associated with five species of leaf-min- ing insects in black locust. Annals of the Ento- mological Society of America 58: 933-934. Webster, R. L. 1909. The lesser apple leaf-folder. lowa (Ames) Agricultural Experiment Station Bulletin 102: 178-212. Weires, R. W., D. R. Davis, J. R. Leeper, and W. H. Reissig. 1980. Distribution and parasitism of gra- cillariid leafminers on apple in the Northeast. An- nals of the Entomological Society of America 73: 541-546. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 24-34 NEW SCORPIONFLIES (MECOPTERA: PANORPIDAE) FROM JALISCO, MICHOACAN, AND OAXACA, MEXICO WESLEY BICHA 121 Old Batley Road, Oliver Springs, TN 37840, U.S.A. (e-mail: bichawj @ornl.gov) Abstract.—Four new species, Panorpa dividilacinia, P. umbricola, P. mixteca, and P. insigna, are described and illustrated from the Mexican states of Jalisco, Michoacan, and Oaxaca. Habitat details are provided. These additions bring the total number of known species of southwestern and south central Mexican Panorpa to 16 and the total of Mexican species to 28. Key Words: There are currently 12 species of Pan- orpa known from southwestern and south central Mexico. These scorpionflies are generally found in moist, shady areas above 1200 m in elevation in the mountainous ar- eas of Mexico. In the nineteenth century P. terminata Klug, which has since been found from a number of locations in the Cordellia Volcanica, was described. In the early twen- tieth century P. immaculata Esben-Peter- sen, found throughout the Sierra Madre del Sur, was added. During the past 45 years Dr. George Byers has greatly expanded our knowledge and the number of known spe- cies from this region, adding P. penicillata Byers from the Sierra Madre Occidental, P. azteca Byers, P. serta Byers, P. apiculata Byers, and P. truncata Byers from the Cor- dellia Volcanica, and P. bimacula Byers, P. gladiata Byers, P. acicularis Byers, P. acu- leata Byers, and P. luteola Byers from the Sierra Madre del Sur. It is possible that P. punctata Klug occurs in this region of Mex- ico. The type location of the single female was unstated and no additional specimens have been located in the century since its description. Extensive field work in this region throughout the past several decades has re- new species, scorpionfly, Mecoptera, Panorpa, Mexico sulted in an additional four species being discovered which are described in this pa- per. Panorpa dividilacinia Bicha, new species (Figs. 1-4) Description—Based on 26 6, 23 2, pinned from alcohol. Head: Dorsum brown; glossy black cross band above antennal bases, extending along inner margin of eyes to posterior mar- gin of vertex. Rostrum yellowish brown, with two longitudinal bands in some indi- viduals; genae pale yellow. Labrum and maxillary palps brown. Antennal scape pale yellowish brown, pedicel dark brown to black, flagellum black, with approximately 40 flagellomeres. Thorax: Pronotum black, pale yellowish along posterior margin; 9—11 black setae at each side on anterior margin. Mesonotum and metanotum black with yellowish white median band, with numerous short, black hairs directed caudad. Scutellum yellowish- white. Pleural surfaces, coxae and mera sor- did white to pale yellow with numerous short, dark setae on anterior coxa and on mesothoraxic coxa and episternum; sparse, pale setae on metathorax, most numerous VOLUME 108, NUMBER | 2 3 “ 2) Figs. 1-5. 1—4, Panorpa dividilacinia, paratypes. 1, Male right forewing. 2, Male genital bulb, ventral aspect. 3, Hypovalves and aedeagus, ventral aspect, left hypovalve removed for clarity, slightly enlarged scale. 4, Female genital plate, ventral aspect (posterior end at top). 5, Panorpa immaculata, hypovalve, ventral aspect. on coxa; small black spot at each end of mesepimeron and metepimeron. Femora and tibiae sordid white with numerous short, apically-directed black hairs and a lesser number of black setae. Tibiae with two brown spines. Tarsi sordid white with numerous apically-directed golden hairs. Wings (Fig. 1) faintly tinged with grey brown, slightly iridescent, markings brown, veins brown, stigma dark yellowish, thyri- dium at first fork of M white; pterostigmal band reduced to triangle extending from C to R,, sometimes broken with a small spot posterior to R;; some individuals with faint, small basal spot at Cu,. Wing markings typ- ically more pronounced in females. Male abdomen: Terga 2—5 glossy black; sterna 2—3 sordid white; sterna 4—5 grading from brown to black; segment 6 glossy black, long, cylindrical, narrowed posteri- orly, without anal horn; segments 7—8 short, yellowish orange. Tergum 3 posterior mar- 26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gin with notal organ slightly pronounced above; tergum 4 anterior margin with small protuberance. Hypovalves (Fig. 2) of ster- num 9 slender, divergent from narrow pro- longation of sternum, parallel, extending half way between ventral fusion of basi styles and bases of dististyles, apices trun- cated at 60° on mesal edge. Tergum 9 nar- rowing apically, extending an additional one-third beyond cerci to just beyond bases of dististyles, apically emarginate. Cerci darker apically. Basistyles joined ventrally. Dististyles outer margin slightly concave until acute tip; with a large, cuplike basal lobe prolonged ventromesally; dorsomesal edge of basal cup with triangular tooth. Ae- deagus (Fig. 3) with straight, slender, sin- gle-branched ventral parameres extending one-third length beyond bases of dististyles; dorsal parameres, long, straight, dorsally flattened, extending just beyond bases of dististyles. Dorsal valves small, rounded, dark; ventral valves cone-shaped. Female abdomen: Terga and sterna 2—7 dark brown to black; 8—9 sordid white; cer- ci brown. Genital plate (Fig. 4) elongate; distal plate yellowish, elongate, with slight apical emargination; basal plate absent; spermathecal apodemes thick, dark brown, extending into and filling apical emargina- tion of distal plate, slightly divergent ba- sally with area between partially sclero- tized, free length one-half length of plate. Measurements: Holotype body length approximately 12 mm. Holotype fore wing length, 12.8 mm. Material examined.—Holotype d, allo- type °, and 25 6 23 @ paratypes, collected from La Polvilla (19°38'N, 100°58’W), 2,135 m elevation, Michoacan, Mexico, July 29, 2001. One 6 paratype, 10 km N of Cheran (19°44’N, 102°1’W) 2,178 m el- evation, Michoacan, Mexico, August 6, 2001. Holotype and allotype deposited in the Florida State Collection of Arthropods (FSCA), Gainesville, Florida; additional paratypes in the California Academy of Sci- ences (CAS), San Francisco, California, Il- linois Natural History Survey (INHS), Champaign, Illinois, the Snow Entomolog- ical Museum (SEM), University of Kansas, Lawrence, Kansas, Texas A & M Univer- sity (TAMU), College Station, Texas, Na- tional Museum of Natural History (USNM), Washington, D.C., and collection of the author. Biology.—Panorpa dividilacinia was found to be abundant, along with the slight- ly less numerous P. azteca, on the top sur- faces of lush herbaceous vegetation in the open shade of the pine forested slopes of the Cordellia Volcanica. Both were ob- served to be active from approximately 9:30 a.m. until 4:30 p.m., in spite of over- cast sky and a light rain. Panorpa serta was not observed to coexist with these insects but was hidden vertically on plant stems among broad-leafed herbaceous vegetation in the dense shade of hardwood forest sev- eral miles away at 6880 ft elevation. Unlike most Panorpa from the eastern United States, but like many Panorpa from Mexi- co, these three species hold their wings to- gether rooflike over their abdomens. Diagnosis.—Panorpa dividilacinia is most similar to P. immaculata in some de- tails of the male genitalia, but superficially resembles P. azteca, P. gladiata, and P. ap- iculata with their generally unmarked wings. However, males of P. dividilacinia may be readily differentiated from these species by the unique shape of the hypo- valves. Further, the basistyles ventral area of fusion is narrowly indented posterome- dially in P. azteca (Byers 1958, fig. 9) and P. apiculata (Byers 2001, fig. 5), but not P. dividilacinia. In the field the insect may be told from P. azteca, with which it coexists, by the glossy black markings on the head. Etymology.—The name is based on a combination of divido, Latin for divided and /acinia, Latin for flap. It is the divided flaplike hypovalves of this species that dif- ferentiates it from P. immaculata (Fig. 5), the species it most resembles. Panorpa mixteca Bicha, new species (Figs. 6-8) Description.—Based on 14 6, 10 2, pinned from alcohol. VOLUME 108, NUMBER 1 iA Figs. 6-8. Panorpa mixteca, paratypes. 6, Male right forewing. 7, Male genital bulb, ventral aspect. 8, Female genital plate, ventral aspect (posterior end at top). Head: Dorsum glossy black. Rostrum yellowish brown; genae yellowish brown. Labrum and maxillary palps brown. Anten- nal scape brown, pedicel dark brown to black, flagellum black, with 40—41 flagel- lomeres. Thorax: Pronotum black, pale yellowish along posterior margin; 8—14 black setae at each side on anterior margin. Mesonotum and metanotum black with yellowish white median band, with numerous short, black hairs directed caudad. Scutellum yellowish -white. Pleural surfaces, coxae and mera sordid white, with numerous short, dark se- tae on anterior coxa and on mesothoraxic coxa and episternum; sparse, pale setae on metathorax, most numerous on coxa; small black spot at each end of mesepimeron and metepimeron. Femora and tibiae sordid white with numerous short, apically-direct- ed, black hairs and a lesser number of black setae. Tibiae with two brown spines. Tarsi sordid white with numerous apically-direct- ed golden, and dark brown hairs. Wings (Fig. 6) faintly tinged with yellow, slightly iridescent, immaculate in both sex- es, veins pale brown, stigma pale yellowish, thyridium at first fork of M white. 28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Male abdomen: Terga 2—5 dark brown to black; sterna 2—5 sordid white; segment 6 black with some lateral grading to yellow- ish brown posteriorly, long, cylindrical, nar- rowed posteriorly, without anal horn; seg- ments 7—8 short, yellowish orange. Poste- rior edge of tergum 3 with notal organ; ter- gum 4 with small protuberance on anterior edge. Narrow prolongation of sternum 9, undivided, extremely short, extending only half length of ventral fusion of basistyles (Fig. 7). Tergum 9 narrowing apically with apical emargination; cerci brown. Basisty- les broadly joined ventrally. Dististyles with outer margin slightly concave until curved acute tip; with large, cuplike basal lobe pro- longed ventromesally; dorsomesal edge of basal cup with angular tooth. Dorsal para- meres cylindrical, straight, extending to ba- ses of dististyles; ventral parameres un- branched; apices slightly curved mesad, aculeate, with one mesal “‘barb”’ on apical one-third; sometimes extending slightly be- yond the bases of dististyles into basal cup. Dorsal valves small; ventral valves conspic- uous, conical, laterally compressed, apices slightly curved ventrally, nearly reaching base of dististyles. Female abdomen: Terga and sterna 2—7 dark brown to black; 8—9 sordid white; cer- ci brown. Genital plate (Fig. 8) elongate, narrow; distal plate parallel, apex with deep emargination forming two thin lateral lobes; basal plate absent; spermathecal apodemes extending into apical emargination of distal plate, slightly divergent basally with area between partially sclerotized, free length one-half of genital plate. Measurements: Holotype: body length approximately 12 mm fore wing length 11.5 mm. Material examined.—Holotype CAS): Food plant.—Juniperus sp. Discussion.—Two specimens, a male and female, from St. George, Utah, are the same as the holotype of S. juniperi. Though it is possible that the absence of vein 2r in the forewing is an anomaly, both specimens from Utah also lack this vein adding some credibility to this character. In the female from Utah, the lancet is the same as S. jun- iperi, but the mesonotum is entirely red. The lack of vein 2r is the basis for associ- ation of the male. Unlike males of other species of Susana, it is entirely black with the tibiae and tarsi white. Another male from the same locality in Utah and the same color as the male asso- ciated with S. juniperi has vein 2r (as “‘un- associated”’ in the preceding key and listed in unassociated males) and is possibly S. jJuniperi or another species. Because that specimen has vein 2r, as do some other sim- ilarly colored unassociated males from Ar- izona and New Mexico, it probably repre- sents the unknown male of another species. Susana marin Smith, new species (Figs. 4, 7) Female.—Length, 7.7 mm. Antenna and head black. Thorax orange with posterior margin of pronotum and mesonotal lateral lobes black. Abdomen black with faint red- dish areas on lateral sides of first four seg- VOLUME 108, NUMBER 1 7 21 22 Figs. 14-22. Susana spp., sheaths and male genitalia. 14, Female sheath of S. cupressi. 15, Female sheath of S. oregonensis. 16, Female sheath of S. rufa. 17-22, Male genitalia. 17, Ventral view of left half of genital capsule and lateral view of penis valve of S. cupressi. 18, Ventral view of left half of genital capsule and penis valve of S. annulata. 19, Penis valve of S. oregonensis. 20, Penis valve of S. diablo. 21, Penis valve of S. bakeri. 22, Penis valve of S. juniperi. 12: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ments. Legs black with femora, except ba- ses, and extreme base of tibiae orange. Wings hyaline; veins and stigma black. Mesopleuron with punctures, punctures on central part separated by short interspac- es, less than diameter of punctures (Fig. 4) Forewing with vein 2r. Sheath truncate in lateral view (as in Fig. 15). Lancet (Fig. 7) with 11 annuli; teeth on annuli 1—3 distinct; annuli | and 2 closer together than annuli 2 and 3; serrulae truncate at apices, central serrulae with 3 or 4 posterior subbasal teeth, apical serrulae with 5—7 small pos- terior subbasal teeth. Male.—Unknown. Holotype.—Female, labeled *‘Calif: Ma- rin Co., nr. Woodacre, Carson Ridge, 1—22— Si, BaJAdelison Collector, ~Cupres= sus goveniana”’ (UCB). Etymology.—Named from the type lo- cality, a noun in apposition. Food plant.—The holotype was collected from Cupressus goveniana Gordon. Discussion.—This species is separated from S. cupressi and S. punctata, the other species with a punctate mesepisternum, by the red mesonotum with the lateral lobes black and the lancet with large annular teeth, with 3—6 large posterior subbasal teeth on the serrulae, and with annuli 2 and 3 farther apart than distance between annuli 1 and 2 (see Figs. 5—7). Susana oregonensis Smith (Bisse Oo rs. 19) Susana oregonensis Smith 1969: 21. &. (Klamath Falls, Oregon; USNM). Diagnosis.—Mesepisternum shining, without punctures. Hind femur orange. Sheath in lateral view truncate at apex (Fig. 15). Lancet (Fig. 9) with about 11 annuli; basal annuli perpendicular to ventral margin of lancet; serrulae truncate at apices, with 4 or 5 posterior subbasal teeth. Male with thorax black; abdomen black with segments 3—5 orange, 5th sometimes black posteri- orly; hind femur orange; penis valve (Fig. 19) curved, with long valvispina, as long as half width of valve, and with large protrud- ing lobe adjacent to valvispina. Distribution.—Oregon east of the Cas- cade Mountains south to Fresno and Mono counties, California; eastern Nevada and southern Utah. Material examined.—CALIFORNIA: Cedar Pass, Warner Mtns., Modoc Co., 6 June 1978, collected on Rosaceae, L. G. Bezark (1 2, 1 6 ; BEZ); Coalinga, III-19— 1940 (1 2, UK), 3-20-1940 (1 2, UK); Lake Eiler, Shasta Co., 9 July 1947 (1 9, UK); Occidental, Sonoma Co., V-19—57 (1 3d, UCB); Sardine Creek, Mono Co., VI-27- 57 (1 6; UCB); Huntington” Lk, Fresno Co:., 7,000 6;-CAS) July Melo eyo= semite Val., VI-27—1921 (1 2: CAS); Glen Algqina, Tahoe, June 23, 1929 (1 2; CAS); Cedar Ridge, Alameda Co., V-3-31 (1 9: CAS); 7 mi W. Hat Creek PO; VilIE3=55; Cupressus macnabiana (1 2, 136; USNM). NEVADA: Modena Summit, Lincoln Co., V-29=19672"6; 719" >" R. '©BechteliGiae NDA). OREGON: Redmond, June 1, 1968, K. Goeden, sticky board trap juniper tree (2 6, ODA, USNM); 2 mi NNE Simnasho, Wasco Co., 1-VI-68, K. Goeden, sticky board trap juniper tree (2 ¢ ODA); Sim- nasho, Wasco Co., June 10, 1963, beating juniper, Ken Goeden (larvae, USNM,; only species in area and adults collected from same locality); 6 mi W. Juntura, 13-VI-68, K. Goeden, Juniperus occidentalis (1 6, USNM); 15 mi NW Warm Springs, Wasco Co., June 2, 1968, K. Goeden, sticky board trap juniper tree (1 d, ODA); Klamath Falls, VI-16—22 (1 2, 1 6; CAS) May 13, ‘24(1 2; CAS). UTAH: Mt. Carmel, V-30- 35, ne Zion Cyn G. 2, CAS): St. Georse V-28-35 (2 2, CAS); Kane Co., Grand Staircase—Escalante Nat’! Mon., Seaman Wash, near spring, N. of Hwy 89, 37°07 02"N, 112°14’59"W, 14 May 2001, H. Barber, R. Lorimer, S. E. Morrison (1 @, BYU). Food plants.—Juniperus occidentalis Hook. Possibly another species of Junipe- rus in southern Utah. A female and a male VOLUME 108, NUMBER 1 were collected from Cupressus macnabiana A. Murray in northern California. Discussion.—The structures of the fe- male lancet and male genitalia are the most reliable for identification of this species. All specimens have a similar lancet, similar male genitalia, and an orange hind femur. Additionally, the hosts are of species of Juniperus. Coloration of the thorax and abdomen varies throughout its range. In the female, the holotype of S. oregonensis, as well as most specimens from eastern Oregon and northern California have the mesoprescu- tum and mesoscutellum orange, mesonotal lateral lobes black, mesepisternum with a large orange spot, and abdomen mostly or- ange with the basal plates black and some black on the apical terga. Most specimens from central California, Nevada, and Utah have the mesonotum and mesepisternum entirely black and the abdomen black at the base and apex with a central orange band. A few from eastern Oregon and California have intermediate color varieties between these extremes. The male has a more uni- form coloration throughout its range. All specimens have the thorax black and a cen- tral orange band on the abdomen. A female from Hat Creek, Shasta Co., California, has the thorax entirely black but a male with the same label data is typical for S. orego- nensis. The leg coloration remains constant in all specimens. Susana punctata Smith (Figs. 3, 6) Susana punctata Smith 1969: 19. 2 (7 mi. EP bie Pine, Inyo "Co.~ California: USNM). Diagnosis.—Mesepisternum with punc- tures (Fig. 3). Black with tegula, posterior margin of pronotum, apex of each femur, fore- and mid tibiae, basal half on hind tib- ia, and line on posterior margin of each ab- dominal segment white. Sheath in lateral view with apex truncate (as in Fig. 15). Lancet (Fig. 6), with 12 annuli; annular 13 teeth large, annuli 5 and 6 with 6 or 7 teeth; serrulae truncate at apices, with 2 large pos- terior subbasal teeth on central serrulae. Male unknown. Discussion.—This species still is known only from the holotype. Susana rufa Smith (Figs. 12, 16) Susana rufa Smith 1969: 21. 2 (5 mi. W. Portal ‘Chiricahua’? Mitsis “Arizona: USNM). Diagnosis.—Mesepisternum smooth, shining, without punctures. Thorax orange with mesosternum black; abdomen orange; hind femur black. Forewing with vein 2r. Sheath in lateral view straight above, rounded below (Fig. 16). Lancet (Fig. 12) with 11 annuli, basal annuli oblique; apices of serrulae truncate, serrulae with 3 or 4 large posterior subbasal teeth; annular teeth large with only about 6 teeth on annuli 3— 6. Male unknown. Discussion.—This species still is known only by the holotype and one paratype from mNEIviexe UNASSOCIATED MALES Several males were studied that cannot be identified. Because the taxonomy is based on females and males are not known for all species, these may represent males of species known only from females or un- known species. Some, however, represent significant records for the genus, and there- fore are recorded here. ARIZONA: Williams, 27-5, Barber & Schwarz coll. (1 USNM). CALIFORNIA: Mojave Desert, V-1939, R. H. Smith (1, LACM); Jacumba, San Diego Co., IV-25- 51 (1, UCB). NEW MEXICO: Torrance Co., Town of Gran Quivira, 9-VIII-65, H. B»Leéch 4) CAS)? UTAH St.. George CE CAS). The specimens from Williams, AZ, Tor- rance Co., NM, and St. George, UT, are black with the tibiae white, have vein 2r in the forewing and correspond to “‘unidenti- 74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fied’? in the preceding key. The specimen from Jacumba is similar to S. oregonensis, but its distant distribution makes identifi- cation questionable. LARVAE Larvae have been examined from various localities and hosts. Smith (1969) charac- terized the larvae of Susana and described those of S. cupressi, S. juniperi, and S. or- egonensis. Wong and Milliron (1972) de- scribed the larva of S. fuscala. Since most have not been associated with adults, larvae cannot be identified. The following records, which are not presented in the species treat- ments, are given because they represent some additional significant food plants and distribution records for the genus. ARIZONA: Yavapai Co., Prescott, VI-19- 73, on Arizona cypress (ADA); Cochise Co., Carr Canyon (1 mi. W. Stat. Hwy 92) near Nicksville, 31-VII-1991, sweeping Juniperus (USNM); Prestcott, Tenopai Co., VI-10-73, on Arizona cypress (USNM); Cochise Co., San Simone, [V-12-72, on Cu- pressaceae (USNM). CALIFORNIA: Im- perial Co., Brawley, larvae on blue Italian Cypress in nursery, 4-11-66 (CDA); Porter- ville, 4-25-66, larvae on cypress (CDA); San Bernardino Co., Chino, 6-1-67, larvae on Hollywood juniper (CDA); Santa Cruz Co., 6-19-67, on Arizona cypress (CDA); Tulare Co., Visalia, on leaf of arborvitae, Thuja plicata (CDA); Stanislaus Co., Em- pire, 5-6-74, on spruce (CDA); Yuba Co., Marysville, 5-22-03, on Italian cypress (CDA); Santa Barbara Co., Highland, Jun- iperus chinensis ‘“Torulosa,”’ ITV-12-1967 (USNM); Woodside, June 2, 1950, Italian cypress (CDA, USNM); San Jose Univer- sity campus, Santa Clara Co., Sept. 1978, ex cypress (USNM); Riverside, April 29, 1953, on Juniperus (USNM); Saticoy, Ven- tura Co., IV-25-1950; Tecate. cypress (USNM); San Diego Co., Escondido, 5-23- 1967, cypress (USNM); Pasadena, 5-16-29, on cypress (USNM). Foop PLANT SUMMARY The following list of food plants are from rearing data, larval collections, and data on specimens which may be only adult collec- tion records. Arborvitae, may refer to a species of Thuja; unassociated larvae. Cupressus arizonicus Greene, Arizona cy- press; Susana cupressi (adult collection); unassociated larvae. Cupressus forbesii Jeps., Tecate cypress; unassociated larvae. Cupressus goveniana Gordon, Gowen cy- press; Susana marin (adult collection). Cupressus macnabiana A. Murray, Mac- Nab cypress; Susana oregonensis (adult collection). Cupressus macrocarpa Hertweg, Monterey cypress; Susana annulata (adults and lar- vae); Susana cupressi (adults and larvae); unassociated larvae. Cupressus sempervirens L., Italian cypress; Susana annulata (adults and larvae; Su- sana cupressi (adults and larvae), unas- sociated larvae. Juniperus chinensis L., Hollywood juniper; unassociated larvae. Juniperus occidentalis Hook., western ju- niper; Susana oregonensis (adults and larvae). Juniperus scopulorum Sarg., Rocky Moun- tain juniper; Susana fuscala (rearings). Thuja plicata Don ex D. Don, western red- cedar; unassociated larvae. ACKNOWLEDGMENTS I appreciate the help of the curators of the following institutions for loan of spec- imens for this study: J. Sorensen (CDA); R. R. Snelling (LACM); R. W. Brooks (UK); H. Goulet (CNC); W. Pulawski (CAS); W. W. Middlekauff (UCB); Arizona Depart- ment of Agriculture, Phoenix (ADA); R. L. Westcott (ODA); Nevada Department of Agriculture, Reno (NDA); R. Baumann (BYU); and L. Bezark, Sacramento, CA. H. R. Jacobson, Chico, California, provided specimens of S. cupressi and S. annulata as VOLUME 108, NUMBER 1 well as his observations on the food plant and collections of the two species. Cathy Apgar, Systematic Entomology Laboratory, USDA, took the photographs and prepared the plates. I thank the following for review of the manuscript: N. M. Schiff, U.S. Forest Service, Stoneville, MS, and S. J. Scheffer and M. A. Solis, Systematic Entomology Laboratory, USDA, Beltsville, MD, and Washington, DC, respectively. LITERATURE CITED Rohwer, S. A. 1911. New sawflies in the collections of the United States National Museum. Proceed- WS ings of the United States National Museum 41: 377-411. Rohwer, S. A. and W. Middleton. 1932. Descriptions of five Nearctic sawflies of the tribe Hemichroini. Proceedings of the Entomological Washington 34: 93-98. Ross, H. H. 1937. A generic classification of the Ne- arctic sawflies (Hymenoptera: Symphyta). Illinois Biological Monographs 15, 173 pp. Smith, D. R. 1969. The genus Susana Rohwer and Middleton (Hymenoptera: Tenthredinidae). Pro- ceedings of the Entomological Society of Wash- ington 71: 13-23. Wong, H. R. and H. E. Milliron. 1972. A Canadian species of Susana on western juniper (Hymenop- tera: Tenthredinidae). Canadian Entomologist 104: 1025-1028. Society of PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 76-81 DESCRIPTION OF THE LARVA OF GOMPHIDICTINUS PERAKENSIS (LAIDLAW) (ODONATA: GOMPHIDAE), WITH DISTRIBUTIONAL NOTES MICHAEL L. FERRO AND ROBERT W. SITES Enns Entomology Museum, Department of Entomology, University of Missouri, Co- lumbia, Missouri 65211, U.S.A. (e-mail: spongymesophyll@gmail.com and SitesR @ missouri.edu). Abstract.—The final instar of Gomphidictinus perakensis (Laidlaw) is described and figured from exuviae and larval specimens collected in Chiang Mai, Kanchanaburi, Pra- chuap Khiri Khan, and Surat Thani provinces in Thailand. This large species is flattened, subovate, and the posterolateral corners of abdominal segments III-IX have an elongate, broadly-recurved flange. Distributional information is given concerning additional collec- tions of adults, larvae, and exuviae from Thailand. Key Words: The Odonata fauna of Thailand has been documented more completely than has any group of insects in Thailand, and might rep- resent the best understood fauna of an order of insects of any Indochinese country. Spe- cifically, this order was the focus of a 21- part series, which included descriptions, il- lustrations, and taxonomic keys to most of the known species of adults (Asahina 1993). Recently, the current state of Thai odonatology was summarized and 315 spe- cies were reported to occur in the country (Haimaléinen and Pinratana 1999), The known Thai fauna of Gomphidae is represented by 21 genera, including the monotypic genus Gomphidictinus Fraser. Laidlaw (1902) described Gomphidia per- akensis from one male from Perak, Malay- sia and subsequently recorded two addition- al males from that region (Laidlaw 1931). Fraser (1942) described a new genus and species, Gomphidictinus wheeleri, from the Federated Malay States. Fraser’s genus and species were synonymized with Gomphidia perakensis by Lieftinck (1954). However, Gambles (1987) regarded Gomphidictinus Odonata, Gomphidae, Gomphidictinus, larvae, Thailand as a valid genus, including only the one species, G. perakensis with its synonym G. wheeleri. Gomphidictinus is currently rec- ognized as a valid genus, although some scientists prefer to retain the species in Gomphidia (e.g., Asahina 1993). Gomphidictinus perakensis was first re- ported from Thailand based on two speci- mens collected in 1965 from “Fang N. Thailand’ (Asahina 1981). Eight more specimens were reported from Thailand, each near running streams in “‘deep jungle” (Asahina 1986). Despite the two separate descriptions and multiple records of adult G. perakensis from Malaysia and Thailand, no record of immatures exists, except the inference that the adults near streams in “deep jungle” (Asahina 1986) likely occur as immatures in those streams. Herein, we present the original descrip- tion of the larva of G. perakensis (Laidlaw) from exuviae, one of which was associated with a recently emerged adult male. MATERIALS AND METHODS All measurements were taken in the dor- sal view using an ocular micrometer and VOLUME 108, NUMBER | represent maximum values unless otherwise stated. The head was measured from the an- terior margin of the clypeus to the posterior margin of the postocciput and the abdomen from the base to the tip of the caudal ap- pendages. Width of the abdomen was de- termined by measuring the distance be- tween the tips of the posterolateral flanges of segment VII. The outer surface of the leg was considered the dorsal aspect. Mid-dor- sal and mid-ventral abdominal lengths were measured from anterior to posterior margins of a sclerite, excluding dorsal hook, if pres- ent. Posterolateral flange width was mea- sured in ventral view from the intersection of the posterior margin with the anterolat- eral corner of the succeeding sternum to the base of the apical spine. In some specimens, the spine of the flange was broken off. Cau- dal appendages were measured from the posterior margin of tergum X to the tip of the appendage. Each site sampled was giv- en a locality number (L-number). Photos of each locality can be viewed at the Locality Image Database link of the Enns Entomol- ogy Museum internet site. Specimens have been deposited in the Entomology Museum, University of Mis- souri—Columbia, and the National Science Museum, Pathum Thani, Thailand. Genetic material has been deposited in the tissue collection at Brigham Young University, Provo, Utah. SYSTEMATICS Gomphidictinus perakensis (Laidlaw) (Figs. 1—4) Gomphidia perakensis Laidlaw 1902: 81— 82, figs.—Williamson 1907: 281-282 (taxonomic notes).—Laidlaw 1931: 208.—Fraser 1942: 99-101, figs.—Lief- tinck 1954: 80.—Asahina 1981: 8.—Asa- hina 1986: 43—45, figs. Gomphidictinus perakensis: Gambles 1987: 51—60.—Pinratana et al. 1988, fig —Hamilainen and Pinratana 1999: 77 (Thailand distribution). Gomphidictinus wheeleri Fraser 1942: 88— 101, figs. Synonymy by Lieftinck 1954. 77 Description.—Length, 28.0 mm; width, 15.8 mm. Overall body shape elongate-oval (Fig. 1). Color generally fuscous with areas of darker brown. Ventral surface covered with fine setae. Head: Length, 4.7 mm; width, 7.8 mm. Posterolateral corner with protruberance projecting slightly above level of eye, cov- ered with stout, dark spines and elongate, light brown setae. Clypeus extending to two-fifths length of third antennal segment, labrum extending to apex of antennae. Compound eyes emarginate laterally. An- tenna concolorous with body; segments I and II subspherical, segment III elongate, slightly expanding toward apex and bowed mesad. Ratio of antennal segment lengths 15:8:55:3. Prementum in ventral aspect gently expanding apically (Fig. 2), covered in short, recumbent, scalelike setae; mid- ventral length 3.5 mm, width at middle 4.2 mm; ligula evenly convex, crenate, with dense fringe of long hairlike setae. Setal fringe with lateral one-fourth twice as long as middle half. Blade (first segment of la- bial palp) cuspidate apically with ca. 10 short, nearly obsolete, squarely-truncate teeth along mesal margin; tooth width and obsolescence increasing apically (Fig. 3). Thick, recurved spines and elongate setae on elevated processes mesad to antennal ba- ses and on anterolateral shelves of vertex laterad of antennal bases. Glabrous portions of head include clypeus except lateral cor- ners; broadly lyriform area on vertex mid- line; broad band bordering mesal margin of compound eye and lateral ocellus; median elongate oval and lateral “‘U”’ shaped areas on ventral deflection of vertex; area form- ing right angle anterior and mesad to pro- truberance of posterolateral corner. Thorax: Dorsally, medium brown with areas of dark brown pigmentation dorsad of procoxal base, forming inwardly curved longitudinal stripes bordering midline of pronotum; ventral edges of meso- and me- tapleuron; midline of mesonotum: costal margin of wingpads. Thick, black, recurved spines on ventroanterolateral and dorsopos- 78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Gomphidictinus perakensis larva. 1, Dorsal view of larva. 2, Ventral view of labium. 3, Labial palp. 4, Lateral view of abdominal segments VI-X and terminal appendages. VOLUME 108, NUMBER 1 terolateral corners of pronotum and ridge of meso- and metaepimeron. Smaller spines on anterior and posterior median humps of pronotum. Rest of-thorax with various de- grees of spinosity. Femora of all legs (most pronounced in profemur) and pro- and me- sotibia with dark band on either side of middle. All femora and tibiae with longi- tudinal rows of setae. Setae on inner surface of tibia elongate and spinelike, other setae scalelike. Length of hind femur 7.0 mm, apex attaining middle of abdominal seg- ment VI. Last tarsal segment of all legs pale in basal third, dark in distal two-thirds. Length of meso- and metathoracic wing- pads 8.2 and 8.7 mm respectively, attaining ca. posterior margin of abdominal segment VI. Wingpad costal margins spinose, else- where glabrous. Wingpads subparallel in larvae (appear divergent in exuviae because of ecdysial fracture). Abdomen: Length, 15.0 mm; maximum width, 15.8 mm. Ovate, triangular in cross section, shurikenate, with well developed mid-dorsal ridge (Fig. 4). Dorsally, with lat- eral margins of segments I-VI densely se- tose, segments VII-IX with stout recurved spines and sparsely setose. Posterior margin of tergites I-VIII with row of short spines except beneath wingpads. Area beneath wingpads shiny with scattered, fine, wisp- like setae. Scattered setae on mesal half of tergites VII-IX, becoming denser posteri- orly on VII; lateral half sparsely setose. Ter- gites III—-VIII with transversely-elongate oval area surrounded by bifurcated row of short spines along posterior margin halfway between mid-dorsal ridge and lateral mar- gin. Mid-dorsal length of tergites I-X 0.8, OOO 5rle22 le 121042.42:25 1.8, and 0.8 mm, respectively. Mid-dorsal ridge well de- veloped (Fig. 4), segment I without process; segment II digitate; segments III—V increas- ingly longitudinally expanded and apex posteriorly directed; segments VI-IX gry- phoform, each ridge extending posteriorly as rounded spine slightly overlapping suc- ceeding segment. Posterolateral corners of abdominal segments III-I[X with elongate, 79 broadly recurved flange. Corner of flange pale, tipped with stout elongate dark spine; each spine encircled with veil of golden se- tae. Flange with spine lengths on segments PRESEN, O57 LZ a2 Ou 2a) 2:3%cand. 333 mm, respectively. Posteriorly directed lat- eral flanges of abdominal segment IX ex- tend to, or slightly beyond, tips of cerci. Epiproct length 1.5 mm, slightly shorter than paraprocts. Cercus length 1.4 mm. Distribution.—Gomphidictinus peraken- sis has been collected from nine provinces scattered throughout the northern, north- eastern, southeastern, and peninsular re- gions of Thailand (Haémalainen and Pinra- tana 1999), and it occurs as far south as Malaysia (Laidlaw 1901, 1931; Fraser 1942). The exuvial nepionotype was col- lected in Prachuap Khiri Khan Province and represents the northernmost record of G. perakensis in the peninsular region of Thai- land. One adult was collected in Phang Nga Province on 28 May 2003, and one larva from Surat Thani Province on 24 May 2005, representing the first records of G. perakensis from these provinces. Additional larvae and exuviae were colleted in Kan- chanaburi and Chiang Mai provinces, from which the species has already been record- ed. Exuvial nepionotype.—THAILAND: Prachuap Khiri Khan Prov., Amphur Bang Saphan, stream from Kha On Waterfall, L- 544, 11°26.351'N, 99°26.052'E—WGS84, elev. 117 m, 18 May 2003, MLE 1 exuviae. The exuviae was collected from the upper surface of a broad leaf ca. 30 cm from the ground and ca. 50 cm from the stream edge. No other exuviae was in the vicinity. The teneral imago was hanging from an adja- cent leaf and did not resist capture. Additional material examined.—THAI- LAND: Kanchanaburi Prov., Amphur Thong Pha Phum, Heuy Ou Long, 14°46.922'N, 98°40.165'E—WGS84, elev. 124 m, 12 May 2003, M. L. Ferro, L-525, 1 exuviae; Surat Thani Prov., Khao Sok National Park, stream from Sipet Chan Waterfall, elev. 42 m, 24 May 2005, 08°54’N, 98°31’E— 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON WGS84, R.W. Sites & T.O. Prommi, L-795, 1 larva; Chiang Mai Prov., Mae Kuarg, 19°00.015’N, 99°17.163'E—WGS84, elev. 521 m, 15 May 2004, A. Vitheepradit, L- 689, | larva; Chiang Mai Prov., Doi Suthep- Pui National Park, Monthathan Waterfall, 18°49'N, 98°55'E—WGS84, elev. 700 m, 19 May 2004, A. Vitheepradit, L-697, 1 larva. Diagnosis.—This species can be recog- nized by its large size, well developed mid- dorsal ridge, and extremely well developed posterolateral flanges of abdominal seg- ments IHI-IX. Because Gomphidictinus is a monotypic genus considered to be closely related to Gomphidia (Gambles 1987), comparisons should be made with species of Gomphidia. The larva of Gomphidia kruegeri Martin, a species known from Chi- na and Thailand, was described and illus- trated by Needham (1930). The larva of Gomphidia kruegeri is slightly larger (length 30 mm, width 18 mm) than that of G. perakensis, otherwise the written de- scriptions are in accord. However, charac- teristics evident in Needham’s illustrations are sufficient to distinguish the two species: the setal fringe of the ligula with lateral one-tenth twice as long as middle, as op- posed to lateral one-fourth in G. perakensis; posterolateral corners of abdominal seg- ments III-[X with much reduced recurved flanges compared to those of G. perakensis; and the posteriorly-directed lateral flanges of abdominal segment IX extend to half the length of the cerci, as opposed to as long as the cerci in G. perakensis. DISCUSSION Gomphidictinus perakensis has been re- ported only from Thailand and Malaysia (Asahina 1986). The two adults collected from this research are new provincial rec- ords (see Haimiléinen and Pinratana 1999). Both specimens were collected at swiftly flowing, intermediate-disturbed, sandy bot- tomed streams with occasional boulders and emergent vegetation (Location-544 and L- 577). Each stream was largely surrounded by forest, which is consistent with the hab- itat description of the adult given by Asa- hina (1986). Larvae were collected from Monthathan Waterfall, Mae Kuarg, and Sipet Chan Wa- terfall. Each larva is dimensionally and pro- portionally nearly identical to the exuviae described above, including wingpad length, and therefore considered to be a Gomphi- dictinus perakensis final instar. Structurally, the larvae and exuviae are identical, al- though the color pattern differs slightly as follows: Larva with greater contrast be- tween the dark bands and background col- or; transversely-elongate oval area sur- rounded by a bifurcated row of short spines along posterior margin halfway between mid-dorsal ridge and lateral margin of ter- gites HI—VIII more distinct and darker than exuviae; single glabrous oval cephalad to the bifurcated row of short spines on seg- ments IV—VIII is very distinct. One well-developed larva (Mae Hong Son Prov., L-306) and one exuviae (Pra- chuap Khiri Khan Prov., L-540) were col- lected which are morphologically similar to the larvae and exuviae of G. perakensis, with some notable exceptions. The relative lengths of the recurved flanges on abdom- inal segments VI and VIII of G. perakensis are 74% and 85% the length of the flange on segment VII, respectively. However, the relative lengths of the recurved flanges on segments VI and VIII are much shorter in the larva (L-306) and exuviae (L-540), and are only 30% and 43% (L-306) and 75% and 70% (L-540) the length of the flange on segment VII, respectively. Additionally, the transversely-elongate oval area sur- rounded by a bifurcated row of short spines along the posterior margin halfway between the mid-dorsal ridge and lateral margin of tergites III—VIII is much reduced in these specimens compared to those of G. perak- ensis. Because complete formal descrip- tions are lacking for the larvae of Diasta- tomma and Gomphidia (except G. krue- geri), Which are thought to be closely re- lated to Gomphidictinus (Gambles 1987), and the degree of interspecific variation in VOLUME 108, NUMBER 1 G. perakensis is unknown, we have not as- signed identities to the specimens from L- 306 and L-540. Two months of sampling throughout Thailand by MLF yielded odonate exuviae at nearly every locality sampled, and ap- proximately 350 exuviae were collected. Although zygopteran exuviae were scarce in general, most were of Calopterygidae and Euphaeidae. Anisopteran exuviae were much more common, especially the Aeshn- idae, Gomphidae, and Libellulidae. Few ex- uviae were weathered, and most appeared to have been left only the night before col- lection. This suggests that rearing, emer- gence traps, and observation of rocks and vegetation at night or early morning may yield many and diverse adult/immature as- sociations in a short period of time. ACKNOWLEDGMENTS We thank T. W. Donnelly for determina- tion of specimens. We are grateful to Ake- kawat Vitheepradit for logistical help in the field and collection of specimens. We also thank Penkhae Thamsenanupap and Pensri Bunlue (Chiang Mai University) and Taeng-On Prommi (Prince of Songkla Uni- versity) for assistance with field work. We are grateful to the National Research Coun- cil of Thailand and the Royal Forestry De- partment for permission to conduct field work in Thailand. Funding for MLF was provided by the Dorris D. & Christine M. Brown Graduate Research Fellowship. Sup- port for RWS was provided in part by NSF project number DEB-0103144 and by MU project number PSSLO232. LITERATURE CITED Asahina, S. 1981. Records of little or unknown Odon- ata from Thailand. Tombo 23(1/4): 3-16. . 1986. A list of the Odonata recorded from Thailand, Part XIV. Gomphidae—2. Tombo 29(1/ 2): 7-53. . 1993. A list of the Odonata from Thailand (Parts I-XX1). Bangkok, Bosco Offset. ca. 460 pp. [No consecutive pagination; contains copies of 430 papers from S. Asahina]. Fraser, E C. 1942. Dr. Raymond Wheeler’s collection of the Odonata from the Federated Malay States, with the description of a new genus and two new species. Proceedings of the Royal Entomological Society of London (B) 11(7): 95-104. Gambles, R. M. 1987. Hagen’s tubercle, a largely over- looked but potentially useful character in gomphid taxonomy (Anisoptera: Gomphidae). Advances in Odonatology 3: 51—60. Hamialdinen, M. and A. Pinratana. 1999. Atlas of the Dragonflies of Thailand: Distribution Maps by Provinces. Brothers of St. Gabriel in Thailand, Bangkok, 176 pp. Laidlaw, EF EF 1902. On a collection of dragonflies made by members of the Skeat Expedition in the Malay Peninsula in 1899-1900. Proceedings of the Zoological Society of London 1(5): 63-92. . 1931. A list of the dragonflies (Odonata) of the Malay Peninsula with descriptions of new spe- cies. Journal of the Federal Malay Museums 16(3/ 4): 175-233. Lieftinck, M. A. 1954. Handlist of Malaysian Odonata: A catalogue of the dragonflies of the Malay Pen- insula, Sumatra, Java and Borneo, including the adjacent small islands. Treubia 22 (Supplement): xiii + 202 pp. Needham, J. G. 1930. A manual of the dragonflies of China: A monographic study of the Chinese Odonata. Zoologia Sinica, Invertebrates of China, SCHESHAW lle 359) laters Pinratana, A., B. Kiauta, and M. Hamialainen. 1988. List of the Odonata of Thailand and annotated bibliography. Bangkok, The Virathan Press. 20 pp. + 20 cl. plates. [No pagination]. Williamson, E. B. 1907. The dragonflies (Odonata) of Burma and Lower Siam -II. Subfamilies Cordu- legasterinae, Chlorogomphinae, and Gomphinae. Proceedings of the United States National Muse- um 33: 267-317. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 82-91 DEVELOPMENT OF A BIOASSAY FOR EVALUATING TICK (ACARI: IXODIDAE) ATTRACTANTS IN THE FIELD J. EF CARROLL, V. B. SOLBERG, AND T. L. Cyr (JFC, TLC) U.S. Department of Agriculture, Agricultural Research Service, Animal Parasitic Diseases Laboratory, Beltsville, MD 20705, U.S.A. (e-mail: jcarroll @anri.barc. usda.gov); (VBS) Department of Entomology, Walter Reed Army Institute of Research, Silver Spring, MD 20910-7500, U.S.A. Abstract.—A method for testing tick attractants under field conditions is described. The uneven distribution of host-seeking ticks, even within habitat types, can compromise eval- uations of experimental baits and necessitate extensive testing. This problem can be rem- edied by releasing known numbers of ticks at predetermined locations and distances from an experimental bait. Lone star tick, Amblyomma americanum (L.), nymphs were released directly downwind of dry ice baits on 2.25 * 2.25 m cotton sheets that had been placed on the ground. Each sheet was aligned with the direction of the wind and edged with a masking tape barrier. Best results in discriminating between dry ice baits and untreated controls were with nymphs released 1.5 and 2 m from the bait and recaptured after | h in a zone <1 m from the bait. The success of this method depends on the wind not stopping for prolonged periods and not radically changing direction. Key Words: The efficacy of any tick attractant system is measured by its performance under field conditions. Using natural populations to evaluate a tick attractant in the field can in- volve extensive replication if the tick pop- ulation sampled is not dense. The tendency for clumped distributions among host-seek- ing ticks, even within habitat types, exac- erbates the likelihood that baits will be placed in unproductive locations. For in- stance, Newhouse (1983) found that along park trails in Georgia adult American dog ticks, Dermacentor variabilis (Say), were nonrandomly distributed in clusters that changed annually. Solberg et al. (2003) also found extreme clumping of blacklegged ticks, Ixodes scapularis Say, in study plots in Maryland. If few or no responsive ticks are within the effective range of an exper- imental bait, little is learned from the trial. lone star tick, Amblyomma americanum, carbon dioxide, dry ice An alternative to using natural populations is to seed the area around an experimental bait with marked host-seeking ticks, there- by assuring that potentially responsive ticks will be exposed to the attractant (Falco and Fish 11991): Carbon dioxide (CO,) has been used as an attractant for ixodid and argasid ticks for decades (Garcia 1965, Semtner et al. 1975, Falco and Fish 1991). Holscher et al. (1980) elucidated electrophysiological responses of three species of ticks to CO,. Along with flagging and dragging, dry ice baiting has been a method of choice for collecting many host-seeking ticks and monitoring their populations (Gladney 1978, Koch 1987, Koch and McNew 1981, Solberg et al. 1992). Highly mobile tick species travel considerable distances to CO, sources. Lone star ticks, Amblyomma americanum VOLUME 108, NUMBER 1 (L), have been reported to be attracted to CO, sources from as far as 21 m (Wilson et al. 1972). Although long duration dry ice traps that operate for days have been de- veloped (Gray 1985), most dry ice baiting is done with small blocks that last for only for 1—2 h (Koch 1987). A drawback in dis- tributing dry ice to numerous sample sites in the field is carrying several kilograms on transects or to remote plots in tick habitat and placing the baits before sublimation significantly diminishes the last pieces of Gry Ice. The purpose of this study was to develop a method for testing attractants in the field that used a modest number of released ticks and provided statistically discriminatory data about the efficacy of the bait. Three variations of the basic premise of releasing ticks directly downwind of a bait in the field were tested sequentially in 2003 and 2004 in an attempt to maximize the num- bers of attracted ticks recaptured. We report the basic features and requirements of a tick attractant field bioassay. MATERIALS AND METHODS We conducted one set of trials to dem- onstrate the variation encountered when us- ing natural populations for tick attraction experiments. Three sets of trials were con- ducted using marked-released-recaptured ticks to define the basic elements of a meth- od to reliably test an experimental tick at- tractant under field conditions. Trials with natural tick populations.—To ascertain variation in nymphal densities at the geographic scale at which a tick attrac- tant would operate, trials using dry ice baits were conducted on seven dates during the late spring and summer of 2004 in mesic forests at Fort A. P. Hill, Virginia and the Patuxent Wildlife Research Center, Laurel, Maryland. The predominant species and life stage present were A. americanum nymphs. To increase the likelihood that host-seeking ticks were in attractable range, bait locations were selected 22 h in ad- vance of testing by flagging for host-seek- 83 ing ticks with 0.5 X 0.5 m crib cloth (a laminate of flannel and rubber). At loca- tions where at least four nymphs were cap- tured in five 30-sec bouts of flagging (each roughly linear) that covered in total a ~15 x 15 m area, a marking flag was placed near the center of the flagged area. A 1 X 1 m piece of white nylon sail cloth was spread, as level as possible, on the sub- strate. A 500—550 g block of dry ice was placed on an inverted styrofoam cup (10 cm diam X 5 cm high X 7.5 cm diam) that had been positioned on the center of the cloth. Elevating the dry ice on the cup may aid in dispersal of the CO,, which is heavier than air. As a control, an inverted cup without the dry ice was placed on a separate sheet at a location =75 m distant where =4 A. americanum nymphs were found by flag- ging. After 2 h, ticks on the upper and un- der sides of the cloths were collected on transparent tape and affixed in a notebook for subsequent identification and counting. A Bacharach CO, Analyzer, Model 2815 (New Kensington, PA) was used to obtain measurements of CO, in the air at ground level. Relative humidity (RH) and temper- ature on the upper surface each sheet were measured with a digital hygrothermometer (Model EA25, Extech Instruments, Wal- tham, MA). A total of 11 pairs of treatment and control sheets was tested. Trials with marked-released-recaptured ticks.—The A. americanum nymphs used in the mark-release-recapture trials were from a laboratory colony at United States De- partment of Agriculture, Agricultural Re- search Service, Knipling-Bushland U. S. Livestock Insects Research Laboratory, Kerrville, Texas. The ticks were maintained at 24°C, ~97% RH and 16:8 h (L:D) until used in the trials. Trials were conducted on large (2.25 X 2.25 m) white cotton sheets (45 45 thread mesh/ cm). Each sheet was marked with three arcs having radii of 0.3, 1.0 and 2.0 m from one corner of the sheet, a pattern referred to hereafter as Configuration 1 (Fig. 1). A marking flag was placed into the 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON wind direction Fig. 1. release points Diagram showing 2.25 * 2.25 m sheet with bait location, recapture zones, tick release points for Configuration 1, and alignment with prevailing wind at start of attractant bioassay. The sheet was edged with masking tape to confine released ticks and exclude ticks from natural population. ground to monitor wind direction. The cor- ner of the sheet nearest the 0.3 m arc was placed at the base of the flag stake, and the sheet was adjusted so that the diagonal be- tween the flag corner the opposite corner aligned with the direction of the wind, as indicated by the flag (Fig. 1). Masking tape (7.5 cm wide) was affixed to the edges of the sheets with the sticky side facing up and a 3—5 cm wide band of sticky surface ex- posed. At each corner, a flag stake was pushed through the tape into the ground to prevent billowing and folding over of the sheet. Two additional stakes were pushed through the tape on each upwind margin of the sheet, where there was a greater tenden- cy for it to fold over in a strong breeze. As dictated by change in wind direction, the cloths were reoriented for each trial before securing them to the ground. Configuration 1, Experiment |. A plastic box @6.7 by wl'6:5 cm tops Z1-0iby 33cm bottom, 10.2 cm high) with a slit (15.6 by 0.6 cm long side, 8.5 by 0.6 cm short side) on each side, 2 cm above the bottom was placed in the upwind corner of the sheet, as close as possible to the masking tape edg- ing. Dry ice (300—350 g) was placed on top of an inverted styrofoam cup inside the box. For the negative control, a box containing an inverted cup was placed on the upwind corner of the sheet. Ten nymphs marked VOLUME 108, NUMBER | with colored dust (Switzer Brothers, Cleve- land, OH) were released from a holding vial 1.01 m downwind from the corner of the sheet with the bait (dry ice or control), on the diagonal between the upwind and downwind corners (Fig. 1). Immediately thereafter a second group of 10 nymphs, marked with a different color dust, was re- leased on the diagonal 2.01 m downwind from the bait corner. The placement of each color group at 1.01 or 2.01 m from the up- wind corner was randomly determined for each sheet. One hour after the ticks were released, they were recaptured on pieces of 2.5-cm wide masking tape. The locations of their recapture were recorded as one of four zones referenced to the upwind corner: Zone | (<0.3 m), Zone 2 (0.3—1.0 m), Zone 3 (1.0-2.0 m), Zone 4 (>2.0 m). All tests were conducted at the same location, a nar- row (5 m) grassy strip between a hedgerow of mature deciduous trees and a hay field. Dry ice and an untreated control were test- ed sequentially in random order (<1 h apart) on each of 4 d. The dry ice was weighed before and after each trial. Ambi- ent temperature and humidity were record- ed for each trial by placing a digital hygro- thermometer on the sheet immediately be- fore the start of each trial. Configuration 1, Experiment 2. The Sheets were similarly marked, taped, aligned and secured, as in Experiment 1. Either dry ice (300—350g) or no attractant (control) was placed in the upwind (bait) corner of each cloth. Each of two cloths was placed 100 m apart and ~3 m from the hedgerow. Unlike Experiment 1, no box and cup platform was used to protect and elevate the treatments. Once the treatments were in place, groups of 15-19 marked nymphs were released 1.01 and 2.01 m downwind from the bait, on the diagonal between the upwind and downwind corners of the sheet. Each of the two groups of ticks was marked with a different randomly as- signed color. A dry ice sheet and untreated control sheet were tested in pairs, with a <10 min lag between the release of ticks 85 on the first and second sheet. One hour after the ticks were released they were recaptured and their locations recorded as in Experi- ment | (Fig. 1). Dry ice was weighed on site with a battery operated scale (Model CS5000, Ohaus Corporation, Pine Brook, NJ) immediately before and after each test. Temperature and RH were recorded on each sheet just before the start of each test. Two trials were conducted on each of 2 d. Configuration 2. Trials were conducted on sheets of the same dimensions and fabric as in Configuration |. However, the dry ice bait was located at the midpoint of one edge of the sheet, and the arcs for the 0.3, | and 2 m radii from the bait were drawn using the midpoint of the edge as origin (Fig. 2). A sheet was aligned so that the wind direc- tion was at a right angle to the edge of the sheet causing the CO, plume to pass per- pendicularly onto the sheet (Fig. 2). An in- verted styrofoam cup was placed at the midpoint of the upwind edge of the sheet, as close as possible to the masking tape edging. A block of dry ice (308-328 g) was placed on top of the cup. Fifty marked A. americanum nymphs were released 1.5 m directly downwind of the cup. One hour lat- er the ticks were collected on 2.5-cm wide masking tape, according to the zone in which they were found and returned to the laboratory for counting. Temperature, RH, wind speed (Kestrel 2000 Pocket Thermo Wind Meter, Nielsen Kellerman, Chester, PA) and background CO, readings were taken just before the dry ice was placed on the cup and just after the ticks were recap- tured. Five trials were conducted with Con- figuration 2, two of which included a paired untreated control. Each trial or pair of trials was conducted on a separate day. There was ~30 min lag between the start of each trial when paired trials were conducted. Statistical analysis.—The criterion we used to evaluate the effectiveness in the bioassays was recapture of significantly more (P < 0.05) ticks in treatments than controls. For instance, when natural popu- lations were tested, we were interested in 86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON @ release point 2m 1m dry ice wind direction Fig. 2. Diagram showing sheet with bait location, recapture zones, tick release point for Configuration 2, and alignment with wind at start of bioassay. The sheet was edged with masking tape to confine released ticks and exclude ticks from natural population. Note that Configuration 2 allows for greater changes in wind direction without attracted ticks becoming stuck on masking tape than Configuration 1 with bait in a corner. whether significantly more ticks were on dry ice sheets than untreated control sheets. When marked ticks were released on sheets, we focused on whether more ticks were re- captured in zones | and 2 on dry ice sheets than the corresponding zones on control sheets. The G statistic was used to deter- mine significance between numbers of ticks in corresponding recapture zones on paired sheets (one baited with dry ice and the other an untreated control) for the release-recap- ture trials, and between dry ice sheets and control sheets when natural tick populations were tested (Bishop et al. 1975). RESULTS Numbers of A. americanum nymphs from natural populations counted on the cloths with dry ice baits placed in forest habitats were highly variable. Even though the locations where the sheets were placed were determined by finding nymphs there VOLUME 108, NUMBER 1 87 Table 1. Efficacies of combinations of the number of A. americanum nymphs released, their release point and recapture area are manifested in the proportion of trials in which statistical significance (P < 0.05) occurred comparing dry ice-baited and control sheets. Configuration/ Distance (m) ticks No. ticks released Comparison experiment® released from bait” in each trial Recapture zone(s)° No. significant trials/total* | 1/1 1.01 10 ] 1/4 2 2.01 10 | 2/4 3 1.01 10 Le tye 1/4 4 2.01 10 Il se 2 4/4 5) 1/2 1.01 15-19 | 2/4 6 2.01 15-19 | 2/4 7 1.01 15-19 Ih Sea 4/4 8 2.01 15-19 estar 4/4 S 2 1.50 50 | 1/2 10 1.50 50 | 4p 2 2/2 “ Configurations depicted in Fig. | and 2; Experiments | and 2 each consisted of 4 trials. > Distances directly downwind from dry ice or control where nymphs were released. © Zone | (<0.3 m from bait), zone 2 (0.3—1.0 m from bait) 4 Number of trials in which significantly more nymphs were recaptured on dry ice-baited sheets than on control sheets over total number of trials. during pre-trial flagging, and the trials last- ed 2 h, four dry-ice sheets had =2 ticks on them. As many as 69 and 71 nymphs were found on other dry ice sheets, with an over- all average of 17.5 + 8.7 nymphs. On sheets without dry ice, O—3 ticks were found with an average of 0.9 + 0.3 nymphs. For 6 of the 11 dry ice-control pairs, there were significantly more (P < 0.05) ticks on the dry ice sheet than the control sheet. In Configuration 1 Experiment 1, when 10 A. americanum nymphs were released 1.01 and 2.01 m downwind of the dry ice bait or control, significantly greater num- bers of ticks from the 1.01 m release were in Zone | (<0.3 m from bait) of the dry ice sheet than the control at | h after release in just one of four trials (Table 1). Of nymphs released at 2.01 m, significantly more (P < 0.05) were in the Zone | of the dry ice sheet than the control in two trials. Of the ticks released 1.01 m from the bait, signif- icantly more (P < 0.05) were recaptured <1 m from the bait (Zones | and 2 com- bined) on the dry ice sheets than the control sheets in one of the trials. However, in all trials, of the 10 ticks released at 2.01 m, there were significantly more (P < 0.05) in Zones | and 2 combined of the dry ice sheets that the controls. In the trials using 15—19 nymphs released at 1.01 and 2.01 m in Configuration | Experiment 2, there was a similar pattern (Table 1). Significantly greater (P < 0.05) attraction to dry ice than the control was observed in half the trials in which ticks were released at 1.01 and 2.01 m from the bait and recaptured in Zone 1. In all the trials, significantly greater num- bers of ticks (P < 0.05) released at 1.01 and 2.01 m were recaptured in Zones | and 2 combined of the dry ice sheet compared to the control sheet. Temperatures ranged from 23.8—37.2°C and 23.3—30.0°C in Ex- periments | and 2 respectively; the RH var- ied from 32-57% and 34-61% in Experi- ments | and 2. For Configuration 2, in which the dry ice was placed at the midpoint of one edge of the sheet and 50 nymphs were released 1.5 m downwind of it, an average of 12.6 + 3.8 ticks (25.2%) was recaptured in Zone | in the five trials (two paired with an un- treated control and three unpaired). An ad- ditional 28.4% of the nymphs, having moved 20.5 m toward the dry ice, were in Zone 2, whereas 1.6% were in Zone 4 (>2 m from the bait). Of the remaining ticks (44.8%) recaptured in Zone 3 (1—2 m from 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the bait) in which they were released, some moved as far as the edge of the sheet (Fig. 2). Among the Configuration 2 dry ice tri- als, there was considerable variation in the numbers of ticks recaptured in Zone 1, which ranged from | to 20. In the two Con- figuration 2 trials using a simultaneous con- trol sheet significantly more (P < 0.05) ticks were recaptured in Zones | and 2 combined on the dry ice sheets than the controls in both trials. There was no signif- icant difference for Zone | in either trial. Both trials with simultaneous treatment and controls had the fewest ticks recaptured in Zone |. During the Configuration 2 trials, temperature and RH varied less than in the Configuration 1 trials, 24.0—30.9°C and 48.5—62.7% RH respectively. Winds were light (=10.5 km/ h) and intermittent, with periods of calm. The direction of the winds often shifted and sometimes drastically (e.g., east to west). The individual dry ice blocks lost 170—273 g, 1.e,. 51.8-89.5% of their original weights. Levels of airborne CO, at the end of each trial at the release point (1.5 m from the dry ice) were indis- tinguishable from the background in all tri- als. At 1.0 m from the dry ice (in line with release point) the level of CO, was high (1350 ppm above background) in only one trial. At 0.3 m from the dry ice, levels were =450 ppm above background in all trials but one during which the wind changed di- rection 180° sending the CO, plume directly off the sheet. DISCUSSION The trials with paired dry ice and control sheets to attract ticks from natural popula- tions showed that dry ice attracted signifi- cantly more A. americanum nymphs in 54.5% of the pairs. There was considerable variability in the numbers of nymphs from the natural population attracted to the dry ice baits. The principal disadvantage of test- ing the efficacy of an experimental attrac- tant with natural populations is that the numbers of ticks available to be attracted is unknown. If ticks were distributed uniform- ly in appropriate habitats, there could be some confidence that a field test of an ex- perimental attractant against a natural pop- ulation of ticks would yield meaningful data. However, these data (0O—73 nymphs on dry ice sheets) showed a clumped distri- bution of A. americanum nymphs. No nymphs were captured at some locations even though ticks had been found by pre- trial flagging in the immediate area where the sheets were placed. Dragging/ flagging for host-seeking ticks is thought to capture only a small portion (8—25%) of the ticks present in the area covered (Sonenshine et al. 1966, Carroll et al. 1991). A given num- ber of ticks attracted could be a small por- tion of a dense population of available ticks or large portion of few available ticks. By releasing known numbers of host-seeking ticks on sheets or marked ticks on the leaf litter or substrate, one can ascertain the ef- fectiveness and reliability of an experimen- tal attractant with some confidence (Falco and Fish 1991). In developing a behavioral bioassay to evaluate attractants, the dimensions of the arena and spatial relationships of the attrac- tant bait and test organism are of particular concern. These trials suggest that using re- captured ticks in Zone | (<0.3 m from the bait) as the standard for attraction is too ex- clusive. In the paired dry ice-control trials with Configurations | and 2, numbers of ticks in Zone 1 discriminated significance between dry ice and untreated controls in no more than half the trials. In contrast, Zones | and 2 (<1.0 m from the bait) of the dry ice sheets had significantly more re- captured ticks that had been released at 1.5 and 2.01 m from the bait than that of the control sheets in all the trials. To reach zone 2 these ticks would have had to have ad- vanced =0.5 m toward the bait within an hour, a good indication of attraction. Drastic shifts in wind direction or cessation of wind during a trial may make the <0.3 m zone an unrealistic standard for assessing attrac- tion. It is desirable to minimize the number of VOLUME 108, NUMBER 1 ticks released in the bioassays, because of time required to mark and recapture them. As seen in the results with Configuration 1, 10 ticks were too few to discriminate be- tween dry ice and no treatment with much consistency. The instances of significance were observed in three of eight compari- sons (four trials, two release points) for Zone | and five of eight comparisons for Zones | and 2 combined, when 10 ticks were released. However, when 15-19 ticks were released, significance was observed in four of eight comparisons for Zone | and all eight comparisons for Zones 1 and 2 combined. Other factors than numbers can- not be excluded as possible influences on the instances of significance. But even us- ing 50 ticks was no guarantee of signifi- cance in all cases. In the two paired dry ice/ control trials with Configuration 2, signifi- cance for zone | was achieved once. These results. show that variability of wind speed and direction greatly influences testing of air-borne attractants in the field. A constant unidirectional wind flow would be ideal for testing with ticks released downwind. For testing attractants with nat- ural populations, changes in wind direction are preferable because more ticks distrib- uted around a source are likely to be ex- posed to the shifting plume of attractant. None of the methods we used, including that with natural tick populations, was use- ful when winds ceased. With no wind there is little dispersion of the attractant, although gravity may allow CO, to flow down hill. At the conclusion of one Configuration 2 trial (dry ice on the side of the sheet), wind was undetectable with the wind meter. In that trial, only | tick (of 50 released) was found <0.3 m from the bait (zone 1), com- pared to an average of 12 ticks in the <0.3 m area when there were intermittent breezes of varying direction. In tests with the CO, source located in the corner of the sheet, we sometimes ob- served that a cluster of ticks (e.g., 5-8) was stuck in the masking tape barrier on one edge of the sheet) 0.3—1 m from the bait. 89 These observations were coincident with shifts =45° in the direction of the wind from the direction at the start of test (par- allel to the diagonal of sheet). As the nymphs advanced up the gradient of the CO, plume, they followed the shifting plume as it passed over the edge of the sheet, and became trapped on the masking tape barrier. By locating the CO, source at the midpoint of one side, shifting winds could swing the plume almost 90° to either side of the starting direction without draw- ing advancing ticks into the masking tape. Nevertheless, the greater leeway for wind shifts obtained by placing the dry ice at the midpoint of the upwind side of the sheet did not accommodate for all conditions that occurred during the trials. A wind shift of = 180° (based on the direction of the mark- ing flag and the visible CO, plume at the end of the trial) occurred during one Con- figuration 2 (dry ice at midpoint of sheet) trial and was associated with a cluster of nymphs caught in the tape in the 0.3—1 m zone. Nonetheless, it seems preferable to lo- cate the bait source on a side of the sheet rather than in a corner. Placing the dry ice in the center of a sheet and releasing ticks around the bait (in groups at the cardinal directions or evenly distributed), as might occur ideally in na- ture, would not necessarily improve the proportion of ticks that came to the bait. However, it would ensure that some ticks are exposed to the attractant if any breeze occurs. A very variable wind would expose the most ticks, but presumably they would take a tortuous route in the shifting CO, gradient to the bait, a route which would be longer than in a unidirectional wind and take more time. If ticks were evenly dis- tributed in a circle around a bait, only about half would be exposed to the attractant if the wind direction changed as much as 180° (e.g., a shift from west to east). Ticks could also move before a shifting plume reached them. A smaller pool of ticks to attract from would increase the likelihood of equivocal results. For best results, using the methods 90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON described in which ticks are released di- rectly downwind of an attractant source, there should be a steady or intermittent breeze that shifts <90° from the direction at the onset of the trial. Similar trials could be conducted using a 1 m? (or smaller) sheet and releasing known numbers of marked ticks on the natural ground cover. Ticks probably traverse a slightly undulating sheet more quickly than they can pass over and through the tangle and clutter of stems, broken twigs and curled leaves on the forest floor. Leaf litter and vegetation would reduce the range from which attracted ticks can reach the bait in a limited time. It is unclear from our tests to what degree, if any, marking the ticks with dusts affected their ability to respond to CO,. However, sufficient numbers of ticks moved >0.5 m and 1.0 m into Zone 2 (0.3— 1.0 m from the bait) to consistently show a significantly greater response than ticks on control sheets. Nymphs and adults of the blacklegged tick, lxodes scapularis Say marked with dusts and released in a wooded habitat were recaptured days and weeks lat- er =5m from their release point (Carroll and Schmidtmann 1996). Exposing sufficient ticks to an attractant is critical to the success of the bioassay. By releasing known numbers of ticks under partially controlled field conditions, more reliable results can be obtained for attrac- tion bioassays than by using natural tick populations. Based on the results of these trials, recommendations for evaluating a tick attractant under field conditions are: 1.) use Configuration 2 (allows for greater shifts in wind direction), 2.) release =15 ticks at a chosen release point, 3.) release ticks 1.5—2.01 m from bait, 4.) use ticks re- captured within a radius of | m from the bait as the basis for discriminating between treatments or treatment and control, and 5.) conduct trials when wind varies <180° and is not interrupted by prolonged periods of calm. ACKNOWLEDGMENTS We thank Kenneth Young, Hasani Martin and Michael Theis, USDA, ARS, Animal Parasitic Diseases Laboratory, Beltsville, MD for their valuable assistance in con- ducting the field trials. We also thank Matt Kramer, USDA, ARS, Biometrical Consult- ing Service, Beltsville, MD for data analy- SiS. LITERATURE CITED Bishop, Y. M. M., S. E. Fienberg, and P. W. Holland. 1975. Discrete multivariate analysis: theory and practice. MIT Press, Cambridge, Massachusetts, 557 pp. Carroll, J. E, E. Russek-Cohen, J. D. Nichols, and J. E. Hines. 1991. Population dynamics of American dog ticks (Acari: Ixodidae) along park trails. En- vironmental Entomology 20: 922-929. Carroll, J. E-and E. T. Schmidtmann. 1996. Dispersal of blacklegged tick (Acari: Ixodidae) nymphs and adults at the woods-pasture interface. Journal of Medical Entomology 33: 554-558. Falco, R. C. and D. Fish. 1991. Horizontal movement of adult xodes dammini (Acari: Ixodidae) attract- ed to CO,—baited traps. Journal of Medical Ento- mology 28: 726-729. Garcia, R. 1965. Carbon dioxide as an attractant for certain ticks (Acarina: Argasidae and Ixodidae). Annals of the Entomological Society of America 55:605—606. Gladney, W. J. 1978. Ticks (Acarina: Argasidae and Ixodidae), p. 102—113. Jn R. Bram [ed.], Surveil- lance and collection of arthropods of veterinary importance. U.S. Dept. Agric. Handb. No. 518. Gray, J. S. 1985. A carbon dioxide trap for prolonged sampling of /xodes ricinus populations. Exp. Appl. Acarol. 1335-1344. Holscher, K. H., H. L. Gearhart, and R. W. Barker. 1980. Electrophysiological responses of three tick species to carbon dioxide in the laboratory and field. Annals of the Entomological Society of America. 73: 288-292. Koch, H. G. 1987. Estimation of absolute numbers of adult lone star ticks (Acari: Ixodidae) by dry ice sampling. Ann. Entomol. Soc. Am. 80: 624—628. Koch, H. G., and R. W. McNew. 1981. Comparative catches of field populations of lone star ticks by CO,-emitting dry-ice, dry-chemical, and animal- baited devices. Annals of the Entomological So- ciety of America 74: 498-500. Newhouse, V. FE 1983. Variations in population den- sity, movement, and rickettsial infection rates in a local population of Dermacentor variabilis (Aca- rina: Ixodidae) ticks in the Piedmont of Georgia. Environmental Entomology 12: 1737-1746. VOLUME 108, NUMBER 1 Semtner, P. J., and J. A. Hair. 1975. Evaluation of CO,- baited traps for survey of Amblyomma maculatum Koch and Dermacentor variabilis Say (Acarina: Ixodidae). Journal of Medical Entomology. 12: 137-138. Solberg, V. B., K. Neidhardt, M.R. Sardelis, C. Hil- debrant, E J. Hoffmann, and L.R. Boobar. 1992. Quantitative evaluation of sampling methods for Ixodes dammini and Amblyomma americanum (Acari: Ixodidae). Journal of Medical Entomology 29:45 1—456. Solberg, V. B., J. A. Miller, T. Hadfield, R. Burge, J. M. Schech, and J. M. Pound. 2003. Control of Ixodes scapularis (Acari: Ixodidae) with topical 9] self-application of permethrin by white-tailed deer inhabiting NASA, Beltsville, MD. Journal of Vec- tor Ecology 28: 117-134. Sonenshine, D. E., E. L. Atwood, and J. T. Lamb. 1966. The ecology of ticks transmitting Rocky Mountain spotted fever in Virginia. Annals of the Entomological Society of America. 59:1234— 1262. Wilson, J. G., D. R. Kinzer, J. R. Sauer, and J. A. Hair. 1972. Chemo-attraction in the lone star tick (Ac- arina: Ixodidae). I. Response of different devel- opmental stages to carbon dioxide administered via traps. Journal of Medical Entomology 9:245— Da2r PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 92-100 TWO SPECIES OF CLINOPOGON BEZZI (DIPTERA: ASILIDAE: STENOPOGONINAE) FROM SABAH, MALAYSIA AUBREY G. SCARBROUGH Department of Biological Sciences, Towson University, 8000 York Road, Baltimore, MD 21252, U.S.A. (e-mail: ascarbrough @towson.edu) Abstract.—The robber flies (Diptera: Asilidae) Clinopogon nicobarensis (Schiner) and C. velatus, n. sp., are reported from Sabah, Malaysia, on the island of Borneo. This is the first record of the genus from Borneo. The species are described and compared with congeners. This is the first publication of detailed illustrations of the terminalia of both sexes in Clinopogon. The importance of the terminalia for identification purposes is em- phasized. An annotated species list of the genus is included. Key Words: Clinopogon Bezzi consists of small, gray tomentose, densely pilose flies that inhabit vegetated zones of beaches (Hull 1962). The genus is poorly studied largely because of the dependency upon external anatomy and the failure to examine the terminalia. Hence, only eight species are known, scat- tered from Madagascar eastward to Fiji (Bezzi 1910, Hull 1962, Oldroyd 1975, Jo- seph and Parui 1984, Daniels 1989, Geller- Grimm Web Database 2004). Six of these were placed in other genera (Bigot 1878, 1879; Fabricius 1775; Schiner 1868; Walk- er 1849, 1861), but later transferred to Cli- nopogon. The remaining two species (Sé- guy 1955, Joseph and Parui 1984) were de- scribed more recently. Most species have limited distributions although five occur in Indonesia. Clinopogon nicobarensis (Shin- er) 1s widely distributed, ranging from is- lands near Madagascar eastward to Taiwan. Recently, Geller-Grimm (2002) synony- mized C. sauteri Bezzi with C. nicobaren- sis (Schiner) and illustrated the intact male terminalia. Of the remaining four, C. barrus (Walker 1849) is unconfirmed because the type is lost and the type locality of the spe- cies is unknown. Diptera, Asilidae, Clinopogon, new species, terminalia, Sabah, Malaysia While searching through unsorted mate- rial collected by USNM colleagues in Sa- bah, Malaysia, on the island of Borneo, I found C. nicobarensis and an undescribed species. The species are described and the terminalia illustrated. The latter species in- creases the number to nine for the genus. This paper provides the first detailed illus- tration of the terminalia of both sexes, dem- onstrates the usefulness of the terminalia to determine species, and documents Clino- pogon in Borneo. METHODS Terminology mostly follows McAlpine (1981). Terms for terminalia and measure- ments follow Cannings (2002). 1) Body length is the linear distance from the ante- rior margin of the face to the extreme pos- terior of the abdomen, including the termin- alia. Width measurements include head width, face width, vertex width, and oral margin width. 2) Head width is the greatest distance between the lateral margins of the eyes in anterior view. 3) Face width is the narrowest distance between the eyes below the antenna. Face-Oral margin Width Ratio VOLUME 108, NUMBER 1 (FOMWR) and Face-Vertex Width Ratio (FVWR) are comparisons of divergence of the oral margin and vertex with the face; Face Head Width.Ratio (FHWR) is a com- parison of the width of the face and head. 4) Vertex width is the distance between the eyes along a line touching the posterior ocelli in dorsal view. Vertex-Head Width Ratio (VHWR) is a measurement of vertex divergence and head. 5) Oral margin width is the greatest distance of the face between the tentorial grooves. Oral Margin-Head Width Ratio (QMHWR) and Oral Margin- Vertex Width Ratio (OMVWR) are com- parisons of divergence of the oral margin with the head and vertex. Key to abbrevi- ations of structures in illustrations are given in the captions of Figs. 1—9. Specimens be- long to the National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C. (USNM). Clinopogon Bezzi Clinopogon Bezzi 1910: 153. Type species Clinopogon sauteri Bezzi, by original designation. Hull 1962 (1): 110; Oldroyd 1975: 128; Daniels 1989: 342; Geller- Grimm 2004. Diagnosis.—Distinguished from all other stichipogonine genera by a combination of the following characters: Medium to large flies, 10O—22 mm long. Face with a promi- nent, dense, shieldlike mat of setae that ex- tends dorsally to just below base of anten- na. Palpus with second segment cylindrical, largely bare, apex oblique. Abdomen broad basally, somewhat flattened, and strongly tapered apically (Hull 1962). Hull (1962) gave a detailed description of the genus. Remarks.—Clinopogon 1s closely related to Stichopogon Loew. They both have 1) the frons and face diverging widely above; 2) a long, dilated flagellum; and 3) tufts of white vestiture on the thorax (Hull 1962). The genus differs from known Oriental Sti- chopogon in the much larger body; the lon- ger, more abundant pile of the body; and the absence of differentiated dorsocentral 98 bristles, bands on the abdomen, and fused cercl. This is the first time that the terminalia of both sexes of Clinopogon have been il- lustrated in detail. These illustrations sup- port the use of the terminalia for identifi- cation purposes. To date, species descrip- tions have relied on external characters (to- mentum, ground color, and _ vestiture patterns), which are mostly inadequate for accurate identification. Clinopogon species are so similar externally that identification is unreliable without dissection and detailed examination of the terminalia. Clinopogon nicobarensis (Schiner) (Figs. 1-9) Redescription.—Male: Black body, gray tomentose and white setose, setae thin, of- ten wavy or proclinate apically. Body length 13.4—21.2 mm; FHWR 1/4.4—-1/4.7; FVWR 1/2.2-1/2.3; FOMWR 1/1.7-1/1.8; VEIWR, V2 172.33 OM Wi S/S: OMHWR 1/2.6—1/2.7. Head: Face in pro- file slightly and gradually produced ven- trally with dense, flat, shieldlike mat of fa- cial vestiture extending from just below an- tennae to just before or to apex of probos- cis; oral margin with a row of thin bristles. Palpus with basal segment tomentose and setose; apical segment cylindrical, mostly bare, with gray tomentum and white setae on basal half ventrally, apex angular with sparse setae. Proboscis blackish, mostly bare, scattered setae ventrally, basal third tomentose. Antenna with scape about as long as pedicel, combined length scape and pedicel one-third as long as flagellum; fla- gellum strongly compressed laterally, wid- est medially, tapered toward each end, 2.7 longer than wide; apex narrower than base, with minute segment and apical spine, com- bined length of latter one-sixth as long as flagellum. Frons strongly divergent dorsal- ly. Ocellar setae thin, only as thick as fron- tal setae. Occiput with two to three pairs of stout, erect, white bristles; setae mostly long, those on ventral half of head dense. Thorax: Black with gray tomentum and 94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 4 Figs. 1-5. Male terminalia of Clinopogon nicobarensis. 1, Tergite 9 [ = epandrium] and cercus, dorsal view. 2—4, Lateral, dorsal, ventral views. 5, Aedeagus, lateral view. Abbrevations: AA = aedeagal apodeme; Ae = aedeagus; Ap = epandrial apodeme; Ge = gonocoxite; Ce = cercus; EA = ejaculatory apodeme; GcA = gonocoxal apodeme; Hp = hypandrium; Gs = gonostylus; Hyp = hypoproct; LHP = lateral hypandrial process; ML = median lobe of gonocoxite. VOLUME 108, NUMBER 1 white setae. Mesonotum 3.5X as long as high; narrow corner of postpronotum bare, ground color yellow to brownish yellow: postalar callus gray tomentose, yellowish brown to reddish; tomentum thin dorsally with faint wide longitudinal stripe medially, tomentum dense elsewhere; mesonotum with short, erect setae dorsally, longer and bristly laterally; dorsocentral bristles thin, undifferentiated, about as long as adjacent setae; 3 lateral bristles, only one supraalar bristle present. Scutellum flat dorsally, wid- er than long, somewhat rectangular with rounded corners; abundant long setae dor- sally, as long as prescutellar setae; seven to eight pairs of marginal bristles laterally, only slightly thicker than setae. Pleuron with mats of long white setae on katepis- ternum anteriorly and posteriorly, on dorsal and posterior margins of anepisternum, and on anterior margin of anepimeron; posterior margin of anepisternum and _ katatergite with a row of bristles. Halter dull yellow, stalk and/or knob dull brown or blackish. Wing: Wide basally, apex pointed, about as long as abdomen; veins brown, cell sur- face with abundant fine microtrichia, most- ly brownish, costal and basal cells narrowly gray. Marginal cell open. Cell r4 trumpet- shaped, wide apically, base narrow, beyond apex of cell d; base of vein R4 abruptly arched anteriorly, ending at wing apex. Cell m3 narrowly open, base with or without short stalk. Alula narrow. Legs: Dark brown to black, white to- mentose with fine, abundant, white setae and sparse white to slightly yellowish bris- tles. Coxae with a mat of long setae; femora ventrally and posteriorly and fore- and mid- dle tibiae ventrolaterally with a fringe of abundant, long setae; hind tibia with sparse long setae, not especially abundant or form- ing a distinct fringe. Fore- and hind femora with an irregular row of stout dorsal bristles curving posteriorly; fore femur with row on apical third, that on hind femur extending from near base to apical fifth; stout anterior and preapical, posterodorsal bristle present on middle femur. Hind femur slightly con- 95 cave ventrally. Tibiae with white bristles only; apex of fore- and middle tibiae slight- ly bent inward and abruptly swollen along inner margin; hind tibiae with straight sides apically, not swollen or bent. Tarsi with black claws, latter mostly white tomentose, apex bare; pulvillus whitish to cream-col- ored, with sparse brownish streaks basally; empodium flat, bladelike on basal two- thirds, abruptly narrowed apically, setalike. Abdomen: Segments as wide as thorax basally and somewhat flattened; tergite 1 without a swollen callus; ground color black, generally densely gray tomentose, somewhat thinner dorsally; vestiture white and slender, setae short dorsally, longer lat- erally, especially on basal three to four ter- gites and sternites 3—5 medially; tergite | with several long thin bristles present in apical corner; segment 8 about half as long as 7, sternite 7 strongly reduced, straplike basally. Terminalia (Figs. 1—5): Tergite 9 [epandrium] longer than tergite 8, hoodlike, with turned-down sides and abundant long, white setae concealing terminalia laterally; apical margin with V-shaped notch medi- ally; short basal apodeme present. Cercus narrow, two-thirds as long as middorsal length of tergite 9, fused medially, strongly sclerotized; apex with narrow notch; sur- face gray tomentose with numerous long setae laterally, setae sparse dorsally. Ster- nite 8 gray tomentose basally; gonocoxite and hypandrium fused, complex; median lobe of gonocoxite elongate; lateral gono- coxal process long, digitate dorsally, flat and pointed laterally; gonostylus elongate, clavate dorsally and ventrally, wide basally, narrow and pointed apically. Hypoproct moderately keeled, grooved medially, apex divergent; in lateral view its ventral edge is even with the ventral margin of the hypan- drium. Aedeagus convex and narrow api- cally, gradually wider posteriorly; stout ventral process present. Female: Body length 16.8—21.8 mm; FHWR_ 1/4.6—1/4.8; FYWR_ 1/2.2-1/2.4; FOMWR 1/1.7—1/2.0; VHWR 1/1.9-1/2.2; OMHWR_ 1/2.2-1/2.7; OMVWRI/1.1-1/ 96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Tg 9 —<—— S/ Tg 10_/ (5 49 - Figs. 6-9. GQ 05mm Female terminalia of Clinopogon nicobarensis. 6, Sternite 9 [genital fork]. 7—9, Lateral [tergite 8 removed], ventral, and dorsal views. Abbrevations: AcSp = Acanthophorite spines; HpV = hypogynial valve; Me = membrane; St8, 9, 10, = sternites 8, 9, 10; Tg9, Tg10 = tergites 9 and 10. 1.4. Body essentially identical to male ex- cept setae generally shorter. Abdomen: Segments 1—2 and 8 with long setae. Hy- pogynal valve oval, sometimes reddish, me- dian fissure present. Terminalia (Figs. 6—9): Tergite 9 uniting acanthophorties dorsally. Acanthophorites with 8—9 pairs of long, black or yellowish spines. Cercus often with spines. Sternite 9 (genital fork, furca) with shallow notch apically, entirely strong- ly sclerotized, base narrow with a raised ca- rina. Sternite 10 asymmetrically capitate. Spermathecae thin, convoluted in a loosely organized “ball? within segment 8, surface simple and without noticeable microstruc- tures; each spermatheca separated basally, VOLUME 108, NUMBER | not joining and forming a common duct, diameter at base wide, tubes narrow grad- ually to simple valve, diameter beyond valve about one-eighth that at base, apex only slightly wider. Accessory glands sim- ple, flaccidlike sacs, gradually widened api- cally, and without distinctive characters. Material examined.—MALAYSIA: 4 6, 4 2, Sabah;/Tanjung Aru _ Beach/ 29.vili.1983/G. FE Hevel & W. E. Strainer (USNM). Remarks.—Clinopogon nicobarensis 1s readily distinguished from congeners by the large body and combined characters of the terminalia (Figs. 1—9). Clinopogon velatus Scarbrough, new species (Figs. 10-18) Description.—Male: As C. nicobarensis except as follows. Body length 15.6—17.0 mm; FHWR 1/4.8-1/5.2; FVWR 1/2.2-1/ 2.3; FOMWR 1/1.8—1/1.9; VHWR 1/1.8-1/ 2.1; OMHWR 1/2.1-1/2.5; OMVWR_ 1/ 1.1-1/1.2. Terminalia (Figs. 10-14): Ter- gite 9 with wide, concave apical notch. Cer- cus about as long as middorsal length of tergite 9; wide, deep apical notch present. Gonostylus with wide, flat, asymmetrical club apically; abundant bristly setae present dorsally. Paired median lobes of gonocoxite forming stout, arched bar in dorsal view, each fused in membrane below aedeagus and with stout podiform flange anteriorly; lateral gonocoxal process short with round apex in dorsal view, constricted preapically, wider and pointed in lateral view; obvious line of fusion of gonocoxite and hypan- drium absent in lateral view. Hypoproct strongly keeled, projecting well below hy- pandrium ventrally, apex parallel, not strongly divergent. Aedeagus stout, wide apically; stout ventral process absent. Female: As C. nicobarensis except as follows. Body length 13.0—16.0 mm; FHWR 1/4.6-1/4.8; FVYWR_ 1/2.2-1/2.4; BOMW RE 1/15 =1/1.73 »-WHWR. 1/2.0; OMHWR 1/2.9; OMVWR 1/1.2-1/14. Ter- minalia (Figs. 15-18): Acanthophorites 97 with five pairs of long, slender, black spines. Cercus with abundant, long, thin se- tae; spines absent. Sternite 9 (genital fork, furca) with large oval area apically, only slightly sclerotized. Sternite 10 slender, dig- itate. Type specimens.—Holotype 3d, MA- LAYSIA: Sabah: 17 km S. Kota Kinabalu/ 9 vin. ([983/GS FE Hevel c/w. EE. Steiner (USNM); allotype 2, MALAYSIA: Sabah; /Tanjung Aru Beach/29.viii.1983/G. FE Hev- el & W. E. Strainer (USNM). Paratypes, 2 2, MALAYSIA: Sabah;/Tanjung Aru Beach/20, 21-viti.1983/G. E Hevel & W. E. Strainer (USNM); 1 6, MALAYSIA: same data as allotype (USNM). Etymology.—Latin velatus for ‘hidden’, referring to the fact that I initially did not recognize it as a species separate from C. nicobarensis. Remarks.—Clinopogon velatus differs from C. nicobarensis by its wide, concave, apical notch on tergite 9 and cercus, and the combined characters of the terminalia (Figs. 10—18), especially the strongly keeled, ven- trally projecting hypoproct, the apically clubbed gonostylus bearing abundant dorsal setae, and the apically stout aedeagus. WORLD SPECIES LIST OF CLINOPOGON C. barrus (Walker) 1849: 353. Distribution: Malaysia. Type ° (Dasypogon), type lo- cality? C. cinctellus (Bigot) 1879: 440. Distribu- tion: Indonesia, Maluku. Type 6 (Sticho- pogon), type locality Tidore, Moluccas (Maluku). C. congressus (Walker) 1861: 147. Distri- bution: Indonesia (Irian Jaya, Sulawasi, Maluku, Java). Type 6d (Dasypogon), type locality Sulawasi (Tidon). Stichopogon albicapillus Wulp 1872: 147 Type &, type locality Indonesia (Java). C. nicobarensis Schiner 1868: 161. Distri- bution: India, Madagascar, Egypt, Mas- carene Islands, Philippines, Nicobar Is- land, Malaysia (Sabah), Taiwan, Yemen (Socotra Archipeligo). Type 2 (Sticho- 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 10-14. Male terminalia of Clinopogon velatus. 10, Tergite 9 [| = epandrium] and cercus, dorsal view. 11—13, Lateral, dorsal, and ventral views. 14, Aedeagus, lateral view. VOLUME 108, NUMBER 1 99 1 8 0.5 mm Figs. 15-18. Female terminalia of Clinopogon velatus, 15, Sternite 9 [genital fork]. 16-18, Lateral [tergite 8 removed], ventral, and dorsal views. pogon), type locality Nicobar Islands cality Sri Lanka (Kandy District, Pera- (Kondul). deniya). Clinopogon sauteri Bezzi 1910: 153. C. plumbeus (Fabricius) 1775: 793. Distri- Type 6d, type locality Taiwan (Takao), bution: Australia. Type? (Asi/us), type lo- C. odontoferus Joseph and Parui 1984: 55. cality Nova Hollandia (Australia, Distribution: Sri Lanka. Type 6, type lo- Queensland). 100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON C. reginaldi (Séguy) 1955: 10. Distribution: Tromelin Island, Madagascar. Type? (Sti- chopogon), type locality Madagascar (Tromelin Island). C. scalaris (Bigot) 1879: 440. Distribution: Fiji. Type? (Stichopogon), type locality Fiji. C. velatus Scarbrough, new species. Distri- bution: Malaysia (Sabah). Type ¢, type locality Malaysia (Sabah). ACKNOWLEDGMENTS I am grateful to EK Christian Thompson, Systematic Entomology Laboratory, USDA, Smithsonian Institution, National Museum of Natural History, Washington, D. C., for work space during my visits and for facilitating a loan of the material used in this study. Robert A. Cannings, Royal British Columbia Museum, Victoria, British Columbia, Canada, reviewed and provided useful comments on the manuscript. LITERATURE CITED Bezzi, M., 1910. Revisio systematica generis diptero- rum Stichopogon (in Latein). Annales historico- naturales Musei nationalis hungarici 8: 129-159. Bigot, J.M.F 1878. Dipteres nouveaux ou peu connus. 10.partie. XV.Tribu des Asilidae. Annales de la Société entomologique de France (5)7: 73-74. . 1879. [Note: Relative to the genera Laphyctis and Laphystia (Loew)]. Annales de la Société en- tomologique de France (5)9:235—236. Cannings, R. A. 2002. The systematics of Lasiopogon (Diptera: Asilidae). Royal British Columbia Mu- seum, Victoria, British Columbia, Canada, 354 pp. Daniels, G. 1989. Family Asilidae, pp. 326-349. In Evenhuis, N. L., ed. Catalog of the Diptera of the Australasian and Oceanian Regions. Bishop Mu- seum Press and E. J. Brill. Fabricius, J. C. 1775. Systema entomologiae sistens insectorum classes, ordines, genera species adiec- tis synonymis locis descriptioibus obsevationibus. Flensburgi et Lipsiae (Flensburg & Leipzig), 832 Pp- Geller-Grimm, FE 2002. Robber flies (Diptera: Asili- dae) of the Socotra Archipeligo, Yemen. Fauna of Arabia 19: 467-489. . 2004. http://www.geller-grimm.de/catalog/ literatu.htm. Hull, EF M. 1962. Robber flies of the world. The genera of the family Asilidae. United States National Mu- seum Bulletin 224 (1—2): 1-907. Joseph, A. N. T. and P. Parui. 1984. On some Asilidae (Diptera) from India and adjoining countries pre- sent in the British Museum (Natural History). Ori- ental Insects 18: 53-71. McAlpine, J. EF 1981. Morphology and terminology— Adults, pp. 9-63. In McAlpine, J. F, B. V. Peter- son, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood. Manual of Nearctic Diptera, Vol. 1. Agriculture Canada. Monograph 27. Ot- tawa. Oldroyd, H. 1975. Family Asilidae, pp. 99-156. In M. D. Delfindo and D. E. Hardy, eds. A catalog of Diptera of the Oriental Region. Vol. 2. University of Hawaii Press. Honolulu. Schiner, J. R. 1868. Reise der Osterreichischen Fre- gatte Novara um die Erde in den Jahren 1857— 1859. Zoologischer Theil 2(1) B; Wien. Kaiser- lich-k6nigliche Hof- und Staatsdruckeri, 388 pp. Séguy, E. 1955. Dipteres de 1’ He Tromelin. Le Natur- aliste malgache 7: 9-12. Walker, E 1849. List of the specimens of dipterous insects in the collection of the British Museum. London. Part 2: 231—484 (299-484). . 1861.Catalogue of dipterous insects collected at Menado in Celebes and in Tond by A. R. Wal- lace with descriptions of new species. Journal of the Proceedings of the Linnean Society of London 5: 258-270. Wulp, E M. van der. 1872. Bijdrag tot de Kennis der Asiliden van den Oost-Indischen Archipel. Tijdschrift voor Entomologie (2)7(15): 129-279. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 101-118 LETHAEINI (HEMIPTERA: LYGAEOIDEA: RHYPAROCHROMIDAE) ASSOCIATED WITH FIGS IN MEXICO, WITH THE DESCRIPTION OF A NEW SPECIES OF CISTALIA Luis CERVANTES PEREDO AND SAGRARIO GAMEZ VIRUES Instituto de Ecologia, A.C., km 2.5 Antigua Carretera a Coatepec # 351, 91070, Xalapa, Veracruz, México (e-mail: cervantl @ecologia.edu.mx, sagrariogamez @ yahoo.com.mx) Abstract.—Descriptions and illustrations of some Lethaeini associated with Ficus spp. in Mexico are presented. Notes on the biology of Cistalia pallidifemur Cervantes and Gamez, Cryphula apicata (Distant), C. trimaculata (Distant), Neopetissius slaterorum O'Donnell, Paragonatas costaricensis Distant, P. divergens Distant, and Petissius spinipes (Stal), and their distribution, mainly within the Mexican Gulf Coast region, are included. Cistalia pallidifemur, n. sp., is described and illustrated. All species included in this study are considered facultative seed predators of figs. Key Words: Members of Lethaeini are normally small in size, with a shining to subshining dorsal surface. They are separated from other rhy- parochromid tribes by the following syna- pomorphies: linear placement of trichoboth- ria on abdominal sternum V, loss of y-chro- mosome, extreme modification of the sperm reservoir, and development of iridescent head areas (Slater and O’Donnell 1978, O’Donnell 1991). Nymphs lack a Y-suture but have lateral evaporative areas. In most genera, there is a conspicuous trichoboth- rium present near each anterolateral pro- notal angle (Slater and Baranowski 1990, Schuh) and Slater 11995), Very little 1s known about the biology of Lethaeini. Sweet (1964) reported Cryphula trimacu- lata Distant as a ground-litter inhabitant of long-lasting old fields vegetated with peren- nial bunch grasses. It definitely prefers to feed on seeds of Panicum spp. but it also feeds on seeds of fescue, millet, and straw- berry. Slater (1972) found that several spe- cies of Neolethaeus in South Africa were obligatory terrestrial and facultative terres- Cistalia, Ficus, Lethaeini, Mexico trial seed predators, mainly of F. sycomorus L. and F. capensis Thunb. Baranowski and Slater (1979) described the biology of Cry- Phula affinis (Distant) and C. bennetti Bar- anowski and Slater in Trinidad, where both species were found under Stachytarphaeta Jamaicensis (L.) Vahl. (Verbenaceae). They mentioned that although both species were found in the same area, they were usually in different microhabitat conditions, with C. affinis preferring relatively moister situa- tions. Slater and Baranowski (1990) report- ed Cistalia signoreti (Guérin), Cryphula tri- maculata (Distant), Paraganotas costari- censis (Distant), and P. divergens (Distant) from Florida. They recounted what Sweet mentioned about the biology of C. trima- culata and pointed out that P. divergens was found in temporary early succession stages of old-field habitats and apparently fed on a variety of seeds. Nymphs of 1 “, 4, and 5" instars of C. signoreti have been described (Slater and Baranowski 1973). Baranowski and Slater (1979) described all immature instars of Cryphula bennetti Bar- 102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON anowski and Slater and C. affinis (Distant). O’ Donnell (2001), while describing the ge- nus Neopetissius, found that most of the material has been collected at light, al- though she had records of N. slaterorum O’Donnell found in litter of Ficus sp. Here we describe the biology of six Le- thaeini that are associated with several spe- cies of figs, and include descriptions of some nymphal stages of Cistalia pallidife- mur, n. sp., C. trimaculata, N. slaterorum, P. divergens, and Petissius spinipes Stal. The description of a new species and the distribution of all species within Mexico also are given. MATERIAL AND METHODS Monthly collecting trips between 2001 and 2003 were made to several localities in the Mexican states of Campeche, Tamauli- pas, and Veracruz. The objective was to collect lygaeoids associated with wild, fruiting fig trees. Around 30 fig species were sampled from localities from sea level to an altitude of 1,500 m. Several types of vegetation were included: low tropical dry forest, me- dium tropical forest, high tropical rain for- est, and cloud forest. The methodology fol- lowed is described in detail in Cervantes et al. (2004). Measurements are given in mm = LS): Additional data of their distribution in Mexico is also reported, based on speci- mens deposited in Coleccion Nacional de Insectos from Instituto de Biologia, Univ- ersidad Nacional Autonoma de Mexico (CNIN) and based on Slater and Brailovsky (2000). Voucher specimens and nymphs have been deposited at Coleccion Entomo- logica del Instituto de Ecologia, A.C. Xal- apa, Veracruz, Mexico (IEXA). Other ab- breviations used are BMNH, The Natural History Museum, London, U.K., and AMNH, American Museum of Natural His- tory, New York, NY, USA. Deposition of the specimens of the new species described here are listed in the description of the spe- cies. RESULTS Cistalia pallidifemur Cervantes and Gamez, new species (Figs. 1A—C, 2A—D) Diagnosis.—Dorsal and ventral surfaces of head and antennal segments I, II, and IV dark brown. Femora with basal two-thirds pale yellow and apical third pale brown. Head covered with small semierect hairs and dorsal surface covered with small sil- very hairs. Parameres with sides of blade smooth. Adult (Fig. 1C).—Body slightly elongat- ed with parallel margins. Head, pronotum, scutellum, clavus, chorion, ventral surface, and antennal segments I, IH], and IV dark brown. Dorsal and ventral surfaces of head covered by small semierect hairs, with dor- sal hairs slightly longer; eyes reddish brown and ocelli yellowish brown; antennal seg- ment III with basal third dark brown, re- mainder pale yellow. Rostrum pale brown, reaching mesocoxae. Lateral pronotal mar- gins with sparse long hairs; dorsal surface covered by small silvery hairs; anterior pro- notal margin slightly concave and posterior pronotal margin straight; lateral pronotal margins weakly rounded; anterior lobe im- punctate, and posterior lobe with small punctures. Coxae and trochanters pale brown, femora with basal two-thirds pale yellow and apical third pale brown; tibiae and tarsi yellowish brown; fore femur with a series of hairs along inner margins; tibiae with long, widely-spaced spines. Evapora- tive area rough, with peritreme orange brown. Scutellum, clavus, and chorion cov- ered by small decumbent hairs; scutellum with small punctures, clavus and chorion with larger punctures forming rows. He- melytral membrane smoky. Ventral abdom- inal surface covered with short hairs, but with several long setae along middle. Male genitalia: Parameres sickle shaped with sides of blade smooth (Fig. 2C). Male measurements (n = 10): Body length 5.57 + 0.09; head length 0.64 + 0.1; width through eyes 1.06 + 0.05; interocular VOLUME 108, NUMBER | 103 Fig. 1. Cistalia pallidifemur. A, Egg. B, First instar. C, Adult. Scale = 1 mm. 104 distance 0.56 + 0.04; interocellar distance 0.39 + 0.2; postocular distance 0.06 = 0.2; antennal segment lengths: I 0.68 + 0.04, II 0:99): 0:3, TM.0.89.= 0:03, 1V 0.91 = 0.07; rostral segment lenghts: I 0.72 + 0.03, II OF73n210:04. «TI O668=5 10.058 0V “0:42 = 0.05; pronotum: length 1.11 + 0.07, width across humeral angles 1.97 + 0.07, width across anterior margin 1.3 + 0.09; scutel- lum: length 1.18 + 0.06, width 1.01 + 0.07; fore leg: femur length 1.45 + 0.07, tibia length 1.38 + 0.08, tarsomere lengths: I 0:53) 0M. Th 0.14: = 020.2 Ss 0:02. Female measurements (n = 10): Body length 5.95 + 0.3; head length 0.65 + 0.09; width through eyes 1.09 + 0.02; interocular distance 0.59 + 0.02; interocellar distance 0.41 + 0.02; postocular distance 0.07 + 0.04; antennal segment lengths: I 0.69 + 6:03, 11 1:01). .0.074m1 (0:88. = O04, TV 0.96 + 0.5; rostral segment lengths: I 0.74 470,04: 10:76: 0:06; TH.O:65 =-0.06;1V 0.44 + 0.04; pronotum: length 1.17 + 0.08, width across humeral angles 2.06 + 0.14, width across anterior margin 1.3 + 0.08; scutellum: length 1.21 + 0.1, width 1.16 = 0:13; fore leg: femur length, 1:46°+,0:08; tibia length 1.45 + 0.08, tarsomere lengths: Ib O52 2210.03; O07, = 0025 O27 Ie 0.01. Types.—Holotype: male, MEXICO, VE- RACRUZ, San Andres Tuxtla, Estacion Biologica Los Tuxtlas, 12.X1I.2001, light traps, A. Sanchez (IEXA). Paratypes: MEXICO: (‘CAMPECHE> =» Escatceaa, 4.V1I.1982, O. Canul, 2 56 (CNIN); km 78 Escarcega-Xpuhil, 31.V.2001, light traps, L. Cervantes; C. Mayorga, 1 ¢@ (IEXA) CHIAPAS: Agua Azul, 22.V.1979, L. Ri- vera, 1 2; Km 9 Palenque-Ocozingo, 5.V.1982, V. Melendez, 2 ¢, 2 @, A. Ibarra, 1d, 1 2; km 7 Tapachula-Talisman, Finca Chinita, 20.IV.1983, H. Brailovsky, 1 6; Tapachula-Huixtla, 26.XII.1985, H. Velas- co, | d. OAXACA: San Pedro Juchatengo, 12. VilL04.L:- Cervantes, Ay Delgado, iC: Mayorga, | 6, 1 2 (CNIN). PUEBLA: San Diego, )\13.V1L.1953.3.H.. Brailovsky, 1.d: Tenango de Doria, 5.11.1986, H. Brailovsky, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON E. Barrera; Id. 249; Yohualehanr 711.1986, E. Barrera, 1 2; Km 10 Xicote- pec-Patla, 2.VII.1994, G. Ortega, E. Bar- rera, I ¢. OQUERETARO.. Chuvaye 21-VI.1998, TE. Cervantes, GOrtega, Id; 5 @; km 8 Neblinas-Agua Zarca, 19.VHI.1998, H. Brailovsky, E. Barrera, 2 2. QUINTANA ROO: Felipe Carrillo Puerto, Buenavista, 22.V1I.1989, A. Cadena, L. Cervantes, 1 2 (CNIN). VERACRUZ: San Andres Tuxtla, Estacion Biologica Los Tuxtlas, 12.X1.2001, light traps, A. San- chez, 1 2? GEXA); 14.V.1985,; L. Cervantes, 1@s) 172V.1985.,- Ex Cervantes 12 = 18°VA1L 985. (La -Céervantest 1 mashes LO-VAG85, TD. Cervantes H 222 0— 26.VII.1985, E. Ramirez, 1 2; 28.VII.1985, Cx Mayorga, Id 19-9 S30 Mines me- Mayorga, 1 6, 1 2; 24.VIII.1985, C. May- orga, 19; V. Melendez, IG 329: VIILI9Sa- C. Mayorga, 1 @; 30.VIII.1985, C. May- orga, 1 63 \1—10.1X.1985, Ae Ihara, Feo; METX. 1985, E. Ramirez, 1°25 6X 19850: Mayorga, V. Melendez, 1 6, 2 2 (CNIN); [6.1X.1987; FE Arias, 3°" 2 =(BIMINED): 17.1X.1987, E Arias, 4 6, 3 2 (AMNH); if SS8. 1G Mayorga’ iia 20.XI.1988, E. Mejorada, 1 ¢; 6.1X.1989, RL Colin, Wee; 12 1989) Ire Colne Rojas, 1 o and 1 9; 18.1X.1989, J-L. Colin, H. Rojas; 1 6220 1X 1989! Colm Rojas, 1 2; 6.201989" J-L, Colm, Ho Rojas ff 9 3exX.1989. AL. Colin. HH. Rojasaiige. l 2 (CNIN); 11.X2-1989,3-E--Colin, H. Ro- jas, 2 2 (NMNH); 1-3.IV.1993, H. Brai- lovsky; 1 -¢,-4--2:;, Sam-Andres, Duxtia: 8.X1.1977, H. Brailovsky, 1 2; San Andres Tuxtla, Km 3 Estacion los Tuxtlas-Cate- maco, 2.XII.2001, under Ficus insipida Willd., A. Sanchez, L. Cervantes, 3 2; San Andres Tuxtla, Ejido Lazaro Cardenas, 12. VIL-1989, IL. Colin, He Rojas, 4ngen3 2; Nanchital, 5.X.1976, light traps, H. Brailowsky, 1 36; Rio Maquinas, 17.VI1I.1991, E. Mejorada, B. Arellano, 1 9 (CNIN); Jalcomulco, 3.XII.1999, L. Cer- vantes, | ¢, 1 2; Misantla, km 18 Misan- tla-Juchique, 26.1.2002; under Ficus pertu- VOLUME 108, NUMBER 1 105 C A, Pygophore of Cistalia pallidifemur. B, Pygophore of C. explanata. C, Paramere of C. pallidifemur. D, Paramere of C. explanata. Scale = 1 mm. Fig. 2. S@tr ley: Ui: (IEXA). Discussion.—This species differs from C. explanata in the form of the parameres. In C. explanata the blade has a small spine on the inner side (Fig. 2D), whereas the py- gophores of both are very similar (Figs. 2A-B). Cistalia pallidifemur differs from C. explanata by the general coloration of the femora; C. explatanata has all femora from pale brown to brown, while C. palli- difemur has the basal two-thirds pale yel- low and the apical third brown. In Slater and Baranowsk1’s key (1973), C. pallidifemur will run to C. alboannulata and C. neotropicalis, because it has slightly more than half of the third antennal seg- Pacheco, L. Cervantes, 1 @ 5 D ment pale, although it is never more than three-quarters pale. Etymology.—Named for the pale basal two-thirds of the femora. Distribution.—MEXICO: Campeche, Chiapas, Puebla, Queretaro, Quintana Roo, and Veracruz. Egg (Fig. 1A) (n = 7).—Oval, pale yel- low, chorium surface smooth, with irregular distributed small punctures. 1.22 + 0.03 mm long by 0.57 + 0.04 mm wide. First instar (Fig. 1B).—Head, pro-, and mesonota pale brown, pro- and mesonota with small pale yellow punctures irregularly distributed; eyes red; antennal segments brownish gray, with intersegmental unions pale yellow with reddish areas; margins of 106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON segments II and I dark brown. Rostrum pale brown. Metanotum pinkish, with lat- eral margin pale brown; thoracic pleurae brown; legs pale yellow; abdominal seg- ments I to III redish, rest of abdomen pink- ish. Rostrum extending beyond metacoxae. Scent glands present as brown bands on ab- dominal segments IIJ-IV, ['V—V, and V—VI. Dorsal and ventral surface of abdomen cov- ered by small decumbent hairs. Three small brownish gray plates present on ventral middle line of segments VII-L[X. Measure- ments (n = 4). Body length 1.75 + 0.21; head length 0.39 + 0.08; width through eyes 0.49 + 0.02; interocular distance 0.34 + 0.02; postocular distance 0.07 + 0.06; antennal segment lenghts: I 0.23 + 0.02, I O25 = O) TH O22 = "0:03, IV 0.44 = 0:02; rostral segment lengths: I 0.34 + 0.02, I Ot 2210.02, ThE-O!22= O03. IV 0:25) = 0.04; pronotum: length 0.25 + O, width across humeral angles 0.52 + 0.03, width across anterior margin 0.423 + 0.03; fore leg: femur length 0.46 + 0.06, tibia length 0.46 + 0.05, tarsomere lengths: I 0.14 + OL027 110. 2720.01: Biology.—Cistalia pallidifemur was col- lected in leaf litter of Ficus insipida Willd. and F. pertusa L. (Moraceae), throughout the year. Individuals usually run under the dead leaves, feeding on seeds of fallen fruits. This species also was collected at light traps. Cryphula apicata (Distant) (Fig. 3) Adult (Fig. 3).—3.5 mm long. Pronotum and scutellum shiny black; pronotum with lateral margins emarginated; posterior mar- gin of pronotum with pale areas. Scutellum with only apex ochraceous; antennal seg- ments brown, with segments I and II slight- ly paler; antennal segment II longest; third and fourth subequal, segment III with apex grayish. Biology.—This species was collected only under Ficus calyculata Mill. Individ- uals were found running on the ground un- der the leaf litter. Distribution.—MEXICO: Puebla, Tamaulipas. Guerrero, Cryphula trimaculata (Distant) (Figs. 4A—D) Adult (Fig. 4D).—Small, 3.0—4.0 mm long. Body shiny, dark castaneous brown. First and second antennal segments ochra- ceous, third and fourth brown. Anterior por- tion of pronotum impunctate and_ black, posterior part closely punctate and ochra- ceous. Pronotal humeral angles with three spots, on scutellum and some veins of hem- elytra pale yellow. Corium ochraceous, coarsely punctate; membrane dark brown, with apex hyaline. Legs ochraceous. Second instar (Fig. 4A).—Body pyriform with maximum width through abdominal segment V. Head brown, with darker areas around each eye. Pro- and mesonota dark brown, two pale yellow maculae on meson- otum, one on each side of middle line; me- tanotum pale brown. Eyes red. Antennal segments I and II pale yellow, segment III brown, and segment IV brown and darkest. Labium and legs pale yellow; femora and apex of labium brownish. Thoracic pleurae brown. Abdomen yellowish with red trans- verse lines. Scent gland plates between seg- ments III-IV, IV—V, and V—VI, but only first and second scent gland openings visi- ble. Tylus half as long as antennal segment I. Rostrum reaching middle of metacoxae. Apex of tylus with a few short hairs. Pron- otum with one tricobothrium at each an- terolateral margin. Trichobrothria present on abdominal segments IT to VIII. Ventral plates present on midline of segments VI— IX. Abdominal venter covered by small brown setae. Hind tibia with spinelike setae only at apex, and middle tibia with more evenly distributed spinelike setae. Measure- ments (n = 1). Body length 2.15; head length 0.375; width across eyes 0.475; in- terocular distance 0.35; postocular distance 0.125; antennal segment lengths: I 0.2, I 02, HI 0.275, IV 0.425; rostral segment lengths: 10:25; d4.0:23, 00h 0275 aLVs O72; pronotum length 0.3; width across humeral b 2. Boe e as 5 bg ie Be t;) re re ws o _¢ Mes a as 22 Par me < RES teae aioe ean op 8S tdpicgastee oar mes coe re a" o & os =< & orc jes?) LH A 3 f= =| Fa i ae ow apo =a Soe s E ge E BS i Bo Sar 2 fe ae eee 5 est : ; repre ee 5 Sp of seth, eee 2 Loe Be 3 n 5 5 Tg hie gece oe PRIS eh ERE ss) De = Siew ss < < ray v lad S 3 = v ea a <0 E 2) 8 A) aS) bn Zz 3 3 0 a = soem exes 8 a va) va) S 5 non S : eee foe) Nn os} =) ob O wn — 4 ep (2 iol} © om —@'2 & = ao I-III pale yellow with bases and apices O75: 108 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Cryphula trimaculata. A, Second instar. B, Fourth instar. C, Fifth instar. D, Adult. Scale = 1 mm. VOLUME 108, NUMBER 1 white; segment IV brown with white base. Labial segments I-III, tibiae, and tarsi pale yellow; labial segment IV and femora pale brown. Pronotum and thoracic pleurae brown; meso- and metanota mixed with ir- regular brown and pale yellow areas: scu- tellar area with dark brown punctures. Ab- domen reddish with pale yellow bands along segment unions. Plates of scent gland openings very similar to second instar. Lat- eral margins of abdomen slightly darker. Two pairs of tricobothria situated in front of eyes. Labium reaching base of mesocox- ae. Fore tibia with numerous spinelike se- tae. Fore femur sulcate, and with four or five small spines on internal margin. Me- sothoracic wing pads covering 4/5 of me- tanotum. Measurements (n = 3). Body lensth= 2-37" -s4045:) head: length 0:53. 0.04; width across eyes 0.58 + 0.02; inter- ocular distance 0.37 + 0.01; postocular dis- tance 0.21 + .0.04; antennal segment leneths2O25) =. 005,500.35 90:05. Il 033922 0103 -1V 0.46 "0.01; rostral seg- menvucneths: 110134 = OOS. 1l 0.35 = 0; Il 0.28 + 0.02, IV 0.28 + 0.03; pronotum length 0.43 + 0.04; width across humeral angles 0.75 + 0.09; width across anterior margin 0.6 + 0.05; fore leg: femur length O'S 75= 90:03; stibianliensth 0.55) 0.5; tar somecc: lengths 109" - 0:01, M019 0.01. Fifth instar (Fig. 4C).—Very similar to fourth instar, except pronotum and thoracic pleurae slightly paler. Mesotharacic wing pads almost all pale yellow with margins pale brown. Punctures of meso- and metan- ota more numerous. Coxae with two or three spinelike setae. Spines of fore femur thicker. Measurements (n = 10). Body lenethy 32570-4031; headnength 0:55 0.04; width across eyes 0.742 + 0.03; in- terocular distance 0.45 + 0.01; postocular distance 0.14 + 0.07; antennal segment leneths 0:29) 220:032all (0:50 20:08 0.45 + 0.04, IV 0.56 + 0.04; rostral seg- ment lengths: 10.45 + 0.04, Il 0.47 + 0.03. Ill 0.42 + 0.04, IV 0.3 + 0.04; pronotum length 0.61 + 0.0.5; width across humeral 109 angles 1.11 + 0.07; width across anterior margin 0.8 + 0.02; scutellum length 0.56 + 0.05; scutellum width 0.64 + 0.06; fore leg: femur length 0.73 + 0.06; tibia length 0.74 = 0.05; tarsomere lengths: I 0.22 + O02. Wee! 25 ==70:027 Biology.—In this species, macropterous forms are present; however, brachypterous forms are more abundant. Cryphula trima- culata was collected in leaf litter of Ficus cotinifolia Kunth., F. perforata L., F. per- tusa, F. tecolutensis (Liebm.) Miq., and F. trigonata L., during February, March, April, and August. Nymphs were collected only on F. pertusa and F. tecolutensis. Both adults and nymphs live under the first layers of litter, very close to the ground, hiding in crevices. They seem to prefer dry ground conditions with little leaf litter. This species also was found close to perennial grasses that grow in clumps. Distribution — MEXICO: Baja Califor- nia, Jalisco, Morelos, Tamaulipas, and Ve- racruz. Neopetissius slaterorum O° Donnell (Figs. 5A—B) Adult.—Head dark brown; anterior pro- notal lobe, almost all scutellum, and macula on corium dark brown; irregular macula on posterior pronotal lobe and apex of scutel- lum reddish brown; two elevated areas of scutellum, and some areas of posterior pro- notal lobe pale yellow. One macula on each side of middle line just above pronotal col- lar, lateral pronotal margins, humeral an- gles, and claval suture cream. Distal third of antennal segment II white, remainder pale brown. Femora amber, with distal end paler; tibiae and tarsi pale yellow. Adult il- lustrated by O’ Donnell (2001). Fourth instar (Fig. 5A).—Pyriform. Head brownish orange; eyes red; antennal seg- ment I brownish yellow; base of segment II brownish yellow, rest dark brown; segment III dark brown; base of segment IV dark brown, rest brownish yellow. Rostrum brownish yellow, apex of segment IV dark brown. Pro-, meso-, and metanota mixed 110 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Neopetissius slaterorum. A, Fourth instar. B, with dark brown and brownish orange are- as, with irregularly distributed dark brown punctures. Thoracic pleurae brownish or- ange; femora brownish yellow; tibiae and tarsi pale yellow. Abdomen grayish, with red bands near lateral margins; interseg- mental unions pale yellow. Abdominal scent glands dark brown, situated between segments HI-IV, IV—V, and V—VI. Four small, dark brown plates ventrally on mid- dle line of segments VI-IX. Antennal seg- ments covered by small hairs, been more abundant on segment III. Rostrum reaching base of abdominal segment III. Mesotho- racic wing pads covering almost all metan- otum. Fore femur with a row of five or six small spines on internal margin. Middle and hind tibiae with long spines irregularly dis- tributed. Abdominal scent glands only func- tional on segments III-IV and IV—V; gland one on segments V—VI without openings. Measurements (n 1). Body length 3.5; head length 0.3; width through eyes 0.8; in- Fifth instar. Scale 1 mm. terocular distance 0.5; postocular distance 0; antennal segment lengths: I 0.45, II 0.6, III 0.5, IV 75; rostral segment lengths: I 0.6, I1 0.7, III 0.6, IV 0.4; pronotum: length 0.5, width across humeral angles 1.1, width across anterior margin 0.9; fore leg: femur length 0.9, tibia length 0.9, tarsomere leneths: Os), no Fifth instar (Fig. 5B).—Pyriform. Very similar to fourth instar. Head brownish or- ange, although a few individuals with dark- er areas. Antennal segments I and IT brown, segment II with a small basal yellow area; segments III and IV dark brown. Dark brown and brownish-orange areas of pro-, meso-, and metanotum defined as longitu- dinal bands; almost all dark brown punc- tures situated over dark brown bands. Me- sothoracic wing pads reaching base of ab- dominal segment III. Measurements (n 4). Body length 6.48 + 0.43; head length 0.72 + 0.03; width through eyes 1.15 + 0.1; interocular distance 0.74 + 0.05; inter- VOLUME 108, NUMBER 1 ocellar distance 0.61 + 0.02; postocular distance 0.1 + 0; antennal segment lengths: FO:693-=50:08; Tl 1.02. = 0.05; 1-Qs9 + 0.06, IV 1.08 = 0.05; rostral segment lenpths: 10:98 = 0.05; Il 1.02 + 0:05, Il VO9s= 0:121Y 0.52 = 0.06; pronotum: length 1.01 + 0.06, width across humeral aneles 2.02°= margin 1.25 + 0.04; fore leg: femur length 1.34 + 0.12, tibia length 1.41 + 0.08, tar- somere lengths: 1 0.38 +.0.06, 1.041 + 0.02. Biology.—Neopetissius slaterorum was collected in the litter of Ficus colubrinae Standl., F. cotinifolia, F. insipida, F. ovalis (Liebm.) Miq., F. perforata, F. trigonata, and F. yoponensis Desy. throughout the year. As with many other Lethaeini reported in this work, adults of this species usually run under leaf litter just above the soil sur- face, hiding under twigs or in crevices. This species was also attracted to light traps. Distribution.—MEXICO: Campeche, Chiapas, Guerrero, Jalisco, Oaxaca, Puebla, Quintana Roo, San Luis Potosi, Tamaulipas, Tlaxcala, Veracruz, and Yucatan. Paragonatas costaricensis (Distant) (Fig. 6) Adult.—Body oval and shiny, general coloration red brown, with numerous dorsal and ventral hairs. Antenna, rostrum, tibiae, and tarsi yellowish brown; femora reddish brown. Hemelytra with pale lines along cla- val suture and on some corial veins. Lateral corial margins pale brown and expanded. Fore femur unarmed; rostrum reaching me- socoxae. Evaporative area covering only basal region of metapleuron. Biology.—Paragonatas costariscensis was collected under the litter of F. trigon- ata. It was found between April and May 2001. It is a relatively rare species, found in small numbers at most study sites, but it was usually very active. This species also was collected at light traps. Distribution.—MEXICO: Campeche, Chiapas, Jalisco, Oaxaca, Puebla, San Luis Potosi, Sinaloa, Veracruz, and Yucatan. 0.12, width across anterior 11 Paragonatas divergens (Distant) (Figs. 7A, B) Adult (Fig. 7B).—Body 4.5—5.0 mm in length. Body surface dull, black; head shiny black; antenna pale brown with segment IV slightly darker, segment II larger than seg- ment HI and IV; pronotum black with lat- eral margins ochraceous, scutellum black; clavus ochraceous with numerous dark punctures and with a parallel line to margin of scutellum; corium black, with veins, base, and union with clavus ochraceous, costal margin and irregular subapical mac- ula gray; hemelytral membrane smoky, base and veins paler. Coxae shiny black ventral- ly, femora dark brown; rostrum, tibiae, and tarsi pale ochraceous, tibiae with numerous hairs; fore femur with two or three short acute spines below a series of elongate se- tae; rostrum reaching mesocoxae. Scent gland auricle slender, acuminate, not strongly hooked posteriorly; evaporative area covering all but outer one-third of me- tapleuron. Fourth instar (Fig. 7A).—Body elongat- ed, maximum width through abdominal segment III, head, pro-, meso-, and metan- otum, and thoracic pleurae brown. Eyes red. Antennal segments brown with white bases. Labial segments I to HI pale brown, seg- ment IV brown. Femora pale brown, tibia and tarsi pale yellow. Abdomen pinkish, with pale yellow unions between segments. Scent gland plates present on segments III— IV, IV—V and V—VI; first two elongate and third very small and without openings. Ven- tral plates present on abdominal segments VI-VIII. Dorsal surface of head, thorax and abdomen, and ventral surface of abdomen covered by small brown hairs. One long tri- cobothrium on each frontal angle of pron- otum. Lateral margins of pro- and mesonota with thick setae. Tibiae of all legs with spinelike setae. Fore femur with a line of spinelike setae on internal margin. Labium reaching middle metacoxae. Measurements (n = 1). Body length 3.8; head length 0.7; width across eyes 0.75; interocular distance PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Paragonatas costaricensis. Adult. Scale = 1 mm. VOLUME 108, NUMBER 1 (Pe eb Ply Ells isa unt er Ree EOC les We 0.5; postocular distance 0.175; antennal segment lengths: I 0.425, II 0.6, III 0.55, IV 0.7; rostral segment lengths: I 0.55, I 0.58, III 0.46, IV 0.4; pronotum length 0.6; width across humeral angles 0.1; width across anterior margin 0.8; fore leg: femur length 0.9; tibia length 0.9; tarsomere lengths: 1 0.25; 11.0.3: Biology.—Paragonatas divergens was collected in the litter of F. continifolia, F. ovalis, F. pertusa, and F. tecolutensis throughout the year. This species also was collected in large quantities at light traps. Distribution.—MEXICO: Campeche, Chiapas, Guerrero, Jalisco, Morelos, Oa- xaca, San Luis Potosi, Tabasco, Tamaulipas, and Veracruz. Petissius spinipes Stal (Figs. 8A—G) Adult (Fig. 8G).—Body oval, slightly elongate; dorsal surface dark brown, almost B Paragonatas divergens. A, Fourth instar. B, Adult. Scale = | mm. black, slightly shiny. Head with a basal me- sial iridescent spot. Antenna, labium, and legs pale brown. Pronotum with lateral mar- gins pale brown; a pair of yellowish-brown maculae on anterior border, some individ- uals with humeral angles yellowish brown. Pronotal calli dark brown, almost black, without punctures, rest of pronotum, and scutellum dark brown and covered with punctures. Hemelytra dark brown with nu- merous punctures; clavus with three lines of webbed punctures and one irregular line. Venter of entire body black. Head slightly declivent. Antennal segment I smallest, oth- er three of similar size. Labium reaching metacoxae. Lateral margins of pronotum slightly expanded, carinate, with a pair of tricobothria on anterior third; transverse im- pression of pronotum well defined. Prono- tum with anterior and posterior margins straight, lateral margins convex. Scutellum 114 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with a mesial elevated line. Metathoracic scent gland with evaporative area rough and expanded anteriorly, longer than posterior mesopleuron. Fore femur with four distal spines, arranged in a line, and two long and proximal seta, on same line. Fore tibia with posterior spines; only hind leg, and tibiae of other legs with spines on all their sur- faces. Tarsi with numerous short setae. Egg (Fig. 8A).—Ovoid, both poles rounded. 1.12. + 0.06 mm long by 0.64 + 0.04 mm wide. White when laid, turning yellowish brown in two days, and six days later finally turning reddish. Corium surface almost completely smooth, except for very small mycropilar processes on anterior pole. First instar (Fig. 8B).—Body slightly elongate, abdomen as wide as thorax. Head, pro-, and mesonota pale brown, although slightly darker towards posterior margin of each segment. Metanotum pale yellow. Eyes red. Antennal segments I and IV pale yellow and segments II and II pale brown, joint between each segment reddish. Labi- um and legs pale yellow; femora and apex of labium sometimes grayish. Thoracic pleurae brown. Dorsal and ventral surfaces of abdomen reddish with a few irregular pale yellow maculae; unions between each segment also pale yellow. Scent gland openings between segments I-IV, IV—V and V—VI narrow and brown, becoming smaller caudally. Tylus as long as half of antennal segment I. Labium slightly longer than metacoxa. Dorsal surface of head with a few short setae and a trichobothrium sit- uated just in front of each eye; pronotum with trichobothrium on each lateral margin near front angle. Measurements (n = 10). Body length 1.6 + 0.14; head length 0.37 + 0.04; width through eyes 0.42 + 0.03; interocular distance 0.27 + 0.03; postocular distance 0.08 + 0.03; antennal segment fensths: 1 O07 2)/01029 1bO 222301037 alll 0.18 + 0.02, IV 0.44 + 0.04; rostral seg- ment lengths: 10.26 + 0.01, If 0.24 + 0.03, III 0.19 + 0.03, IV 0.21 + 0.02; pronotum: length 0.19 + 0.04, width across humeral angles 0.33 + 0.02, width across anterior margin 0.3 + 0.01; hind leg: femur length 0.4 + 0.07, tibia length 0.44 + 0.06, tar- somere lengths: I O02 = OFN.O! LS een0102- Second instar (Fig. 8C).—Very similar to first instar, although abdomen wider than thorax. Head, pro- and mesonota turn dark brown. Antennal segments I to III pale brown, segment IV pale yellow. Labium and legs pale brown. Abdomen with orange areas around scent gland openings; a few pale brown maculae appear on lateral mar- gins of each segment. Labium reaching sternite III. Fore femur wider than other femora, with a row of small spines on its external margin. Measurements (n = 10). Body length 1.85 + 0.15; head length 0.46 + 0.05; width through eyes 0.51 + 0.01; interocular distance 0.34 + 0.04; postocular distance 0.07 + 0.06; antennal segment lensthiss 1-025) =240/027.110:32, 50025 sim 0.3 4 0.0271V 0.51 =-0:02; rostral segment leneths: 10:34. 01035 0:337== 0025.mm 0.3 + 0.04, IV 0.28 =. 0.02; pronotum: length 0.28 + 0.02, width across humeral angles 0.43 + 0.03, width across anterior margin 0.39 + 0.02; hind leg: femur length 0.59 + 0.02, tibia length 0.68 = 0.02, tar- somere lencths? IF02 "=" O02 41024 5= 0.02. Third instar (Fig. 8D).—Slightly pyri- form, maximum width across abdominal segment V. Dark brown areas of body and antennal segments I to III turning darker. Pronotum and mesonotum with a pair of ir- regular pale yellow bands on each side of middle line. Lateral margins of pronotum pale yellow. Thoracic pleurae dark brown, almost black. Metanotum and first abdom- inal segment pale yellow; metanotum with a few pale brown areas near base. Abdomen pale brown. Abdominal venter with small mesial pale spots, and with numerous setae bent caudally. Ventral spiracles visible on segments II-VIII. Segment IV with a tri- cobothrium situated below spiracle, appear- ring in line with anterior and posterior tri- cobothria of segment V. Segment VI with one anterior and two posterior tricobothria. VOLUME 108, NUMBER 1 115 Fig. 8. Petissius spinipes. A, Egg, dorsal view. B, First instar. C, Second instar. D, Third instar. E, Fourth instar. EK Fifth instar. G, Adult. Scale = 1 mm. 116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Posterior segment and segment VII with two posterior tricobothria. Measurements (n =" 10). Body “length’ 2.79" = 0.32; head length 0.6 + 0.08; width through eyes 0.62 + 0.05; interocular distance 0.42 + 0.04; postocular distance 0.08 + 0.07; antennal segment lengths: 1.033" = "0.07, I 0:43" 0.08, II 0.44 + 0.02, IV 0.6 + 0.05; rostral segment lengths: I 0.42 + 0.05, II 0.44 + 0:05; Ill 0:42 = O05, AV 0:32 0:02; pron- otum: length 0.42 + 0.02, width across hu- meral angles 0.76 + 0.06, width across an- terior margin 0.62 + 0.05; hind leg: femur length 0.77 + 0.09, tibia length 0.94 + 0.19, tarsomere lengths: I 0.3 + 0.04, I O28" = O03: Fourth instar (Fig. 8E).—Oval, with maximum width across abdominal segment V. Very similar to third instar, although an- tennal segment IV may be pale yellow or dark brown; unions between segments red. Pro-, and mesonota, and visible part of me- tanotum with numerous dark brown punc- tures. Anterior margin of pronotum some- times black, with pale yellow bands, ap- pearing only as spots near posterior margin. Mesonotum with a pair of pale yellow bands along wing pads; two bands and two small pale yellow spots on area of scutel- lum. Metanotum with a pale yellow band on each side of middle line. Scent gland openings between segments V—VI with only a small dark brown macula. Labium reaching metacoxae. Metathoracic wing pads covering almost all of metanotum. Tibiae with numerous spines. Conexivum more apparent than in previous instar. Mea- surements (n = 10). Body length 3.64 + 0.4; head length 0.6 + 0.12; width through eyes 0.86 + 0.06; interocular distance 0.5 + 0.06; postocular distance 0.09 + 0.08; antennal segment lengths: I 0.44 + 0.04, II O66" 01055 MIMO0:629 "0:06; IV. 0! 16ee= 0.06; rostral segment lengths: 10.58 + 0.12, IO:627 = 0108 Til O56) == 0075 TV 0.41°= 0.02; pronotum: length 0.63 + 0.08, width across humeral angles 0.98 + 0.06, width across anterior margin 0.71 + 0.09; hind leg: femur length 1.11 + 0.17, tibia length 1.33 + 0.11, tarsomere lengths: I 0.44 + 010351034" = 0103. Fifth instar (Fig. 8F).—Slhghtly pyri- form. Similar to fourth instar. Antennal seg- ment IV dark brown. Apex of femora and tarsi pale yellow. Apices dark brown me- sothoracic wing pads with four, longitudi- nal, pale yellow bands. Scutellum with two pale yellow bands, one on lateral margin and one mesially. Abdominal segments red- dish brown, with joints pale yellow. Mesial maculae of sternites VI—VIII becoming larger and appearing as a continuous mesial band. Other characteristics as in fourth in- star. Measurements (n = 10). Body length 4.45 = 0.88; “head length 0/64 + 0:15; width through eyes 0.89 + 0.15; interocular distance 0.55 + 0.08; interocellar distance 0.04 + 0.03; postocular distance 0.03 + 0.01; antennal segment lengths: I 0.41 + OL MW 0.81 = O47 W-0:78) = 0.097 INGOs + 0.1; rostral segment lengths: I 0.64 + 6:08; Tl 0.59" = 0.147 lb 05>’ = Ole Ii 0.38 + 0.05; pronotum: length 0.66 + 0.02, width across humeral angles 1.14 + 0.19, width across anterior margin 0.81 + 0.26; scutellum: length 0.62 + 0.05, width 0.81 + 0.06; hind leg: femur length 1.25 + 0.24, tibia length 1.48 + 0.43, tarsomere lengths: 1O46 = O63 O38" 0:05. Biology.—Petissius spinipes was collect- ed under the litter of F. cotinifolia in central Veracruz, during April. In southern Vera- cruz, this species was common throughout the year but was more abundant in May and June. First-instar nymphs were usually found under leaf litter of several species of figs, and adults and fifth-instar nymphs were found on superficial layers of leaf lit- ter. Associated fig species included F. in- sipida, F. tecolutensis, and F. yoponensis. In the laboratory, the life cycle of P. spi- nipes took about 53 days. The egg when first laid was white; after three days some red marks started to appear, first on the an- terior pole and then the entire egg; eggs re- mained red for five days before hatching. The first molt appeared after six days. First- instar nymphs were not very active and al- VOLUME 108, NUMBER 1 ways were found feeding inside the fruit. After six days they molted to the second instar, which lasted 11 days, and became slightly more active. Third instars showed similar behavior and lasted 7 days. Fourth instars lasted 8 days and fifth instars 12 days. Their behavior in the field showed that when disturbed, nymphs and adults usually hide in soil crevices. Although this species seems to develop while feeding on fig seeds, it is not always associated with them. Like other Lethaeini species reported in this study, it is considered a facultative terrestrial seed predator of figs. Distribution.—MEXICO: Campeche, Chiapas, Guerrero, Hidalgo, Jalisco, More- los, Nayarit, Oaxaca, Puebla, Queretaro, Quintana Roo, San Luis Potosi, Tamaulipas, and Veracruz. DISCUSSION All Lethaeini -reported in this paper are considered facultative terrestrial seed pred- ators, are extremely rapid in movement, and adults usually run under dead leaves and hide in soil crevices. They seem to be as- sociated with numerous species of Ficus. Cryphula apicata and Paragonatas costar- icensis were the only rare species and were found associated only with F. calyculata and F. trigonata, respectively. Neopetissius slaterorum was associated with seven spe- cies of figs; Cryphula trimaculata, with five species; P. divergens and P. spinipes, with four; and Cistalia pallidifemur with two. These species, as with some Myodochini reported by Cervantes and Gamez (2005), are facultative feeders, and they differ from the Ozophorini (Cervantes et al. 2004) in which all the species reported are obliga- tory and feed exclusively on figs. Although Sweet (1964) reported that Cryphula trimaculata is univoltine with an obligatory adult diapause in Connecticut, the asynchronous fruiting of species of figs recorded as hosts in this study, along with the favorable weather, provide a continuous food supply and permit multiple genera- tions. Nymphs of five species are reported, 117 although not all instars of each are de- scribed. Fourth-instar nymphs of C. trima- culata, P. divergens, and P. spinipes can be distinguished by their general coloration. The posterior pronotal lobe of P. divergens is pale brown, whereas in C. trimaculata and P. spinipes it is mixed with brown and pale yellow. Nymphs of P. divergens are more pubescent than the other species. ACKNOWLEDGMENTS We thank Aaron Sanchez (Benemerita Universidad Autonoma de Puebla) for col- lecting some of the specimens and for the illustrations of P. spinipes. We also thank Iliana Pacheco for collecting most of the specimens from La Mancha, Veracruz. Fi- nancial support for the field work was granted by a CONACYT project (34238-V) assigned to the first author. LITERATURE CITED Baranowski, R. M. and J. A. Slater. 1979. Notes on the biology of two species of Cryphula (Hemip- tera: Lygaeidae) in Trinidad with the description of a new species. Florida Entomologist 62(3): 224-231. Cervantes, P. L. and S. Gamez. 2005. Three species of facultative Myodochini (Lygaeoidea: Rhyparo- chromidae) associated with figs in Mexico. Pro- ceedings of the Entomological Society of Wash- ington 107(2): 362-375. Cervantes, P. L, I. Pacheco, and A. Sanchez. 2004. Immature stages and life cycles of five species of Ozophora Uhler (Hemiptera: Rhyparochromidae. Ozophorini) associated with figs in Mexico. Pro- ceedings of the Entomological Society of Wash- ington 106(3): 654—674. O'Donnell, J. E. 1991. A survey of male genitalia in Lethaeine genera (Heteroptera: Lygaeidae: Rhy- parochrominae). Journal of the New York Ento- mological Society 99(3): 441—470. . 2001. A new genus and five new species of Neotropical Lethaeini (Heteroptera: Lygaeoidea: Rhyparochromidae). Florida Entomologist 84(1): 133-146. Schuh, R. T. and J. A. Slater. 1995. True Bugs of the World (Hemiptera: Heteroptera). Classification and Natural History. Cornell University Press. Ith- aca, New York, 261 pp. Slater, J. A. 1972. Lygaeid bugs (Hemiptera: Lygaei- dae) as seed predators of figs. Biotropica 4(3): 145-151. Slater J. A. and R. M. Baranowski 1973. A review of 118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the genus Cistalia Stal (Hemiptera: Lygaeidae). Florida Entomologist 56(4): 263-272. . 1990. Lygaeidae of Florida (Hemiptera: Het- eroptera) Arthropods of Florida and Neighboring Land Areas, Vol. 14. Florida Department of Ag- riculture and Consumer Services, Gainesville, Florida. 211 pp. Slater J. A. and H. Brailovsky. 2000. Lygaeidae (He- miptera), pp. 319-333. Jn Llorente, B. J. E., E. S. Gonzalez,. and N. Papayero, eds. Biodiversidad, taxonomia y biogeografia de Artropodos de Méx- ico: hacia una sintesis de su conocimiento. Uniy- ersidad Nacional Autonoma de México. Facultad de Ciencias. Slater J. A. and J. E. O'Donnell. 1978. A new species of Cistalia from Brazil and comments on system- atic characters in the Lethaeini (Hemiptera: Ly- gaeidae). Florida Entomologist 61(2): 49-55. Sweet, M. H. 1964. The biology and ecology of the Rhyparochrominae of New England (Heteroptera: Lygaeidae). Part Il. Entomologica Americana 44: 67-71. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 119-124 CUNEOGASTER, A NEW GENUS OF THE SUBFAMILY MICROGASTRINAE (HYMENOPTERA: BRACONIDAE) FROM THE NEOTROPICAL REGION WON-YOUNG CHOI AND JAMES B. WHITFIELD Department of Entomology, 320 Morrill Hall, University of Illinois, 505 S. Goodwin Ave., Urbana, IL 61801, U.S.A. (e-mail: choi @life.uiuc.edu) Abstract.—Cuneogaster, a new genus of the subfamily Microgastrinae from the Neo- tropical Region, is described and illustrated, with C. inae, n. sp., as the type species. The features and relationships of Cuneogaster and several somewhat similar microgastrine genera are discussed. Key Words: species Microgastrinae is a cosmopolitan parasit- oid subfamily recorded trom tropical to arc- tic climates, but especially diverse at the ge- neric level in the tropics (Whitfield et al. 2002). Mason (1981) has estimated that the actual number of world species may be 5,000—10,000, of which only about 1,500 species have been described. Many genera or species groups are endemic to single zoogeographical regions. Taxonomic treat- ments have mainly been focused on tem- perate regions, and the fauna remains poor- ly studied in the highly diverse tropical re- gions. An ongoing revision of the genus Diol- cogaster Ashmead from the New World has revealed some specimens that somewhat re- semble the genus Diolcogaster and often key to this genus using the present micro- gastrine keys (Mason 1981, Whitfield 1997) but fit poorly using Mason’s (1981) diag- nosis. Saeed et al. (1999) redefined the ge- nus Diolcogaster in their revision of the Australasian species, more accurately de- limiting the genus Diolcogaster. Our un- usual Neotropical specimens clearly do not fit within their redefined Diolcogaster. parasitoid, New World, Microgastrine, Cuneogaster inae, new genus, new Here, we describe a new genus, Cuneo- gaster Choi and Whitfield, from the Neo- tropical Region based upon a new species, C. inae Choi and Whitfield. Most speci- mens have been assembled from M. Shar- key and B. Brown’s Colombian Insect bio- diversity project (http://www.uky.edu/ ~myshar0/colombia/welcome.html) except two specimens, from neighboring countries, one from Panama and the other from Ve- nezuela. Relationships of Cuneogaster to other microgastrine genera are discussed. MATERIAL AND METHODS The holotype is deposited in the insect collection of the Humboldt Institute, Villa de Leyva, Colombia. Paratypes are depos- ited in the insect collection of the Humboldt Institute; Illinois Natural History Survey, Champaign, Illinois, USA; National Muse- um of Natural History, Smithsonian Insti- tution, Washington, D.C., USA; Texas A&M University, College Station, USA; and the Canadian National Insect Collec- tion, Ottawa, Canada. Terms for general morphology and wing venation follow Sharkey and Wharton (1997). Additional terms include: lunule (Mason 1981), and 120 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON axillary trough of mesonotum and axillary trough of metanotum (Townes 1969). Specimens for scanning electron micros- copy were dried under room conditions, and mounted on carbon-based SEM subs. Specimens were coated in a Denton Desk II TSC turbo-pumped sputter coater with 40 nm of gold/palladium for 90 seconds and examined under a Philips XL30 field emis- sion environmental scanning electron mi- croscope with Hi-Vac mode. Cuneogaster Choi and Whitfield, new genus (Figs. 1-9) Type species.—Cuneogaster inae, 0. sp. (described below). Diagnosis.—Median and _ lateral ocelli forming almost equilateral triangle. Glossa long and bilobed apically. Antennal plac- odes two-ranked. Mesoscutum without no- tauli. Medioposterior band of scutellum smooth. Metanotum with phragma of scu- tellum slightly exposed laterally. Metano- tum smooth or weakly sculptured. Propo- deum smooth with complete medial longi- tudinal carina, but transverse carinae never present. Fore wing second submarginal cell closed (areolet present). Hind wing vannal lobe slightly convex, margin with evenly destributed row of setae. Hind coxa large, longer than tergite I. Hind tibial spurs un- equal in length, with inner spur always lon- ger than outer spur. Hind tarsal claw with a tooth at midlength, with broad basal lobe. Tergite I narrowing posteriorly, median lon- gitudinal groove of tergite I always present. Tergite II shorter than tergite III, tergite II widened posteriorly, smooth and polished, without well defined median field. Suture between tergite II and tergite HI indistinct in dorsal view. Hypopygium short, as long as hind basitarsus, evenly sclerotized. Ovi- positor short, mostly hidden within hypo- pygium. Ovipositor sheaths short, with se- tae on apical half. Note.—Cuneogaster possesses a unique combination of features that will easily dis- tinguish it from any known genus of Mi- crogastrinae. This new genus appears close- ly related to the genus Diolcogaster Ash- mead by its possession of the following characters: two-ranked antennal placodes, propodeum with medial longitudinal carina but without transverse carinae, second sub- marginal cell of fore wing closed, large hind coxa (always longer than tergite 1), median longitudinal groove of tergite I, short and evenly sclerotized hypopygium, and short ovipositor and ovipositor sheath. It can be separated from Diolcogaster by its long and apically bilobed glossa and posteriorly narrowing tergite I. It is also re- sembles Pseudapanteles Ashmead by its possession of the latter two characters, but differs in the small and evenly sclerotized hypopygium and the short ovipositor sheaths with setae only on the apical half. However, while females are easily diag- nosed, there is currently no unique diag- nostic character system that separates male Cuneogaster alone from other genera that share several key characters. In a preliminary morphological and mo- lecular (16S, 28S and COI genes) phylo- genetic analysis of the genus Diolcogaster and relatives (W.-Y. Choi and J. B. Whit- field, in preparation), we attempted to re- solve phylogenetic placement of Cuneogas- ter. Cuneogaster was supported as mono- phyletic. It was also grouped in a larger clade with Buluka De Saeger, Diolcogaster, Exix Mason, Larissimus Nixon, and Pro- tomicroplitis Ashmead, but the morpholog- ically similar Pseudapanteles was not relat- ed to Cuneogaster. The analysis did not strongly confirm relationships among many genera due to lack of molecular data in some rare genera such as Buluka and Lar- isstmus. Etymology.—The genus name is de- rived from the Latin word ‘cuneus’ (mean- ing wedge shape, because of the apically tapering tergite I) and the greek word ‘gas- ter’ (meaning abdomen). The gender is neuter. VOLUME 108, NUMBER 1 Seppe SEEN: ee DLS Tenis = qe qe RE cess SSE ECS RE ee estes. gS, ae Rare =. Cuneogaster inae. |, Cuneogaster inae Choi and Whitfield, new species (Figs. 1-9) Female.—Body length 1.7 wing length 1.9 mm. Color: Body mostly yellowish brown except dark brown scutellum and metano- tum. Antenna dark brown except yellowish mm. Fore- Fore and hind wings. 2, Head, Head, dorsal view. Scale frontal view. 3, brown pedicel. Maxillary and labial palpi pale yellow. Mesosoma and metasoma yel- lowish brown to dark brown. Legs yellow- ish brown. Head (Figs. 2—4): 1.0—1.1X as wide as scutum. Vertex smooth to weakly punctate and polished, with scattered setae; tangent to posterior margin of median ocellus pass- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 4-9. Cuneogaster inae. 4, Fifth antennal segment, dorsal view. 5, Hind tarsal claw, lateral view, with a tooth indicated by arrow. 6, Mesosoma, dorsal view. 7, First—third tergites, dorsal view. 8, Metasoma and ovipositor mechanism, lateral view. 9, Male genitalia, dorsal view. Scale bars = 40 jm. ing in front of lateral ocelli; distance be- tween lateral ocelli 0.4 distance from lat- eral ocellus to edge of eye. Face 0.8—0.9X as broad at midheight as long medially, weakly punctate with scattered setae and with faint median longitudinal carina in dorsal one-third to half. Clypeus 1.7—1.8X as broad as its height. Malar space 0.2 eye height in frontal view. Eyes 1.3—1.4 high- er than width. Temple smooth to weakly VOLUME 108, NUMBER 1 punctate with scattered setae. Glossa long and bilobed apically. Antenna slightly lon- ger than forewing; first flagellomere 3.4 x as long as wide. Mesosoma (Figs. 5—6): 1.1—1.2X longer than high. Scutum 1.2—1.3X as wide as long, weakly punctate with scattered setae. Dorsal scutellum 1.1X as wide as long, smooth medio-anteriorly to punctate poste- riorly with scattered setae. Axillary trough of mesonotum generally coarsely carinate. Lunule of scutellum smooth and 0.3—0.4x as high as axillary trough of mesonotum, with nearly uniform height. Medioposterior band of scutellum smooth. Metanotum gen- erally smooth. Axillary trough of metano- tum smooth with scattered setae in posterior half. Medioposterior band of metanotum smooth, flat to slightly convex. Medioan- terior pit of metanotum without longitudinal carina. Propodeum 2.4 as wide as long, smooth with sparse setae. Medial longitu- dinal carina apparently formed by incom- plete fusion of two longitudinal carinae. Pronotum laterally granulate with ventral groove. Propleuron smooth to weakly punc- tate without propleural flange. Mesopleuron mostly smooth and polished with scattered setae anteriorly, anterodorsally and ventral- ly. Metapleuron mostly smooth and _ pol- ished with scattered setae posteriorly. Hind coxa 2.1—2.2X as long as wide, 2X as long as first tergite, smooth and polished later- ally, punctate dorsally and postero-ventral- ly. Inner hind tibial spur 1.6—1.8X as long as outer spur, 0.6 as long as hind basitar- sus. Hind tarsal claw with single tooth in median portion, with broad basal lobe. Wings (Fig. 1): Fore wing evenly setose. Stigma 2.5X as long as wide, vein r arising from middle of stigma, vein RI 1.3—1.4x as long as stigma. Vein 1M 0.7% as long as vein (RS+M)a, 2.0X as long as vein m- cu. 3RSa present. Areolet subtriangular. Vein r-m spectral. Hind wing evenly setose. Vannal lobe slightly convex with even fringe of setae. 1r-m 0.6X as long as 1RSa, 1.2X as long as 2r-m, 0.4 as long as IRSb. [23 Metasoma (Figs. 7-8): Tergite I narrow- ing posteriorly, 3.7 as long as apical width anterior two-third smooth, glabrous, posterior one-third smooth to weakly ru- gose with sparse setae. Median longitudinal groove strongly present over anterior two- thirds, weakly impressed to absent on pos- terior one-third. Tergite I] 0.6 as long as tergite I, 0.8X as long as tergite III, 1.7 as wide at posterior end as long, median tergite widened posteriorly, smooth and polished with sparse setae. Tergite III smooth and polish with sparse setae. Suture between tergite II and tergite III indistinct dorsally. Tergite IV to VII smooth with sparse setae. Hypopygium smooth with sparse setae, medioventral length 1.0 as long as hind basitarsus. Ovipositor sheaths short with setae in apical half. Male.—As for female; Fig. 9 shows male genital capsule. Material examined.—Holotype: 2°, CO- LOMBIA: Magdalena, PNN [= Parque Na- cional Natural] Tayrona Pueblito, 11°20'N T4202 W,, 50 m5 7 Sa 2000. R Hen fiquez ee... M.789™ baraty pes lurc © ©- LOMBIA: same data as holotype; 2 4, Cundinamarca PNN Chingaza Valle del Fraviejon, 4°31°N 73°45°W 3,170 m, 31.vi11.2000—13.1x.2000, A. Pérez Leg. M.732; 1 ¢, Magdalena, PNN Tayrona Zai- no, 11°20’N 74°2’W 50 m, 28.vii.2000— 14.vi1.2000, R. Henriquez Leg. M.567; 4 36, Magdalena, PNN Tayrona Pueblito, 11°20)N. 74°2'W_ 0225. 4m: .26:%<2000= 22.xi.2000, R. Henriquez Leg. M.944; 1 6, Magdalena, PNN Tayrona Canaveral, Li 20"N 7422" WwW 30" im, 194x%22000= 30.ix.2000, R. Henriquez Leg. M.628; 1 d, Magdalena, PNN Tayrona Zaino, 11°20'N 74°2’W 50 m, 17.x.2000—3.xi.2000, R. Henriquez Leg. M.789; 2 6, Bolivar, SFF [ = Santuario de Fauna y Flora] Los Co- lorados Venado, 9°54'N 75°7’W 320 m, 2.%.2000—17-x.2000, EE. Deuluteut Mees. M.766; 2 3d, Magdalena, PNN Tayrona Zai- nos bio 2OUIN 7452" We S50 in= 294%. 2 000— 17.x.2000, R. Henriquez Leg. M.793; 4 6, Magdalena, PNN Tayrona Zaino, 11°20'N 124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 74°2'W 50 m, 3.xi.2000—22.xi.2000, R. Henriquez Leg. M.941; 1 3d, Magdalena, PNN Tayrona Zaino, 11°20'N 74°2'W 50 m, 4.xii.2000—15.xii.2000, R. Henriquez Leg. M.966; 4 2, 1 3d, Magdalena PNN Tayrona Cafiaveral, 11°20'N 74°02'W, 30 mee 2=—22,1- 2001, Rk; «Menriquez bez; M.1207; 1 2, 3 6, Magdalena PNN Tay- rona Pueblito, 11°20’N 74°02'W, 30 m, 4— 15.xii.2000, R. Henriquez Leg., M. M. 962. 1 6, PANAMA: Las Cumbres, 26.v- 8.vi.1982, H. Wolda. 1 6, VENEZUELA: Aragua Cumbolo, 100 m, 12.1.1996, R. Wharton. Hosts.—Unknown. Comments.—The species is named after Dr. Kyong-in Suh, for her unlimited support of W.-Y. Choi. ACKNOWLEDGMENTS We thank Mike Sharkey of the University of Kentucky, Bob Wharton of Texas A&M University, and Henri Goulet of the Cana- dian National Insect Collection for loan of specimens. We also thank Josephine Rod- riguez of the Department of Entomology, University of Illinois, and Scott Robinson of the Imaging Technology Group, Beck- man Institute, University of Illinois, for aid in producing the SEM and ESEM photos. This work is supported in part by USDA Townes, H. NRI-CGP grant number 2003-35316-13679 and NSF DEB 0346712 awarded to JBW, and NSF DEB 9972024 and DEB 0205982 awarded to Michael J. Sharkey and Brian V. Brown. LITERATURE CITED Mason, W. R. M. 1981. The polyphyletic nature of Apanteles Foerster (Hymenoptera: Braconidae): a phylogeny and reclassification of Microgastrinae. Memoirs of the Entomological Society of Canada 115: 1-147. Saeed A., A. D. Austin, and P. C. Dangerfield. 1999. Systematics and host relationships of Australasian Diolcogaster (Hymenoptera: Braconidae: Micro- gastrinae). Invertebrate Taxonomy 13(1): 177— 178. Sharkey, M. J. and R. A. Wharton. 1997. Morphology and terminology, pp. 19-37. Jn Wharton, R. A., P. M. Marsh, and M. J. Sharkey, eds. Identification Manual of the New World Genera of the Family Braconidae (Hymenoptera). International Society of Hymenopterists Special Publication 1, 439 pp. 1969. The genera of Ichneumonidae, part 1. Memoirs of the American Entomological Insti- tute 11: 1-300. Whitfield, J. B. 1997. Chapter 29. Subfamily Micro- gastrinae, pp. 333-364. Jn Wharton, R. A., P. M. Marsh, and M. J. Sharkey, eds. Identification Manual to the New World Genera of the Family Braconidae (Hymenoptera). International Society of Hymenopterists Special Publication 1, 439 pp. Whitfield, J. B., PB. Mardulyn, A. D. Austin, and M. Dowton. 2002. Phylogenetic relationships among microgastrine braconid wasp genera based on data from the 16S, COI and 28S genes and morphol- ogy. Systematic Entomology 27(3): 337-359. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 125-130 TWO NEW SPECIES OF MYRMELEON LINNAEUS, 1767 (NEUROPTERA: MYRMELEONTIDAE) FROM CHINA, WITH A KEY TO CHINESE SPECIES BAO RONG AND XINLI WANG Department of Entomology, China Agricultural University, Yuanmingyuan West Road, Beiing 100094, China (e-mail: brbrong@ yahoo.com.cn). Abstract.—Two new species of the genus Myrmeleon Linnaeus, 1767, from China are described, M. circulis and M. trigonois. A key to the known species of Myrmeleon from China is presented. Key Words: The genus Myrmeleon Linneaus, 1767, is worldwide in distribution with about 160 known species, 27 of which occur in the Palearctic and Oriental regions. Eleven spe- cies are known in China. Myrmeleon bore (Tjeder) and M. formicarious L. occur throughout most of China, Myrmeleon im- manis Walker is found only in the Palearc- tic part of China, and the other eight species occur only in the Oriental Region of China. The new species described below were dis- covered in Fujian, Guangxi, Guizhou, and Hubei provinces and in Yunnan and Sich- uan provinces, all in the Oriental Region. Myrmeleon is characterized by the fol- lowing characters: Vertex slightly to mod- erately raised; antennae with well defined club; wings narrow to moderately broad; posterior Banksian line in forewing usually strongly developed, anterior Banksian line rarely evident; posterior Banksian line in hind wing sometimes well developed: cos- tal cells simple; Rs of forewing arises be- yond cubital; forewing vein CuP originates at or very near basal crossvein; forewing vein 2A runs close to 1A for short distance, then bends at sharp angle toward 3A; bas- itarsus of hind leg shorter than apical tar- somere; female posterior gonapophyses Neuroptera, Myrmeleontidae, Myrmeleon, new species, China usually slender, sometimes very short, an- terior gonapophyses rounded, spermatheca slender with duct usually coiled or convo- luted; male ectoproct often expanded ven- trally, sometimes simple, gonarcus arched, mediuncus usually well developed. All types are deposited in the Insect Col- lection of the China Agricultural Universi- ty, Beijing China. KEY TO SPECIES OF MYRMELEON FROM CHINA 1 Head black, yellow or brown with distinct manrksy (Sinaia tose sae2s. eee en ne 2 — Head black without distinct marks ........ 8 2 Forewing costal cells wider than high above radial sector Be apes se M. punctinervis Banks — Forewing costal cells much higher than wide above radial sector (as in Figs. 1,6) ...... 3. Tibial spurs of foreleg reaching to about mid- M. immanis Walker — Tibial spurs of foreleg about equal to or short- dlevof 2ndvtarsomere)y. 5 4) er in length to basitarsus 4 Head black, occiput black with a yellow an- mulus? (Figs 2)). cea tns eek. M. circulis, n.sp. Head yellow with brown or black marks ... 5 5 Pronotum without stripes except narrowly pale M. alticola Miller and Strange — Pronotum with distinct yellow stripe (Fig. 7) brown anteriorly 6 Hind wing with 4 crossveins before origin of Rs (one specimen with hind wing with 5 crossveins before origin of Rs) Sh Se hy OER A cer M. bimaculatus Yang — Hind wing with no less than 5 crossveins be- fore origin of Rs (Fig. 6) 7 Pronotum with two median yellow stripes in- tercrossed (Fig. 7) M. trigonois, n.sp. — Pronotum with one median yellow stripe .. . FSET One a ee CR RCS. Beet aee M. fuscus Yang 8 Tibial spurs of hind leg shorter than basitarsus RUN ooence Sroka ise M. wangi Miller and Stange — Tibial spurs of hind leg equal to or longer than basitarsus 9 Tarsomeres pale brown expect at apices .... A et eee M. heppneri Miller and Stange — Tarsomeres all black or dark brown except PalesDLOwPGOrsalllivemnen eer eon n eer tet 10 10 Tarsomeres with pale brown markings dorsally Bey LS Knee OnE, Set Conen, Cre a Maa ors kiss Atremane 11 —elarsomencestrall black ovaries) erento 12 11 Male pilula axillaries small, knob not much wider than pedicel with setae about as long as knob ; mesonotum usually all dark brown pos- terior M. taiwanensis Miller and Stange — Male pilula axillaries large, knob much wider than pedicel with matlike setae which are much shorter than knob; mesonotum pale brOwnyPOStenlOnN ces a oer ae) es es M. persimilis Miller and Stange M. bore (Tjeder) 12 Length no more than 33 mm .. — Length more than 35 mm er iowe tics a Pas Meise tee ame M. formicarious Linnaeus Myrmeleon circulis Rong and Wang, new species (Figs. 1-5) Diagnosis.—Black; vertex inflated; frons, vertex black; occiput black with a yellow annulus. Pronotum black with a median stripe. Description.—Male body length 32 mm, forewing 34 mm, hind wing 31 mm. Fe- male body length 30—34 mm, forewing 35— 39 mm, hind wing 33-36 mm. Head (Fig. 2): Vertex inflated; frons, vertex black; occiput black with a yellow annulus; clypeus and labrum pale brown. Maxillary palpus yellow brown, terminal segment black; labial palpus yellowish, ter- minal segment long, spindle-shaped, black. Antenna short, less than 1/4 forewing length, slightly thickened apically, flagel- lum with about 34 segments, black, pale brown on base and on inner surface of scape. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Thorax: Pronotum (Fig. 2) slightly shorter than broad, black except a median stripe and lateral edges pale brown; setae on pronotum long, curved. Mesothorax with tergum black anteriorly, with long black setae; pale brown posteriorly with long white setae. Metathorax with soft white setae on tergum. Legs: Foreleg with coxa yellow brown except black on outer surface, trochanter yellow brown, femur with black stripe and long white and short black setae, basally with a long sensory hair, tibia brown except yellow brown on outer surfaces, with white and black setae, tarsus brown, basitarsus yellow brown, about % length of distal tar- somere; tibial spurs about equal in length to basitarsus, claw about % length of tibital spur. Middle and hind legs similar to fore- leg, hind leg without long sensory hairs. Wings (Fig. 1): Hyaline, longitudinal veins mostly alternating pale and dark brown except some pale brown; nearly all crossveins pale brown; membrane without marking; stigma white, wider than high; posterior Banksian line conspicuous in both wings; hind wing with 5 crossveins before origin of Rs; pilula axillaries with knob much wider than long, with dense setae. Abdomen: Black with apices of tergites and sternites pale black; shorter than fore- wing; ectoproct produced ventrally; para- meres and mediuncus as in Fig 5. Female genitalia (Fig. 3): Ectoprocts round with long, erect black setae; posterior gonapophyses digitiform, with long setae; anterior gonapophyses large; lateral gona- pophyses separated from each other by nar- row membranous area; pregenital plate small. Spermatheca (Fig. 4) curved, broad at base, slightly tapering toward apex. Type material.—Holotype: ¢, Fujian: Jianzhen, Dongfeng, 28,07,1986, leg. M.Xie. Paratypes: 4 °, Fujian: Xishui, Jin- jiang, 24~30,09,2000, leg. Q.Zh.Song; 1 2, Guangxi: Longzhou, Nonggang, 18,05,1982, leg. Ch.K. Yang; 12, Guizhou: Hezhang, 28,06,1978, leg. Unknown; | @, Fujian: Yanluxia, 15,05,1981, leg. Un- VOLUME 108, NUMBER | | 5 Figs. 1-5. Myrmeleon circulis. 1, Wings. 2, Head and pronotum. 3, Female genitalia, ventral. 4, Spermatheca. 5, Male genitalia, caudal. Scale bars: 1 = 1 cm; 2 = 1 mm; 3-5 = 0.5 mm. 128 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ) 10 Figs. 6-10. Myrmeleon trigonois. 6, Wings. 7, Head and pronotum. 8, Female genitalia, ventral. 9, Sper- matheca. 10, Male genitalia, caudal. Scale bars: 6 = 1 cm; 7 = 1 mm; 8 = 0.5 mm; 9-10 = 0.25 mm. VOLUME 108, NUMBER 1 known; 1 2, Hubei: Zigui, Jiutouling (110 m), leg. J. Yao. Remarks.—This species is similar to M. wangi but it can be separated from the latter by the stripes on the pronotum and the male genitalia. In M. wangi, the pronotum is nearly completely black except narrowly pale brown anteriorly and laterally, and the gonarcus is slightly curved (see fig. 25 in Miller et al. 1999); in M. circulis, the pron- otum is black with a median stripe and the gonarcus is strong curved. Etymology.—The specific name refers to the black occiput with a yellow annulus. Myrmeleon trigonois Rong and Wang, new species (Figs. 6—10) Diagnosis.—Black, vertex inflated, pale brown with several black spots, pronotum slightly shorter than broad, with two me- dian intercrossed yellow stripes. Description.—Male body length 31 mm, forewing 30 mm, hind wing 27 mm. Fe- male body length 30 mm, forewing 35 mm, hind wing 32 mm. Head (Fig. 7): Vertex inflated; frons black; vertex and occiput pale brown with several dark marks. Clypeus and labrum pale yellow brown. Maxillary palpus yel- low brown, terminal segment brown; labial palpus yellowish, terminal segment long, spindle-shaped, dark brown. Antenna yel- low on base and on inner surface of scape. Thorax: Pronotum (Fig. 7) slightly shorter than broad, dark brown, with two median intercrossed yellow stripes, lateral edges pale brown; setae on pronotum long, curved. Mesothorax with tergum black an- teriorly with long black setae, pale black posteriorly with long white setae. Metatho- rax with soft white setae on tergum. Legs: Foreleg coxa, femur and tibia dark brown on outer surface, otherwise pale brown, femur with long white and shorter black setae, basally with a long sensory hair, tarsus brown, basitarsus yellow brown, about % length of distal tarsomere; tibial spurs about equal in length to basitarsus, 129 claw slightly curved, about *%4 length of ti- bital spur. Middle and hind leg similar to forelegs, hind leg longer than foreleg with- out long sensory hairs. Wings (Fig. 6): Hyaline, longitudinal veins mostly alternating yellow and brown: nearly all crossveins pale brown; membrane without marking; stigma white, wider than high; posterior Banksian line conspicuous in both wings; hind wing with 5 crossveins before origin of Rs; pilula axillaries with knob much wider than long, with dense se- tae. Abdomen: Shorter than forewing, brown to black with apices of tergites and sternites pale brown; covered with short white hairs, terminalia with sparse short white and black setae, ectoproct produced ventrally; para- meres and mediuncus as in Fig. 10. Female genitalia (Fig. 8): Ectoprocts round with long erect black setae; posterior gonapophyses digitiform, with long setae; anterior gonapophyses round; lateral gona- pophyses separated from each other by nar- row membranous area; pregenital plate small. Spermatheca (Fig. 9) curved. Type material.—Holotype: ¢,Yunnan: Degqin, Meilishi (2,200 m), 20.07.1982, leg. S.Y.Wang. Paratypes: 4 2°, Sichuan: Batang (2,600 m), 14.08.1982, leg. L.Ch.Niu & S.Y.Wang; 1 ¢, Sichuan: Batang (2,500 m), 13.08.1982, leg. S.Y.Wang. Remarks.—This species is similar to M. formicarious, but it can be separated from the latter by the stripes on the pronotum (Aspock et al. 1980). In M. formicarious, the pronotum is black without a median yel- low stripe, but the new species has two me- dian intercrossed yellow stripes on the pronotum. Entymology.—The specific name refers to the two triangular marks on the prono- tum. ACKNOWLEDGMENTS We are very grateful to Mr. Qiongzhang Song, Mr. Ming Xie, Mr. Chikun Yang, Mr. Chunlai Niu, and Mr. Shuyong Wang for collecting specimens. 130 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Miller, R. B., L. A. Stange, and H. Y. Wang. 1999. New species of antlions from Taiwan (Neuroptera: Aspéck H., U. Aspéck, and H. Holzel. 1980. Die Neu- Myrmeleontdiae) . Journal of the National Taiwan ropteren Europas. Goecke & Evers, Krefeld, Ger- Museum 52(2): 47-78. many. Vol. I, pp. 281-311; Vol. Il, pp. 182-210. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 131-138 ADULT CADDISFLY (TRICHOPTERA) PHENOLOGY AT THE HANFORD REACH NATIONAL MONUMENT, WASHINGTON STATE RICHARD S. ZACK, DAVID E. RUITER, DENNIS L. STRENGE, AND PETER J. LANDOLT (RSZ) The James Entomological Collection, Department of Entomology, Washington State University, Pullman, WA 99164-6382, U.S.A. (e-mail: zack@wsu.edu); (DER) 6260 South Grant Street, Centennial, CO 80121 (e-mail: druiter@msn.com); (DLS) Battelle Pacific Northwest National Laboratory, Battelle Boulevard, Richland, WA 99352; (PJL) USDA, ARS, 5230 Konnowac Pass Road, Wapato, WA 98951 Abstract.—Adult caddisflies were sampled on the Wahluke Wildlife Area and Saddle Mountain National Wildlife Refuge subunits of the newly created (2000) Hanford Reach National Monument using 15-watt “black lights” from April 2002 through April 2003. A diverse fauna consisting of nine families, 21 genera, and 33 species were collected. Protoptila coloma Ross, Agraylea multipunctata Curtis, Hydroptila xera Ross, Ceraclea alagma (Ross), Nectopsyche lahontanensis Haddock, Oecetis cinerascens (Hagen), and Ylodes reuteri (MacLachlan) represent new records for Washington State. Species com- position and phenology are presented in tabular form. Key Words: During the period April 2002 through April 2003, entomological survey studies were conducted on the Hanford Reach Na- tional Monument located in south central Washington State. The Monument was cre- ated in 2000 from portions of the Hanford Nuclear Site and is administered by the United States Fish and Wildlife Service and the United States Department of Energy. The Monument includes a variety of sub- units but those surveyed for this study were the Wahluke Wildlife Area and the Saddle Mountain National Wildlife Refuge that are located in portions of Franklin, Grant, and Adams counties that lie to the north of the Columbia River (Fig. 1). Newell et al. (2001) conducted a survey of caddisflies at the two largest spring- stream systems on the Monument—Rattle- snake and Snively Springs. These springs are found in the Fitzner-Eberhardt Arid Lands Ecology Reserve subunit of the Monument, and are situated approximately Trichoptera, Hanford, survey, Washington State, caddisflies 20 km from where our studies were con- ducted. Newell et al. (2001) found a total of 26 taxa based on adults but noted that previous benthic studies (Gaines 1987a, b; Gaines et al. 1989, 1992; Newell 1998) had revealed only four genera of caddisflies based on larval collections from the springs. Newell et al. (2001) postulated that most of the caddisflies they obtained during their survey work did not originate from the spring systems, but from the nearby Colum- bia River. Because of habitat differences found on the Wahluke and Saddle Moutain subunits and their closer proximity to the Columbia River, we decided to undertake a more inclusive study of the Monument in order to verify and add possible new re- cords, as well as verify and add to available phenological data. SITE DESCRIPTION Climate at Hanford is best characterized as semi-arid with hot and dry summers and 132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rr | Wahluke ical Wahluke and Saddle Mountain subunits. cold winters. Precipitation ranges from 30— 35 cm at the crest of Rattlesnake Ridge to less than 12 cm along the Columbia River. Temperatures range from an average of 3°C in January to 33°C in July; temperatures of 32°C or above occur an average of 56 days per year (ERDA 1975). The Wahluke and Saddle Mountain sub- units of the Hanford Reach National Mon- ument are bordered on the south by one of the few free-flowing stretches of the Co- lumbia River. The units primarily consist of Central Hanford Richland WA | Subunits of the Hanford Reach National Monument. Sampling during this study took place in the sandy soils supporting native shrub-steppe plant communities consisting mainly of shrubs including sagebrushes and_ rabbit- brushes, Sandberg’s bluegrass (Poa sand- bergii Vasey), perennial bunchgrasses, and a variety of forbs; invasive cheatgrass (Bro- mus tectorum L.) also is common in dis- turbed areas. Details of the flora are found in Sackschewsky et al. (1992). Aspects of both the flora and fauna are treated in Downs et al. (1993). The Wahluke and Sad- dle Mountain subunits contain numerous VOLUME 108, NUMBER 1 lakes and wetlands created and sustained by raised water tables associated with major ir- rigation canal return systems referred to as the Saddle Mountain Wasteway and WB-10 Wasteway. This wastewater return provides the only source of water on the sites besides the Columbia River. This wastewater return provides for a series of lakes and shallow pools that are interconnected by numerous canals. The system prevents wastewater from empyting into the Columbia River. Ir- rigation return flow quality is discussed by Johnson and Prescott (1981). Numerous collection sites were estab- lished and maintained throughout the Wah- luke and Saddle Mountain subunits from 1 1 April 2002 through 15 April 2003. Sites were established at various locations in- cluding within 100 m of the Columbia Riv- er, in the riparian areas surrounding one of the larger lakes (700 acres), next to several of the shallow irrigation runoff ponds, along several of the canals, and at a variety of areas throughout the shrub-steppe habi- tat. These latter collecting sites were estab- lished at distances of less than 5 m to no more than | km from a source of water. Specific sites were trapped on an irregular basis but, at least three different sites were trapped on a weekly basis. Collecting was not conducted in January or February. Col- lection information from specific trapping sites was maintained but, as we did not find specific trends in caddisfly species per site, these data are not included in this paper. MATERIALS AND METHODS Adult caddisflies primarily were collect- ed through the use of 15-watt “‘black light” traps that were established for both cad- disfly and moth collections. Black light traps consisted of six gallon plastic buckets over which a metal funnel was placed. A 15-watt, fluorescent tube was placed verti- cally over the funnel. In some cases, the bulb was surrounded by a series of four vanes while in other traps, a bulb was sim- ply hung over the funnel. We did not mon- itor for differences in capture rates between the two types of traps, but no evident dif- ferences were noted. Ethyl acetate was used as a killing agent in the traps. Traps were usually established within an hour of sun- down and collected within an hour of sun- rise. Running times for traps depended on the date of collection (1.e., shorter in the spring and longer in the summer and fall). Trap contents were collected each morn- ing and placed into containers for return to the laboratory. Caddisflies were sorted un- der a microscope at the laboratory. If the number of caddisflies in a trap was less than 200 and appeared to consist of at least three taxa, all of the specimens were retained and identified. If the number of specimens in the trap was greater than 200, specimens were sorted into morphologically similar units and up to 50 specimens of each were retained; the remainder were discarded. In several instances, over 3,000 caddisflies were found in a single trap. Relative abundance figures (Table 1) are subjective and are based on the number of specimens of a taxon collected during a giv- en night and during the season. If a species was collected on only one date during the year or if less than 10 total specimens were collected, it is listed as rare (R) in the table. If a species was collected on at least two separate dates or between |1 and 50 were taken in total, it is listed as occasional (O). If a species was collected on at least two separate dates or between 51 and 200 were collected, it is listed as common (C). If a species was collected on at least two sepa- rate dates or more than 201 specimens were collected, the species is listed as abundant (A). All specimens were identified by DER, and voucher material is housed in the James Entomological Collection, Washington State University. RESULTS AND DISCUSSION Table 1 lists the 40 taxa that were col- lected during this study; 33 of which were identified to species; specimens of Leuco- trichia were only identified to the generic level. Other taxa listed at the generic level PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 134 e e Ps (syueg) DADAD $1J29aE e e e ‘ds ayodsdojoan e Psi yooppepy sisaunjuoyv) ayodsdojoan e e Ps | UYJUD-[[IH Vviuquiypyjyp sapiovisKp e Ps I (SSOY) VUSYDID DajIvIAD AV dese OO Lda @ e ‘ds pwojsopidaT e e P| (syueg) Wwinasauid pDuolsopidaT AV CILVNOLSOCIdH 1 e a (SSOY) VAlAP VIJIIGOJIVIS ‘ds piyoijoonaT e 6 e ‘ds pjpudoupay SSOY Véax DINdOApPAL Sssoy vsOsiv YjIIdOApAH ssoy vYIID YjIUdOAPAH ssoy xplp vjudoipaAH ® e e Onw~Oo = Soa s ee Beton NS NM AN — — ~~ : aa oa) ~ @8€ 6 Sh a SENT Ces | Sa ey oo yD ways | | a fos] Fe ee ee asa tN Tr aM FE A NS ae SSSeoqs see fon) OMaAnanm See ic Pa ee ~ ' _ TANIA OMM or AA YH rae eat Seo = are tl NN + — NANMA A SSS Ss SS eee GN (GN am om = = lial Mc | cal AANA \O ama NG) ACG) —~ (oa) SIR), ae NN i IGN = fo Va) Ly = a (ence ar | - CO ~~ WT gud | fos amnXN N NNR ANNAN SS SRS OR UR a a a 10 1-3 (2) Specimens studied are as follows: Costa Rica, Heredia, Puerto Viejo, Finca La Sel- va. ViIlE7-71, 1. 232) 35 WePe! ex natural containers (A. Berrios Arias); Panama, Da- rien, Puero, Rio Pucro, II-24—64, 1 2,1 6, LePe ; ex rockhole (A. Quinonez). Description of the Pupa of Lutzia trigripes (Fig. 3) Range and modal number of branches in Table 3. Cephalothorax: Light brown, darker posteriorly on scutum, setae 1, 3-5, 7-9-CT long, 2-CT moderately long, 6-CT short, all single, 10-CT long, with 4—6 br, 11,12-CT long, single; Trumpet: Medium brown, reticulate except for small tracheoid patch measuring 0.15—0.22 subbasally, length 1.01—1.25, x 1.11 mm, index 2.88— 4.31, x 3.64, pinna 0.4—0.53, x 0.45 mm. Abdomen: Light brown, darker on HII, intersegmental membranes II-III and I-IV reticulate medially; length 5.05—6.59, x 5.72 mm; setae O-II-IX, 9-H-VI, 14-II- VIII minute, single, seta 2-I—VII, 11-II-— VII short, single; seta 1-I dendritic, with 7,8 primary branches, 1-II short, pedunculate, usually 4 br (3-6), 1-HI—VII long, 1-H usually 6—8, seldom 9—11 br, 1-IV usually 4, 5 br, seldom triple or 6 br, 1-V—VI single to triple, 1-VII single; seta 1-IX present; seta 3-I-III long, stout, single or double, 3- IV, VI-VIII long, [V with 4—6 br, 1-VI, VIII single, 3-V moderately long, double to qua- druple; seta 4-I short, usually with 4 br (3— 7), 4-H-V,VIU long, 4-IL,IV,V,VII single or double, 4-III with 3—5 br, seldom double, 4- VI moderately long, single or double; seta 5-I-III short, 5-I,II single to triple, 5-III double to quadruple, seldom with 5 br, 5- IV—VII long, 5-IV with 3-5 br, 5-V—VII single or double, rarely triple; seta 6-I-VI long, 6-I,VI stout, single or double, 6-II-V single, rarely double, 6-VII moderately long, with 2—4 br, seldom single; seta 7-I very long, single, seldom double or triple, 7-H, VI,VII long, single or double, 7-III,V short. usually 4—6 br, seldom double or with 7 br, 7-[V moderately long, single to triple, seldom quadruple; seta 8-III-VII VOLUME 108, NUMBER | 151 Pa ST Krtisak? UL Fig. 3. Pupa of Lutzia tigripes. A, Cephalothorax. B, Metanotum and abdomen. CT = cephalothorax, GL = genital lobe, Pa. = paddle. short to moderately long, 8-III,I[V usually rarely 7 br, 9-VIII usually 8—10 br, seldom with 3—6 br, V—VII usually single to triple, 7 or 11-16 br. Paddle: Ovoid., margin seldom with 4—6 br; seta 9-VII, VIII long, smooth, midrib not reaching apex, length stout, aciculate, 9-VII usually with 3-6, 1.19—-1.54, x 1.37 mm, index 1.03-1.26, x ia tigripes. Range is followed in parenthesis by model numbers. j Setal branching of the pupa of Lut Table 3. Abdominal segments Paddle IX Vil Vill VI IV Il Cephalothorax Seta (1) 24 (3) 3-6 (4) l 4—6 (5) 2 (1) 6-11 (8) | 3-5 (3) Il 73 (il) 3-6 (4) | Ie 1-3 (1) Teron) | ) N 2 ()) il. 3-7 (4) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ee) I Lalas = | ™ — —— foe) oom @) = g © -- AN nN on GEG ech ae sé | _—~ oF Ss fon) >a ey + SO Ciltaen an NON | fixe loa) —~ —~ MO c- AQ. -I =a Va) =e iE Tee fae) = | — PE SSeS EIR) SS ECO weal el Nam ol a maa = C: = on] wy = SS mmo la = =i =e Se = NM + WNOMm OD 2 (1) IF IL, 2 (GD) 1-3 (?) 2-6 (6) 10 2A) 1.17, emarginate at apex where setae are attached Specimens studied are as follows: Yem- en, Vaizz, XTI-1-51) I> 2 sleds Pekewex basin (K.L. Knight), Liberia, Roberts Field, no date, #43a, 1 6, PeLe, #33a,b, no date, 1 2,1 3d (Gelfand); Senegal, M’bao, no date, #10257, 1 2, PeLe, ex polluted ditch. KEY TO KNOWN PUPAE OF THE GENUS LUTZIA il. Intersegmental membranes of abdominal segments II/III and ILI/IV reticulate; seta 3- I[,1ll usually single; paddle emarginate api- cally (except in Lt. shinonagai) ........ 2. — Intersegmental membranes of II/III and III/ IV smooth, without reticulation; seta 3-IL, [II usually double; paddle straight apically (Subgenus! uid) wee | eee 5 2(1). Seta 5-II laterad of seta 4-II; postero-lateral corner of VIII not produced; seta 9-II-VI pointed at tip (subgenus /nsulalutzia) .... OD sala teehee ake ee ee ee ee shinonagai - Seta 5-II mesad of seta 4-II; postero-lateral corner of VIII rather sharply produced; seta 9-II-VI blunt at tip (subgenus Metalutzia) 3(2). Seta I-III, ['V 0.5 or less length of following segment; seta 5-[V—VI shorter than follow- Ing SESMeNt,...2 wanes se ee trigripes ~ Seta I-III, IV 0.7 length of following seg- ment; seta 5-[V—VI longer than following Lerswmiy fy Fe... e's Seen e cane Tee 4 4(3). Longest setae on abdominal segments [V— VI usually longer than following tergum . . cothe Ralahipe Fos @ aoe a Sha ee kee See vorax ~ Longest setae on abdominal segments [V— VI usually shorter than following tergum . . = si 4 hope bye ewe eS © cea halifaxii, fuscana 5(1). Seta 11-I,VII double or triple; trumpet without subapical expansion ..... allostigma = Seta 11-III, VII single; trumpet with subapi- cal expansion, flared apically ........ bigoti KEY TO THE FOURTH INSTAR LARVAE OF THE GENUS LUTZIA Since I had access to larval exuviae of the species of Lutzia, or to the descriptions of the larva of Lt. shinonagai.and Lt. vorax, and since a key to all the species worldwide has never been produced, it seemed appro- priate to include one in this work. Descrip- tions of the larvae for the New World may be found in Bonne and Bonne-Wepster VOLUME 108, NUMBER 1 (1925), Senevet and Abonnenc (1939) and Lane (1953), and for the Old World, Bram (1967), Tanaka et al. (1979), Hopkins 952). ite Siphon index 3.0 or greater, setae shorter than width of siphon at point of attach- MEM SUDSCMUSELUIZIG) sae ie ee fae ae 2 = Siphon index 2.0 or less, setae much longer than width of siphon at point of attach- ment (@OldwWorldsspecies) ieee ae alee oe) 3 . Pecten confined to basal 0.4 of siphon, usu- ally. aI OnumMOTe. =.= = cesses < Seas: allostigma — Pecten extending to 0.7 length of siphon usually 10 or fewer bigoti 3(1). Pecten closely spaced, confined to middle 0.3 of siphon; with 18—20 comb scales (sub- genus /nsulalutzia) shinonagai - Pecten more widely spaced, extending to near apex of siphon, with more than 30 comb scales (subgenus Metalutzia) ...... 4 4(3). Siphon with 10 or more pecten spines tigripes Siphon with 9 or fewer pecten spines... . Ls Ree eee fuscana, halifaxii, vorax DIFFERENCES BETWEEN OLD WORLD AND NEW WorLD Lu7Z/A PUPAE Intersegmental membranes.—The inter- segmental membranes of the abdominal ter- ga II/UI, and/or III/IV have a definite retic- ular pattern in about the medial 0.15 in Lt. fuscana, Lt. halifaxti, Lt. vorax and Lt. ti- gripes, While Lt. bigoti and Lt. allostigma pupae have no such pattern, although a pig- mented spot may be present medially on II/ Il. Paddle.—Paddles of the New World pu- pae are longer and wider that those of the pupae of the Old World species. The Lt. bigoti and Lt. allostigma paddles combined measure 1.63—1.98, x 1.78 mm long and 1.25—1.88, x 1.55 mm wide and they are not emarginate apically. Paddles of Lt. fus- cana, Lt. halifaxii, Lt. vorax and Lt. tigripes measure 1.19—1.56, x 1.36 mm long and 0.96—1.36, x 1.15 mm wide for the com- bined sizes of the three species. Data on Lt. fuscana and Lt. halifaxii given by Toma and Miyagi (1986) and Tanaka (2003) agree with these dimensions. Seta 1-IX.—This seta is present in the Old World species and absent in the New World taxa. Larvae.—The larvae of Old and New World species are also quite different. Lt. bigoti and Lt. allostigma larvae have a si- phon with an index of 3.0, a row of short, 6- to 14-branched, siphonal setae along the whole length while the larvae of Lt. fus- cana, Lt. halifaxii and Lt. tigripes have si- phon indices of 2.0 or less, a row of closely set, very long, single to triple setae, extend- ing along most of its length. It is quite obvious that the New World Lutzia immatures are morphologically very different from the Old World species, al- though the predatory behavior of the larvae remains the same. In the evolutionary pro- cess it is obvious that the isolation of the New World species has resulted in the de- velopment of significant morphological dif- ferences from the Old World species. ACKNOWLEDGMENTS I am indebted to Stephanie Clark-Gil Bloem for collecting and preparing the specimens of Lutzia bigoti; to the National Geographic Society, Dr. Greg Courtney and S.P. Pradhan for supporting and participat- ing in the collection of Nepal specimens and to T. Gaffigan for providing specimens from the collections of the National Muse- um of Natural History, Smithsonian Insti- tution. The illustration was prepared by B. Bower-Dennis and T. Litwak and the man- uscript reviewed by G.E O’Meara and J. Rey. This is Florida Agricultural Experi- ment Station Journal Series No. R-19543. LITERATURE CITED Baisas, E E. 1974. The mosquito fauna of Subic Bay Naval Reservation, Republic of the Philippines. U.S. Navy, First Medical Service Wing Technical Report No. 72-2, 170 pp. Belkin, J. N. 1962. The mosquitoes of the South Pa- cific (Diptera, Culicidae). Berkeley, University of California Press, Vol. I,, 608 pp; Vol Il, 412 pp. Bonne, C. and J. Bonne-Wepster. 1925. Mosquitoes of Surinam. Mededeeling Koloniaal. Instituut Am- stersdam, Afdeeling Tropische Hygiene Number 13, 558 pp. Bram, R. A. 1967. Contributions to the mosquito fauna 154 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of Southeast Asia—II. The genus Culex in Thai- land (Diptera, Culicidae). Contributions of the American Entomological Institute 2(1): 1-296. Clark-Gil, S. and R. E Darsie, Jr. 1983. The mosqui- toes of Guatemala, their identification, distribution and bionomics, with keys to adult females and larvae in English and Spanish. Mosquito System- atics 15: 151-281. Darsie, R. EF, Jr. 1998. Descriptions of the pupae of six species of Armigeres Theobald, subgenus Leices- teria Theobald (Diptera: Culicidae). Proceedings of the Entomological Society of Washington 100: 234-240. Darsie, R. FE Jr. and S. P. Pradhan. 1990. The mosqui- toes of Nepal, their identification, distribution and biology. Mosquito Systematics 22: 69-130. Delfinado, M. D. 1966. The culicine mosquitoes of the Philippines, Tribe Culicini (Diptera, Culicidae). Memoirs of the American Entomological Institute 7: 1-252. Edwards, KE W. 1932. Genera insectorum. Diptera. Family Culicidae. Fascicle 194, Belgium. 253 pp. . 1941. Mosquitoes of the Ethiopian Region III. Culicine Adults and Pupae. London, Brtitsh Mu- seum (Natural History), 499 pp. Heinemann, S. J. and J. N. Belkin. 1977. Collection records of the project Mosquitoes of Middle America” 7. Costa Rica (CR). Mosquito System- atics 9: 237-287. Heinemann, S. J. and J. N. Belkin. 1978. Collection records of the project Mosquitoes of Middle America”’ 10. Panama, including Canal Zone (PA, GG). Mosquito Systematics 10: 119-196. Hopkins, G. H. E. 1952. Mosquitoes of the Ethiopian Region I—Larval Bionomics of Mosquitoes and Taxonomy of Culicine Larvae. Second Edition. British Museum (Natural History), 355 pp. Knight, K. L. and R. W. Chamberlain. 1948. A new nomenclature for the chaetotaxy of the mosquito pupa, based on a comparative study of the genera (Diptera: Culicidae). Proceedings of the Helmin- thological Society of Washington 15: 1-18. Knight, K. L. and A. Stone. 1977. A Catalog of the Mosquitoes of the World (Diptera: Culicidae). 2™4 Edition, Thomas Say Foundation, Vol. VI, 611 pp. LaCasse, W. J. and S. Yamaguti. 1950. Mosquito fauna of Japan and Korea. U.S. Army, Office of Surgeon General, Headquarters, Eighth Army, 12 pp. Lane, J. 1953. Neotropical Culicidae. 2 Volumes, Sao Paulo, Brazil. 1112 pp. Penn, G. H. 1949. Pupae of the mosquitoes of New Guinea. Pacific Science 3: 3-85. Sennevet, G and E. Abonnenc. 1939. Les moustiques de la Guyane Frangaise.—II. Le genre Culex. Ar- chives of the Institute Pasteur Algerie 17: 62—134. Tanaka, K. 2003. Studies on the pupal mosquitoes of Japan (9) Genus Lutzia, with establishment of two new subgenera, Metalutzia and Insulalutzia (Dip- tera, Culicidae). Japanese Journal of Systematic Entomology 9: 159-169. Tanaka, K., K. Mizusawa and E. S. Saugstad. 1979. A revision of the adult and larval mosquitoes of Ja- pan (including the Ryukyu Archipelago and the Ogasawara Islands) and Korea (Diptera: Culici- dae). Contributions of the American Entomologi- cal Institute 16: 1-987. Theobald, E V. 1903. A monograph of Culicidae or Mosquitoes III, London, xv + 354 pp. Toma, T. and I. Miyagi. 1986. The mosquito fauna of the Ryukyu Archipelago with identification keys, pupal descriptions and notes on biology, medical importance and distribution. Mosquito Systemat- ics 18: 1-109. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 155-164 HAPLOCHELIDAE, A NEW FAMILY OF CRETACEOUS BEETLES (COLEOPTERA: MYXOPHAGA) FROM BURMESE AMBER ALEXANDER G. KIREJTSHUK AND GEORGE POINAR, JR. (AGK) Laboratory of Insect Systematics, Zoological Institute of Russian Academy of Sciences, St. Petersburg 199034, Russia (e-mail: AK3929@ AK3929.spb.edu); (GP) De- partment of Zoology, Oregon State University, Corvallis, OR 97331, U.S.A. (e-mail: poinarg @science.oregonstate.edu) Abstract.—A new family, genus and species of flightless beetles (Coleoptera: Lepi- ceroidea: Haplochelidae: Haplochelus: Haplochelus georissoides) are described from Cretaceous Burmese amber. Haplochelus georissoides is the first fossil that can be reliably placed in the suborder Myxophaga. The new family is characterized by its small size (under 2 mm in length), long frons (extended anteriorly far beyond eyes), ventrally dis- placed, declined and reduced mouthparts, 7-segmented antennae with a triangular terminal club bearing a dense layer of setae, long mesosternum, very short metasternum, fused elytra with no evidence of a suture, and 1-segmented tarsi with a single and long claw terminating all legs. The new species has similarities with members of the extant Lepicerus Motschulsky, 1855, although it is distinct enough to be placed in a separate family. Key Words: Haplochelidae, new family, Haplochelus, new genus Haplochelus georis- soides, new species, Lepiceroidea, Myxophaga, Burmese amber, Lower Cretaceous A small beetle with unique characters un- known in any extinct or extant family was discovered in Cretaceous Burmese amber. It is described here and its affinities within the Coleoptera are discussed. Amber from Bur- ma (Myanmar) occurs in lignitic seams in sandstone-limestone deposits in the Hu- kawng Valley. Palynomorphs obtained from the amber beds where the fossil piece orig- inated have been assigned to the Upper AI- bian (~ 110—100 mya) (Cruickshank and Ko 2003). Nuclear magnetic resonance (NMR) spectra of amber samples taken from the same locality as the fossil indicated an ar- aucarian (possibly Agathis) plant source (Lambert and Wu, personal communication). MATERIALS AND METHODS The amber was polished in order to better view the specimen. The amber piece con- taining the fossil is roughly rectangular in outline, measuring 14 mm long, 8 mm wide, and | mm in depth. The specimen is well preserved and complete, although some portions of the legs are obscured by the dark under portion of the body and por- tions of the abdomen appear to be some- what distorted. Observations, drawings, and photographs of the fossil were made with a Nikon SMZ-10 R stereoscopic microscope and a Nikon Optiphot TM compound mi- croscope (with magnifications up to 650%). All measurements are in mm unless other- wise noted. Lepiceroidea Hinton, 1936 This superfamily is proposed here to in- clude the Lepiceridae, including two extant species of the genus Lepicerus Motschul- 156 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sky, 1855 (Lepiceridae -Hinton, 1936 = Cyathoceridae Sharp, 1882) and Haplo- chelidae, n. fam. with one extinct species described below. Diagnosis.—This superfamily differs from the Sphaerioidea in the characters list- ed in the key below as well as in the fol- lowing features: (Sub) sinuated lateral con- tour of elytra, subhorizontal epipleura rath- er wide, with deep excision for reception of posterior legs, medial portion of anterior pronotal rim strongly projecting forward; and short intercoxal process on prothorax (not extended behind middle of procoxae). The lateral view of members of the Lepi- ceroidea is quite characteristic due to the peculiar shape of the head and the height- ened distal part of elytra, the apices of which are strongly curved downwards (be- yond the ventral plane of body). Haplochelidae Kirejtshuk and Poinar, new family Description.—Body small, under 2 mm in length, with rather convex dorsum; sub- flattened ventrally; dorsal integument with very coarse irregular sculpture, represented on elytra by somewhat regular tubercles and dense fossae, impunctate and scarcely pubescent; underside with coarse, rather dense fossae and very large punctures, lack- ing visible pubescence; frons anteriorly ori- ented, rather long (extended anteriorly far beyond eyes) and declined ventrally; mouthparts reduced, dislocated ventrally, concealed; gula comparatively narrow and deeply excavate; ocelli absent; antenna 7- segmented, with ultimate antennomere sub- triangular, widest at subtruncate apex; all coxae small, apparently oval and rather nar- rowly separated; thorax with very long me- sosternum, rather short metasternum; legs moderately raised, with distinct trochanter and spinose tibiae, tarsi 1-segmented, long and straight, with a large terminal, scarcely curved single claw; abdomen with 4 appar- ent ventrites; scutellum not exposed; elytra apparently completely fused, completely covering abdomen, with a deep longitudinal excavation along middle and 3 longitudinal excavations lateral to median one; epipleura deeply and elongately excavate, with deep excisions for intermediate and posterior legs. Type genus.—Haplochelus, n. gen. Etymology.—Derived from the Greek haplos for simple, single, and the Greek chele for claw, referring to the single clawed condition. Diagnosis.—The above-mentioned ima- ginal characters separate this family from all previously described extant and extinct beetles. In general form and many external structural characters, the fossil resembles members of the extant family Lepiceridae (Reichardt 1976), especially regarding the tarsomeres and single claw on each tarsus, the peculiar shape of the head, the charac- teristic excisions in the elytra for reception of the legs, and the elytral apices projecting far downwards (beyond the ventral plane of body). However the family differs from the lepicerids in the following characters: A long, narrow and very deeply excavate gula, completely fused elytra (without a trace of a suture) with faint longitudinal costae, 7-segmented antennae, coxae nar- rowly separated, and a very long mesoster- num and a very short metasternum. In ad- dition, the family Haplochelidae includes apterous species, while the known represen- tatives of Lepiceridae have moderately de- veloped posterior wings. The following key separates the new family Haplochelidae from the adults of other families in the Myxophaga. Ite Tarsi 1-segmented, terminating in one claw; mouthparts dislocated ventrally; ultimate antennomere much larger than preceding ones; pronotum widened anteriorly; dorsum coarsely sculptured and without distinct punctuation; superfamily Lepiceroidea .... 2 = Tarsi 3- or 4-segmented, terminating in two claws; mouthparts oriented apically; ulti- mate antennomere of various shapes, but never much larger than preceding ones; pronotum widest at base and more or less gradually narrowing anteriorly; dorsum var- VOLUME 108, NUMBER 1 iable in sculpture and punctation; superfam- liy SRO , M,, R,,;, R; and from R, to slightly anterior to R, (total of 7 pale fringe spots). Halter: Pedicel pale, scabellum and capitellum dark, capitellum dark-scaled. Legs: Coxae and trochanters without scales; femora, tib- iae and tarsi dark-scaled, apices of tibiae indistinctly pale, tarsomeres 1—4 with mi- nute faint dorsoapical pale spots. Abdomen: Integument dark with uniform covering of golden setae, without scales. Male.—Similar to female except for ob- vious sexual differences; other differences include the following. Head: Eyes more 201 widely separated, decumbent falcate scales slightly more numerous. Proboscis longer and more slender, about 1.9 mm, approxi- mately 1.4 length of forefemur. Maxillary palpus largely dark-scaled, with pale scal- ing as follows: narrow dorsolateral patch at apex of palpomere 3, large dorsolateral patch closer to anterior than to posterior margin of palpomere 4, and covering entire dorsolateral surface of palpomere 5. Wing: Generally paler and scaling reduced, fringe spots less distinct. Genitalia: Not dissected, see Harrison (1980: Fig. 16). Pupa (Figs. 1A, B).—Character and po- sitions of setae as figured; numbers of branches in Table |. Cephalothorax: Light- ly and more or less evenly pigmented, ap- pendages and metanotum with darker areas. Seta 7-CT long, about twice length of 6- CT, usually double, sometimes triple; 8-CT single; 1O-CT with 1—3(2) branches. Trum- pet: Moderately pigmented; length 0.24— 0.40 mm (mean = 0.37 mm); meatus very short, 0.04—0.10 mm (mean = 0.07 mm); pinna long, 0.25—0.33 mm (mean = 0.29 mm). Abdomen: Length 2.09—2.53 mm (mean = 2.29 mm); lightly pigmented, an- terior margins of more anterior segments darker; terga and sterna minutely spiculate. Seta O-III-VII long, usually with 2,3 branches (1—4), inserted anterior to seta 2 on segment III, far laterad of seta 2 on seg- ments IV—VII, more or less directly anterior to seta 5 on segments V—VII; seta 1-II—-VI with multiple thin flexible branches, 1-V,VI usually and 1-VII always single, long, ex- ceeding length of following tergum; 6-1, II long, more than twice length of seta 7, 6-I often double (single to 6-branched), 6-II single; 6-III—-V always and 6-VI,VII usually branched, number of branches generally progressively decrease from 6-III to 6-VII; seta 7-III—V short, normally branched, 7- VI,VII single, long, about length of follow- ing sternum, 7-III—VI inserted on lateral side of fold line before posterior margin of segment, 7-VII inserted on fold line at pos- terior margin of segment; setae 8,10,11-II usually absent, on one side when present, 202 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. A,B, Pupa of Anopheles minimus, modelled after exuviae of neotype female. A few missing setae were drawn from a paraneotype. A, Left side of cephalothorax, dorsal to right. B, Dorsal (left) and ventral (right) aspects of metathorax and abdomen. C, Map of southern Asia showing the type localities (see text for names) of An. minimus and its synonyms (1, minimus; 2, vincenti: 3, formoasensis I; 4, christophersi; 5, alboapicalis). CT = cephalothorax; Pa = paddle; T = trumpet; I-IX = abdominal segments I-IX; 0—14 = setal numbers for specified areas, e.g., seta 3-I. Scales in mm. VOLUME 108, NUMBER 1 203 8-II usually double (1,2), 10-II represented by alveolus in 3 specimens and double seta in One specimen, 1 1-II single when present; seta 9-I relatively long, about half length of = 6-I, with 3—6(4) branches; 9-II,III small, + Healt at lest: Male peglike; 9-IV—VII long, curved, simple and a sharply pointed, length progressively in- creasing from about 0.4 of segment IV to 0.6 of segment VII; 9-VIII slightly shorter lok = than 9-VII, about length of 4-VIII, plumose with 9—14(10) close-set branches arising from broad flattened central stem; 10-II ab- s sent. Genital lobe: Length 0.16—0.18 mm in female; 0.33—0.37 mm in male, with nip- a ple at apex. Paddle: Lightly pigmented (hyaline), buttress and midrib slightly dark- 5 er, midrib distinct to near seta 2-Pa; length ‘ES a ae eee Staite 0.61—0.70 mm (mean = 0.65 mm), width ie 0.40—0.46 mm (mean = 0.43 mm), index 1.4—1.65 (mean = 1.53); marginal serra- - tions begin 0.28—0.35 from base and end ae 0.47—0.54 from base where they are re- ~ placed abruptly by short hyaline filaments; refractile index 0.27—0.33 (mean = 0.30). = Seta 1-Pa long, sinuous, with hooked tip, (eee about one-third length of paddle, arising us from shallow emargination at apex. Larva, fourth-instar (Fig. 2).—Character a and positions of setae as figured; numbers VEVVyY LF of branches in Table 2. Head: Slightly wid- = er than long, width 0.57—0.66 mm (mean = _ 0.61 mm), length 0.54—0.60 mm (mean = 0.57 mm); integument with mottled pattern of moderately to darkly pigmented areas, la with some intervening lightly pigmented ar- eas; collar and mentum darkly pigmented. Setae 2,3-C single, simple; 3-C 0.5—0.6 length of 2-C; 4-C single, reaching just be- yond alveolus of 2-C, inserted in line with 5-C; 8-C usually dendritic, with 4—7(6) branches; 13-C inserted posterolateral to 11-C. Antenna: Moderately to darkly pig- mented; strong barb-like spicules on mesal surface, weaker ones on ventral surface; length 0.20—0.24 mm (mean = 0.21 mm). Seta 1-A short, single, simple, length about diameter of antenna, inserted about 0.3 from base on outer dorsolateral surface; 4- A with 4—7(5) branches. Thorax: Integu- Paddle Pp l 2-5 (3) IX Vill ] Vil 4 1 3-5 3 3-5 3 1—4 (2) 2 VI 2 3 3 7 1-3 Y~S~wYewTewr ~— 3-7 —- ( 2-6 (4) Abdominal Segments IV 6-8 (7) Dreon(2) 3-9 (7) I 1 (2) 12-21 I 16-32 (22) 4-8 (6) 4-8 (7) 3-8 (4) OQ-2 (0) 2 0, 1 (O) 5 = 1-6 (2) Range (mode, where apparent) of numbers of branches for pupal setae of the neotype series of Anopheles minimus. I n 5-5 (5) 3-5 (4) 2-5 (4) 2-6 (3) Cephalothorax (Ei nc = not counted; a = alveolus only. Table 1. Setae No 0 1 2 3 4 5 6 1 8 9 10 204 Fig. 2. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fa) Sr —<——s ~ a vA é : ZZ w ze a SS ————— oa = = <-> ay S — aS Wi 4 \ 0:5 Fourth-instar larva of Anopheles minimus, reconstructed from exuviae of neotype female. A few setae missing on the right side of the exuviae were drawn from the left side, and vice versa. A, Head, dorsal (left) and ventral (right) aspects of left side. B, Thorax and abdominal segments I-VI, dorsal (left) and ventral (right) aspects of left side. C, Abdominal segments VII-X, left side. D, Seta 1-IV. A = antenna; C = cranium; M mesothorax; MATP = median accessory tergal plate; NP = notal plate; P = prothorax; S = spiracular lobe; Sa = saddle; T = metathorax; TP = tergal plate; I-VIII,X = abdominal segments I-VIII,X; 0-15 = setal numbers for specified areas, e.g., seta 5-C. Scales in mm. VOLUME 108, NUMBER 1 Table 2. Range (mode, where apparent) of numbers of branches for fourth-instar larval setae of the neotype series of Anopheles minimus. Abdominal Segments IV 2,3(2) 17-23 Thorax Head Setae Vill VII 1,2(2) 15-19(16) 3- 5(3) VI 1,2(2) 15-21(17) 1,2(1) i 1,2(2) 16-22(21) II 1,2(1) 14 7-14(13) 2,3(2) 16-21(18) 11-16(13) 13-20(18) 4-7(5) 1,2(2) 25-37(28) 19-22(19) 1 1 2,3(3) 1 2- 4(2) 2- 5(4) 1 11-18(16) 1,2(1) 1,2(1) 3-6 1 1 9-13(11) 1 4-7(6) 5-8(6) 25-31(29) 2-5(3) 4-7(5) 3-5(4) 32-41(35) St = = = fon) ca 3 = 3-5(4) 29-39 1 3-6(4) 3,4(3) 31-34(34) 16-26(23) 27-39(34) 1 1 1 1 11-16(15) 27-42(28) 12-17(14) — Nm = — SSS Sw — — aa ee 7-97) 9-13(10) sO *Range of branches for individual setae (9 pairs). 205 ment hyaline, smooth. Mesothorax with conjoined pair of median notal plates, sometimes also with pair of submedian no- tal plates; metathorax usually with separat- ed pair of notal plates, apparently without submedian notal plates. Setae 1,2-P inserted on narrowly separated tubercles; support plate of pleural setal groups 9—12-PM,T with short spine; 9-P,T relatively sparsely plumose, 9-P with 9—13(10) branches, 9-T with 7—9(7) branches usually all arising from distal half of rachis; 10-P, 9,10-M and 10-T long, single, simple; 11-P large, sig- nificantly larger than 11-M,T, divided dis- tally into 2—5(3) branches. Seta 4-M with 3—6 branches arising at base; 12-M single or double, branched at midlength. Seta 3-T with short thick stem bearing 11—18(16) lanceolate leaflets; 12-T shorter than 12-M, with 3—5(3) branches from near base. Ab- domen: Integument hyaline, smooth; tergal plates of segments II—-VIII very large, about 0.6—0.7 width of segment, enclosing small median accessory tergal plate (not always enclosed on segment II), all of segments I-— VII usually but not always with distinct submedian accessory tergal plates. Seta O- II—VII well developed, normally branched, largest on segments IV and V, inserted close to posterolateral edge of tergal plate; 1-I- VII fully palmate with moderately pig- mented leaflets, leaflets with distinct shoul- ders and long slender filaments (blades and shoulders narrower on segment I), blades usually with distal patch of darker pigment near shoulder (Fig. 2D); 3-I-III,V,VI fairly long, single, 3-[V usually double or triple (2—4), 3-VII triple; 9-I,I] inserted antero- mesal to seta 7; 2-H, HI, VII branched, 2-IV— VI single (2-VI rarely double); seta 6-IV— VI branched at base, triple (2 of 16 seta 6- VI 4-branched). Pecten plate with 12—16 spines (usually 14 or 15) with basal denti- cles on dorsal side, long spines usually at each end with several interspersed among short spines. Seta 1-S large, with 7-11 branches (usually with 7, 8 or 9 branches); 2-S small, with 7—10(8) branches. Saddle moderately to darkly pigmented, length 206 0.20—0.24 mm (mean =*0.22 mm). Seta 1- X single, simple, inserted on saddle, much longer than saddle; 2-X with 19—22(19) branches, most basal branches on dorsal side of rachis, relatively straight, with fine tapering tips; 3-X with 8-13 (often 11 or 12) long, thick, slightly curved, apically hooked branches; 4-X (ventral brush) with 9 offset pairs of setae, longest branches on anterior side of main stems. Dorsal and ventral anal papillae equal in length, short, shorter than saddle. Molecular analysis —Among the nine mosquitoes available for study (five fe- imaless specimen mos” CH6—Ik +355; 7, and —13; four males, specimen nos. CH6— 8, —9, —10, and —12), DNA from individ- ual —5 consistently failed to amplify by PCR, indicating poor DNA extraction. Of the remaining eight specimens, six yielded good sequence traces for the D3 region, all of which matched An. minimus species A at all of the five nucleotide positions that distinguish it from An. minimus species C (Sharpe et al. 2000, Chen et al. 2002, 2003). Sequencing of the COI fragment proved more difficult and only three sequences were obtained, of which only one was of good quality. Sequence from this individu- al, CH6—3, matched An. minimus A at the four sites with fixed differences between species A and species C in Thailand, in the region of 294 bp sequenced (Sharpe et al. 2000). These four bases all match An. min- imus species A sequences obtained in China (Chen et al. 2002, 2003). Based on the availability of COII and D3 sequences, specimen number CH6—3 was selected as the neotype for An. minimus (see above and below). The sequences obtained for this specimen are deposited in GenBank under accession numbers AMO039906 (D3) and AM039907 (COID). Systematics.—Anopheles minimus is the nominotypical member of a sibling species complex comprised of three or possibly four genetic species (see introduction) that are virtually isomorphic; hence, unambig- uous diagnosis of this species is only PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON achievable by means of genetic and molec- ular markers (see above). Having said this, An. minimus is partially, albeit dubiously, distinguished from species C of the com- plex by the absence of a humeral pale spot (HP) on the wings. Green et al. (1990) found that this spot was absent in 95% of An. minimus females (as species A) but was missing in only 22% of species C females from Kanchanaburi Province in western Thailand. Similarly, Sharpe (1997) recorded the absence of a HP spot in 91% of An. minimus (as species A) but only 37% of species C collected at the same locality (Ban Phu Rat) visited by Green et al. (1990). Chen et al. (2002) noted the ab- sence of this spot in a comparable percent- age of An. minimus females (92.7%, as spe- cies A) from southern China, but it was missing in a significantly greater number of species C females (84.4%). Finally, Van Bortel et al. (1999) observed an even higher degree of similarity between the two spe- cies in northern Vietnam where 99% of An. minimus (as form I) and 91.8% of species C (as form II) lacked HP spots. From these observations, it is obvious that the presence or absence of HP spots cannot be used to identify or distinguish the two species with any degree of confidence. Van Bortel et al. (1999) and Chen et al. (2002) also recorded the presence/absence of a presector pale spot on the wings of males and females and showed that this character is even less re- liable for distinguishing the two species. Anopheles minimus is very similar to three other species of the Myzomyia Series that occur within its range of distribution in the Oriental Region, i.e. An. aconitus, An. fluviatilis, and An. varuna. As pointed out by Harrison (1980), no morphological char- acters are completely reliable for distin- guishing the adults of these species. Fur- thermore, the adults of An. pampanai are also often misidentified as An. minimus be- cause the distinguishing features of the wings are not easily discerned. Consequent- ly, adults of An. minimus (as well as those of other members of the Minimus Com- VOLUME 108, NUMBER 1 plex) cannot be distinguished from _ the adults of these species with certainty with- out associated larval and pupal exuviae. Distinguishing morphological features are provided in the identification keys of Har- rison (1980). However, because of the un- certainties associated with morphological differentiation, the various types of molec- ular assays developed by Sharpe et al. (1999), Van Bortel et al. (2000), Kengne et al (2001), Phuc et al. (2003), and Garros et al. (2004) should be used for the unequiv- ocal identification of An. minimus (= their An. minimus species A). The names of four nominal species (one with an unjustified replacement name, see synonymy) are currently regarded as junior synonyms of An. minimus, but only one of these, formosaensis I Tsuzuki, is certain to denote the same biological species. Tsuzuki (1902) described An. formosaensis I from adult mosquitoes collected at an undis- closed location on the island of Taiwan (lo- cality 3 in Fig. 1C). Evidence shows that species C of the Minimus Complex does not extend into eastern areas of China, and only An. minimus occurs in Taiwan (Chen et al. 2002). The syntype specimens (adults) of An. vincenti were collected at Van Linh in the former French protectorate of Tonkin (Lav- eran 1901), which in 1946 formed the northern part of Vietnam bordering on Chi- na (locality 2 in Fig. 1C). Electrophoretic studies of the octanol dehydrogenase (Odh) enzyme locus indicate that An. minimus and species C both occur at this locality (Nguy- en Duc Manh, personal communication). Whether some or all of the syntypes of An. vincenti are conspecific with either An. min- imus Or species C is unanswerable. As in- dicated above, adults of these species are indistinguishable in northern Vietnam, and molecular methods are unlikely to be useful in resolving the identity of the syntypes be- cause they are mounted in balsam on a sin- gle microscope slide (Harrison 1980). For these reasons, vincenti should remain in 207 synonymy with An. minimus until the ap- plication of this name is resolved. Anopheles christophersi and variety al- boapicalis were described by Theobald (1902 and 1910, respectively) from speci- mens collected in localities of northeastern India that reside in present-day West Bengal (localities 4 and 5, respectively, in Fig. 1C). Before Garros et al. (2005b) and Chen et al. (2006) showed that An. fluviatilis species S was conspecific with An. minimus species C, only An. minimus (as species A) was known to occur in India. Species C is now known to have a wide distribution in the northern states of Madhya Pradesh, Orissa, and Rajasthan. Although this species (as ei- ther minimus C or fluviatilis S) has not been reported from the more easterly state of West Bengal, its possible occurrence in this geographic area cannot be ruled out. The location of the type of An. fluviatilis is un- known (Knight and Stone 1977, Harrison 1980), and it probably does not exist. James (1902) apparently described this species from several places in India, including lo- calities in West Bengal where type speci- mens of An. christophersi and variety al- boapicalis originated. Hence, it is possible that one or more of the names fluviatilis, christophersi, or alboapicalis may apply to either An. minimus or species C. Fixing the identity of An. minimus by neotype designation does not resolve the taxonomic identity of species C, i.e., wheth- er it should be denoted by an available name or recognized as a new species. In contrast, there is clearly no available name for species E of the complex from Ishigaki Island, Japan, and it should be formally de- scribed and named as a species new to sci- ence: Material examined.—Nine individually reared specimens (5 females, 4 males), each with associated larval and pupal exuviae on microscope slides. Neotype, female (CH6— 3), CHINA: New Territories, Hong Kong, Ham Hang Mei (22° 13’ N, 114° 13’ E), 11 April 2002, stream margin in full sun with dead leaves (L. Rueda & J. Pecor). Other 208 specimens, same data as neotype: 4 females (CH6-1, CH6—-5, CH6-7, CH6-13); 4 males (CH6—8, CH6—9, CH6—10, CH6—12). The specimens are deposited in the USNM. Sequences for DNA extracted from a hind- leg taken from the neotype are deposited in GenBank: accession no. AM039906 for D3, AM039907 for COIL. ACKNOWLEDGMENTS We are indebted to Leopoldo Rueda and James Pecor (Walter Reed Biosystematics Unit, Smithsonian Institution, Washington, DC) for collecting, rearing, and loaning the specimens to us that made this study pos- sible. Bruce A. Harrison (Public Health Pest Management, North Carolina Depart- ment of Environment and Natural Resourc- es, Winston-Salem, North Carolina) is gratefully acknowledged for reviewing the manuscript and providing constructive comments. We are equally grateful to The- resa M. Howard (The Natural History Mu- seum, London) for preparing the illustra- tions. LITERATURE CITED Baba, K. 1950. A study of anopheline mosquitoes in relation to the epidemiology of malaria in Canton Delta, South China. The Transactions of the Kan- sai Entomological Society 15: 1-11. Baimai, V. 1989. 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Insect Molecular Biology 9: 335-340. Yamada, S. 1925. A review of the adult anopheline mosquitoes of Japan: systematic descriptions, their habits, and their relations to human diseases (Part II.). Scientific Reports to the Government Institute of Infections Disease (Tokyo) 4: 447—493. Yu, Y. 1987. Studies on the two forms of Anopheles (Cellia) minimus Theobald, 1901 in China (Dip- tera: Culicidae). Mosquito Systematics 19: 143— 145. Yu, Y. and M. Li. 1984. Notes on the two forms of Anopheles (Cellia) minimus Theobald, 1901 in Hainan Island. Journal of Parasitology and Para- sitic Diseases 2: 95—98 (in Chinese). Zhou, S., L. Tang, and X. Zheng. 2002. Molecular var- iation based on mtDNA-COII gene of members of the Anopheles minimus group. Chinese Journal of Parasitology and Parasitic Diseases 20: 168—170 (in Chinese). PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 210-225 A QUANTITATIVE EVALUATION OF INTRASPECIFIC AND INTERSPECIFIC VARIATION IN EYE MORPHOLOGY FOR EFFERIA COQUILLETT (DIPTERA: ASILIDAE) SPECIES K. C. HOLSTON AND C. R. NELSON (KCH) Swedish Museum of Natural History, Section for Entomology, P.O. Box 50007, S-104 O05 Stockholm, Sweden (e-mail: kevin.holston@nrm.se); (CRN) Department of In- tegrative Biology, WIDB 401, Brigham Young University, Provo, UT 84602, U.S.A. (e- mail: rileynelson@byu.edu). Abstract.—Qualitative taxonomic characters based on differences in linear eye mea- surements were converted to qualitative scores for Efferia group species, and statistical analyses of intraspecific and interspecific variation were perfomed. Sexual dimorphism noted for E. albibarbis is the first documented for an eye morphology character in Asi- lidae, and may be analogous to sexual dimorphism described in a holoptic-dichoptic con- text for other species of Diptera. Interspecific variation in E. albibarbis Macquart and E. aestuans (Linnaeus) demonstrates that robust comparisons among species cannot rely on single exemplars, and differences among specimens based on sex and geography should be considered during subsequent eye morphology studies in Asilidae. The variables iden- tified in this study demonstrate that a wide variation in eye morphology exists among Efferia group species that can be examined in a quantitative context. Detailed accounts of intraspecific variation are considered crucial for comparisons among species of Asili- dae, and interspecific comparisons of eye morphology are likely to demonstrate biologi- cally significant results in studies of monophyletic taxa and ecological guilds. Key Words: intraspecific variation, eye morphology, robber fly, Efferia albibarbis, Ef- feria aestuans Compound eyes of robber flies (Diptera: Asiloidea: Asilidae) are sensory organs of great importance for these active visual predators, and studies of eye morphology are relevant to further taxonomic, ecologi- cal, and phylogenetic assessments of Asi- lidae. Robber flies hunt primarily from ex- posed perches on vegetation or the ground, where they often move their heads and re- orient their bodies in response to move- ments of potential prey. The subsequent at- tack flight has been described as “‘target- directed”” or ‘“‘ballistic interception’’ (Buschbeck and Strausfeld 1996, 1997), and differences among species have been noted concerning hovering ability, habitat preferences, and prey handling. The com- pound eyes of robber flies are large and prominent with respect to dorsal and ventral margins of the head in anterior view, such that the vertex appears “‘excavated”’ (Wood 1981, Lehr 1988, Majer 1997). Eyes in both sexes are well-separated at the vertex (Wood 1981), which has been considered an evolutionarily derived characteristic for Asilidae (Yeates 1994), although holoptic males (i.e., the margins of the compound eyes meet dorsally at vertex) and dichoptic females (i.e., compound eyes separated) fre- quently characterize species of Diptera VOLUME 108, NUMBER 1 (McAlpine 1981). The predatory behavior of robber flies may promote similar eye morphology between males and females, but sexual dimorphism has been noted for other body parts directly involved with pre- dation, such as wings and legs (Hull 1962, Lavigne 2002). Intraspecific differences in eye morphol- ogy, however, have not been documented, and interspecific differences are generally summarized as qualitative taxonomic char- acters to differentiate among genera. The degree to which the compound eyes con- verge toward the vertex is often used in tax- onomic characterizations of Asilidae, with the vertex width relative to face width used to distinguish genera having compound eyes that are widely divergent towards the vertex (e.g., Stichopogon, Lasiopogon, Townsendia, Willistonina) (Wood 1981). Quantitative variables for eye morphology would facilitate precise comparisons among specimens and species and allow eye mor- phology to be incorporated in biological in- vestigations of robber fly species. The wide geographical distribution and abundance of Efferia albibarbis (Mac- quart), and its distinctive eye morphology makes this species an appropriate choice for describing intraspecific variation in Efferia. Fisher and Hespenheide (1992) considered E. albibarbis the most common robber fly in North and Central America, and this spe- cies 1s often well represented in collections of North American Asilidae. Wilcox (1966) noted in his key to the “‘Albibarbis Group” that E. albibarbis has a “‘frons at vertex as wide as at antennae” in contrast to the nar- rower vertex in other Albibarbis Group spe- cies. Wilcox (1966) also reported eye, face, and vertex widths for 81 Nearctic Efferia species, but the utility of these measure- ments in taxonomic or biological studies has not been evaluated. Efferia aestuans (Linnaeus) is a North American species that is also well represented in collections, and a comparison of eye morphology among Ff- feria species with emphasis on intraspecific variation in E. albibarbis and E. aestuans 2a would provide an informative reference for further studies. Face, eye, and vertex widths have been reported in descriptions of robber fly spe- cies and can be expressed as ratios that de- scribe the relative proportions of these mea- surements. Qualitative taxonomic diagnos- tic characters used in keys, such as “‘face wide, about width of eye” or “‘frons greatly expanded toward vertex’’ are based on the relative widths of these three features (Wood 1981). The distance between the compound eyes across the face (‘‘face width’’) and across the vertex (‘vertex width’’) and the width of the eye are mea- sured from endpoints on the eye margin; these measurements describe, therefore, three major aspects of the anterior eye mor- phology. Ratios derived from these linear measurements yield values corresponding directly to these taxonomic characters that eliminate the effect of specimen size from comparisons. For example, if face and eye width are equal, then the 1:1 ratio can be expressed as the quotient of face width/eye width, or 1.0; a 1:2 ratio for a specimen with eye width twice that of the face yields a value of 0.5. In this study, interspecific and intraspe- cific differences in relative face, eye, and vertex widths for Nearctic Efferia species were examined to evaluate the utility of eye measurements in a quantitative context. MATERIALS AND METHODS Nomenclature.—Although the Nearctic Efferia species featured in this study have been divided recently among seven genera, these genera represent groups of Efferia species considered artificial groups by pre- vious authors (Hine 1919, Wilcox 1966, Martin and Papavero 1970, Fisher and Hes- penheide 1992, Artigas and Papavero 1997). Name combinations for species cor- respond to the genus level classification of Asilidae by Fisher and Hespenheide (1992) such that all species names are presented in combination with Efferia except Triorla in- terrupta (Macquart). The genus names re- i) in) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON euifse Ue removed), showing the three distances measured (a = eye). cently proposed for Efferia species groups or removed from synonymy with Efferia by Artigas and Papavero (1997) are not used. The term “Efferia group species” is used, therefore, in reference to the Nearctic Ef- feria species plus T. interrupta. Names and circumscriptions for species groups in Ef- feria follow Wilcox (1966) except that 7. interrupta is considered separately from Ef- feria and Staminea group species. Eye morphology measurements.—Mea- surements for the left compound eye, face, and vertex widths were recorded for 103 specimens of E. albibarbis and 48 speci- mens of E. aestuans housed at the Illinois Diagram of a robber fly head (E. albibarbis, anterior view with antennae and upper setae of head width of vertex, b = width of face, c = width of left Natural History Survey (INHS) (Fig. 1). Localities from which specimens were col- lected were grouped by geographic area: 1) Midwestern United States (MWUSA), 2) Baja California Norte, Mexico (BAJA), 3) Central America (CAMER), and 4) south- western United States (SWUSA), (Table 1). Measurements of the head in anterior view were recorded using an ocular micrometer at 50X and are reported as ocular micro- meter units (om). Wilcox (1966) reported measurements of the eye, face, and vertex for 81 Nearctic Efferia group species in ocular micrometer units at a magnification of “about 30 VOLUME 108, NUMBER 1 Table 1. Localities from which specimens of E. aestuans and E. albibarbis were examined, grouped by the geographic region noted for comparisons. Legend: MWUSA = midwestern USA, SEUSA = southeastern USA, SWUSA = southwestern USA, CAMER = Cen- tral America, BAJA = Mexico, Baja California Norte. MWUSA Indiana (Clinton), Illinois (Adams, Bond, Brown, Champaign, Dubois, Eddyville, Edgar, Ef- fingham, Hancock, Iroquois, Jackson, Marion, Mc- Henry, McLean, Pope, Pulaski, Richland, Scott, St. Clair, Wabash), Michigan (Berriens). SWUSA Arizona (Mohave, Pima, Santa Cruz), New Mexico (McKinney, Otero). CAMER Guatamala (Esquintla, Quiche), El Salva- dor (Cajunte), Costa Rica (Guanacaste, Puntaren- as). BAJA Baja California Norte. times.’’ Measurements were recorded for a single male specimen of most species; a fe- male was measured for E. femorata (Mac- quart) and E. clementi (Wilcox and Martin). Eye morphology variables.—We have used four variables to compare differences in eye morphology among specimens, using ratios derived from linear measurements to control for differences in specimen size. A straightforward quantitative description of eye convergence toward the vertex is the face width across the antennal bases divid- ed by the vertex width (face/vertex). An ad- ditional description of eye convergence is obtained by subtracting the face width di- vided by eye width (face/eye) from the ver- tex width divided by eye width (vertex/eye). This “‘Convergence-Divergence Index” (CDI) is distinguished from face/vertex by its use of two variables that incorporate eye width (CDI = face/eye—vertex/eye). Scores showing low or no variability within and strong differences among groups indicate variables that are most appropriate for tax- onomy. Proportional change in eye morpholo- gy.—The relationship between face/vertex and CDI scores allows correlation plots to depict proportional changes in eye width among specimens examined, which allows the variables describing eye convergence to 218 be evaluated based on predicted patterns. Differences in eye width determine the range of variation in CDI scores and the placement and spread of observations in correlation plots. Pattern 1: If face, vertex, and eye width measurements are constant or proportional among specimens, then face/vertex and CDI scores will be identical and all observations share a single point in the correlation plot. Pattern 2: If only eye width varies and face and vertex widths are constant or pro- portional among specimens, then face/ver- tex scores will be identical and the distri- bution of observations is linear and perpen- dicular to the x-axis. Pattern 3: If eye width is constant and face and vertex widths increase arithmeti- cally (e.g., face and vertex increase by 2 omu), then CDI scores will be identical and the distribution of observations is linear and perpendicular to the y-axis. This pattern also characterizes groups with identical dif- ferences in face/eye and vertex/eye scores. Pattern 4: If vertex and eye widths are constant or proportional and only face width varies, then the distribution can be described by the following linear equation: y = (vertex/eye)x—(vertex/eye). The slope of this line is m = vertex/eye, the y-inter- cept is b = —vertex/eye, and the x-intercept is at (1,0). “Pattern 4” also characterizes groups of specimens having arithmetic in- creases in face, vertex, and eye widths. Dis- tributions become more curvilinear as eye measurements change in larger increments, and the closest fits are expected for small incremental increases. An additional characteristic of face/ver- tex and CDI correlation should be noted. The point indicating equal face and vertex widths is (1, 0), where the face/vertex score equals 1 and the CDI equals 0. As the widths of the face and vertex become in- creasingly similar (e.g., face/vertex —> 1), differences in CDI scores for an identical range of eye widths are smaller due to clos- er face/eye and vertex/eye scores near the x- intercept. The fit of observations in a face/ 214 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON vertex and CDI correlation plot to the linear equations described above would be closer near the x-intercept even when overall var- iation in eye width does not change. Statistical descriptions and analyses.— Box plots were constructed to summarize descriptive statistics for Efferia group spe- cies, with inner fences for the data extend- ing | step beyond the interquartile range. Correlation plots were developed to de- scribe the relationship between the face/ver- tex ratio and CDI, with linear regressions calculated in Microsoft® Excel 2000, ver- sion 9.0. Pairwise comparisons between groups of specimens measured during this study (separated by species, geographic re- gion, and sex) were performed using un- paired two-tailed ¢-tests. These comparisons test for significant differences in eye mor- phology scores between EF. albibarbis and E. aestuans and between conspecific males and females. RESULTS Intraspecific variation.—In E. aestuans, there is a higher correlation between face and vertex widths in males (y = 0.6901x + 8.6234, R? = 0.8902) than females (y = 0.6956x + 8.0324, R? = 0.8879), a lower correlation in face and eye for males (y = 1.4152x + 11.518, R*? = 0.6644) than fe- males (y= 01.4298x. 13.858,> R2 = 0.8420), and a lower correlation in vertex and eye widths in males (y = 0.3046x + 26.632, R? = 0.5230) than females (y = 0.4063x + 19.776, R? = 0.7354). In E. al- bibarbis, there is a lower correlation be- tween face and vertex widths in males (y = 0.9942x + 1.4494, R? = 0.6968) than fe- males (y = 0.8615x + 7.6243, R? = 0.9048), a higher correlation in face and eye formmaless(y°— 1037x511 9346 SRA — 0.7584) than females (y = 0.7326x + 17.986, R? = 0.4741), and a higher corre- lation in vertex and eye widths in males (y =O 1x 14.957, RJ —10:5824) ethan females (y = 701541797x04; 925.768; -R20— OS9Siy). Intraspecific variation in E. aestuans and E. albibarbis indicate moderate levels of variation in the variables studied, with no overlap in face/vertex and CDI scores (Ta- ble 2). The face/vertex and CDI scores are higher for E. aestuans than for E. albibar- bis, showing that the compound eyes in E. aestuans (face/vertex: 1.1020 to 1.2745, mean = 1.1910 + 0.0408; CDI: 0.0806 to 0.1872, mean = 0.1326 + 0.0241) con- verge more strongly towards the vertex than for E. albibarbis (face/vertex: 0.8841 to 1.0526, mean = 0.9745 + 0.0434; CDI: =(01228 to’ 0:0476, mean’ = —0.02427"= 0.0413). The face/eye and vertex/eye scores show that eye width is generally greater than face or vertex widths in both E. aes- tuans (0.7037 to 0.9643, mean = 0.8345 + 0.0531; vertex/eye: 0.5977 to 0.8750, mean = 0.7019 = 0.0576): and E. albibarbis (face/eye: 0.7746 to 1.0678, mean = 0.901 1 + 0.0642; vertex/eye: 0.8060 to 1.0702, mean = 0.9225 + 0.0620) and that E. al- bibarbis has more similar face, vertex and eye widths. The linear relationship between face/eye and CDI scores indicates a trend toward proportional change in eye measurements for E. albibarbis and E. aestuans following the “Pattern #4” distribution, but variation in scores shows that increases are arithmetic (Fig. 2). The distribution for all E. albibar- bis specimens can be described by a linear regression (y = 0.9548x — 0.9547, R? = 0.9941) with a slope similar to the observed mean (0.9547 + 0.0513) and median (0.9547) vertex/eye scores. Similarly, the distribution for E. aestuans can be de- scribed by a linear regression (y = 0.6987x — 0.7, R* = 0.8225) close to the observed mean (0.7019 + 0.0576) and median (0.6992) vertex/eye scores. The correlation plot for face/eye and CDI shows lower fit to a linear regression for E. aestuans than for E. albibarbis although higher standard deviations are observed for E. albibarbis, which is predicted from the calculation of the CDI for distributions closer to the x- intercept (see ““Materials and Methods’’). 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Correlation plot for face/eye and CDI scores, showing separation of distributions and “‘Pattern 4” linear trends for E. albibarbis (triangles—lower scores) and E. aestuans (diamonds—higher scores). icant differences between E. albibarbis and E. aestuans (P < 0.001) and significant in- traspecific differences for E. albibarbis (Ta- ble 3). Differences between males and fe- males were not significant for E. aestuans (P > 0.413) but significant between males and females of E. albibarbis in face/vertex scores (P = 0.014) and nearly significant in face/vertex scores (P = 0.062). Males and females from MWUSA constitute the larg- est geographic group of E. albibarbis spec- imens, and have a higher significant differ- ence in face/vertex (P = 0.004) and are sig- nificantly different in face/eye (P = 0.008) and CDI scores (P = 0.003). Significant differences between males and females in face/vertex scores were also detected for BAIA (2 — .0:041)eand CAMER (2 — 0.043) groups and in CDI scores for the CAMER group (P = 0.040). Summary sta- tistics and correlation plots of face/eye and CDI scores show that scores for MWUSA E. albibarbis males extend farther into the realm of divergent eye scores (i.e., lower face/vertex and CDI scores for males) whereas there is a complete overlap of scores for males and females of E. aestuans (Fig. 3, Table 3). More females than males have face/vertex scores greater than 1.0 (Nynrates = 14, Ngemates = 18), more males have face/vertex scores between 1.0 and 0.965 (Males = 11, Memales = 3), and only males have scores below 0.965 (jae; = 35 Deemates = (0). Interspecific variation.—The four eye morphology variables identify outliers rath- er than discrete groups of Efferia species, and all but two Efferia group species (7. interrupta and E. albibarbis) have com- pound eyes that are closer at the vertex than at the frons (i.e., face width > vertex width) (Fig. 4, Table 4). The face/vertex ratio and CDI indicate divergence of the eyes at the vertex for 7. interrupta, neither divergence nor convergence for E. albibarbis, and con- vergence for all other Efferia group species. The lowest face/vertex scores are recorded for T. interrupta (0.980) and E. albibarbis VOLUME 108, NUMBER 1 PLING CDI face/vertex CDI 1,20 face/vertex Fig. 3. Correlation plots of face/eye and CDI scores for MWUSA E. albibarbis (upper plot, triangles) and E. aestuans (lower plot, diamonds) comparing distributions of males (solid symbols) and females (open symbols), with lower scores for males than females in E. albibarbis. (1.000); and the highest scores are observed for E. leucocoma (Williston) (1.500), E. zonata (Hine) (1.500), and FE. peralta Wil- cox (1.600), with E. peralta as an upper outlier among Efferia group species (Fig. 4). The lowest CDI scores were also ob- tained for 7. interrupta (-0.0189) and E. al- bibarbis (0.000), but E. leucocoma has the highest CDI score (0.308) with 7. interrup- ta, E. albibarbis, and E. leucocoma as out- liers for the CDI (Fig. 4). The face/vertex and CDI scores for E. albibarbis and E. aestuans include scores calculated from Wilcox (1966) measurements. There is, however, broad overlap in distributions of scores for the eight Efferia species groups in all eye morphology variables (Table 5), and the ranges of face/vertex and CDI scores for E. albibarbis and E. aestuans en- compass scores for 7. interrupta and 37 other Efferia species (Fig. 5). The correlation plot for Efferia group species shows a linear relationship for face/ vertex and CDI scores, and the slope of the 218 Table 3. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Results (p-values) from unpaired two-tailed t-tests of face/eye, vertex/eye, face/vertex, and CDI scores; with comparisons between E. aestuans and E. albibarbis, between all males and females of E. aestuans and E. albibarbis, and between males and females of E. albibarbis grouped by geographic region. Significant differences between groups (~ = 0.05) are indicated by asterisks (*). Face/Eye Vertex/Eye Face/Vertex CDI E. aestuans vy. E. albibarbis <0.001* <0.001* <0.001* <0.001* E. aestuans v. E. albibarbis, 3 3 <(0.001* <0.001* <0.001* <0.001* E. aestuans vy. E. albibarbis, 2 & <0.001* <0.001* <0.001* <0.001* E. aestuans, 66 v. E. aestuans, 2 & 0.835 0.882 0.576 0.413 E. albibarbis, 33 vy. E. albibarbis, 2 & 0.062 0.741 0.014* 0.203 MWUSA, 66 v. MWUSA, @ 2 0.008* 0.695 0.004* 0.003* BAJA, 66 v. BAJA, 22 0.017* 0.522 0.041* 0.158 CAMER, 66 v. CAMER, @ 2 0.695 0.130 0.043* 0.040* SWUSA, 36 v. SWUSA, 22 0.687 0.351 0.072 0.075 regression line through the x-intercept with best fit to: the data (Gy — 0:6227x — 0:625, R2 = 0.6996) has a value lower than me- dian (0.6739) and mean vertex/eye scores (0.6651 + 0.1209) (Fig. 6). Efferia species with the best fit to this linear regression have vertex/eye scores that are closest to Ol6227 (EZ. adeci: 0.6000, E. wilcoxi: 0.6207, E. latrunculata: 0.600, E. inflata: 0.6000, E. pilosa: 0.6250, E. kansensis: 0.6000, and E. leucocoma: 0.6154) (Table 4). As seen in the regressions for E. aes- tuans and E. albibarbis, the correlation plot shows greater spread of observations with increasing distance from the x-intercept, and vertex/eye scores indicate a lack of pro- portionality of vertex and eye widths among Efferia group species. DISCUSSION Intraspecific variation and interspecific distinctions.—All four variables can be used to distinguish specimens of E. albi- barbis from E. aestuans. The significant H = - — a = |e a 1 0,95 1,05 4113 11-25 “es 1,45 1,55 1,65 = a = lon a T <= SS ia ea = — = = T r 1 -0,05 oO 0,05 0,1 0,15 0,2 0,25 0,3 0,35 Fig. 4. Boxplots summarizing distributions of face/vertex (upper plot) and CDI (lower plot) scores, as listed for all species in Table 4. Symbols: face/vertex—T. interrupta, 0.980 (first square); E. albibarbis, 1.000, (tri- angle), E. cana, 1.070 (second square); E. aestuans, 1.143 (diamond); E. leucocoma and E. zonata, 1.500 (third square); and EF. peralta, 1.600 (upper outlier, fourth square). Symbols: CDI—T. interrupta, —0.0189 (lower outlier, first square); E. albibarbis, 0.000 (lower outlier, triangle); E. cana, 0.062 (second square); E. aestuans (0.116, diamond); E. peralta, 0.267 (third square); and E. leucocoma, 0.308 (upper outlier, fourth square). VOLUME 108, NUMBER 1 differences in eye morphology emphasize that FE. albibarbis and E. aestuans are dis- tinct species although the taxonomic status of these species is clearly established using a suite of traditionally used morphological characters (Wilcox 1966). Similarly, E. al- bibarbis and other species that are outliers in distributions of eye morphology scores can be diagnosed without reference to the variables defined in this study. These vari- ables, however, are more informative de- scriptors of eye morphology than summa- ries of linear measurements (e.g., the aver- age eye width for specimens examined), which lose diagnostic properties that are in- dependent of specimen size. Further docu- mentation of eye measurement ratios can improve the scope of current species diag- noses, but ratios should be selected based on their relevance to further taxonomic and biological investigations. Intraspecific sexual differences.—The lower minimum, median, and mean face/ vertex, CDI, and face/eye scores in male specimens of E. albibarbis indicate differ- ences in eye morphology between males and females. Significant sexual differences were detected in face/vertex, CDI, and face/ eye scores, indicating that males have either a narrower face or broader vertex and eye in comparison to females. The correlation between face and vertex widths for males of E. albibarbis indicates that face and ver- tex widths do not increase arithmetically as face and vertex widths for E. albibarbis fe- males or for either sex of E. aestuans. Mod- erate variation in vertex/eye scores was ob- served for E. albibarbis and E. aestuans, and there is no significant difference in means of vertex/eye scores between males and females MWUSA specimens (P = 0.695) or in comparisons of all E. albibar- bis specimens (P = 0.741) (Table 3). Males and females of E. aestuans, in contrast, are not significantly different in scores for any of the four eye morphology variables (P = 0.413) despite a lack of proportional change in eye measurements (Table 3). Lower face/vertex, CDI, and face/eye 219 scores are due to larger vertex and eye widths in males of E. albibarbis rather than narrower face widths. Face width and eye widths are related by a shared endpoint on the eye margin (Fig. 1), and a narrower face width that is not complemented by a larger eye width would result in sexual differences in ratios of overall head width (face width + eye width) to vertex width as in face/ vertex scores. To investigate this possibility, two tailed t-tests for (facet+eye)/vertex scores were performed. A lack of signifi- cant difference in (face+eye)/vertex scores between males and females from MWUSA (P = 0.334) or all E. albibarbis specimens (P = 0.186) shows that lower scores for eye divergence variables in males correspond to increases in both eye and vertex widths. An arithmetic increase in both vertex and eye width explains the lack of significant dif- ference between males and females in ver- tex/eye scores. Furthermore, a positive cor- relation between face and eye widths in E. albibarbis demonstrates that eye width in- creases with increasing face widths, and males have a stronger positive correlation in vertex and eye widths than females. The correlations between face/vertex and CDI scores also indicate that face, eye, and ver- tex measurements increase arithmetically: the fit to a linear regression is not due to proportional changes in vertex/eye scores. Sexual differences in eye morphology have not been documented in previous stud- ies of Asilidae and suggest that directional selection in E. albibarbis may have shifted face/vertex, CDI, and face/eye scores to lower means in males. The distribution of face/vertex and CDI scores for fly species with holoptic males would be bimodal and non-overlapping between males and _ fe- males. Eye morphology in robber fly spe- cies 1s considered identical between males and females, and the predicted distribution of face/vertex and CDI scores would be un- imodal, with complete overlap and identical means for male and female distributions. In E. albibarbis, however, males have lower median and mean scores than females, with 220 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 4. Scores for face/eye (f/eye), vertex/eye (v/eye), face/vertex (f/v), and CDI calculated from measure- ments reported in Wilcox (1966) for North American Efferia species (and *Triorla interrupta). Species f/eye v/eye tv CDI aestuans 0.9302 0.8140 1.1429 0.1163 albibarbis 0.8333 0.8333 1.0000 0.0000 antiochi 0.8889 0.7556 1.1765 0.1333 apache 0.8605 0.7209 11935 0.1395 apicalis 0.7660 OS532 1.3846 0.2128 argentifrons 0.8776 0.7551 1.1622 0.1224 argyrosoma 0.8889 057333 2A 0.1556 arida 0.9091 0.7636 1.1905 0.1455 armata 0.9063 0.7500 1.2083 0.1563 aurimystaceus 0.8684 0.6316 1.3750 0.2368 auripila 1.0000 0.9184 1.0889 0.0816 azteci 0.7143 0.6000 1.1905 0.1143 basingeri 0.8039 0.6863 1.1714 0.1176 basini 0.9091 0.7500 e221 0.1591 beameri 0.7368 0.5789 22a 0.1622 belfragei 0.8649 0.7027 1.2308 0.1622 benedicti 0.8261 0.6522 1.2667 0.1739 bexarensis 0.9592 0.7143 1.3429 0.2449 bicaudata 0.8776 0.7959 1.1026 0.0816 bicolor 0.7879 0.5303 1.4857 0.2576 bryanti 0.8261 0.6739 1.2258 0.1522 cabeza 0.8235 0.7059 1.1667 0.1176 caliente 0.8500 0.7000 1.2143 0.1500 californica 0.8333 0.6875 e221 0.1458 cana 0.9583 0.8958 1.0698 0.0625 canella 0.7200 0.5800 1.2414 0.1400 clementi 0.7778 0.5556 1.4000 0.2222 coquilletti 0.7429 0.6286 1.1818 0.1143 costalis 0.7209 0.5349 1.3478 0.1860 coulei 0.9118 0.8235 1.1071 0.0882 cressoni 0.6667 0.4615 1.4444 0.2051 davisi 0.8333 0.6667 1.2500 0.1667 deserti 0.8222 0.7556 1.0882 0.0667 ehrenbergi 0.6757 0.5405 1.2500 0.1351 femoratus 0.8167 0.6333 1.2895 0.1833 frewingi 0.8837 0.6977 1.2667 0.1860 gila 0.8824 0.7451 1.1842 0.1373 halli 0.8780 0.7805 1.1250 0.0976 helenae 0.9434 0.7170 1.3158 0.2264 inflata 0.7917 0.6250 1.2667 0.1667 interrupta* 0.9434 0.9623 0.9804 —0.0189 Jubata 0.7091 0.5091 1.3929 0.2000 kansensis 0.8500 0.6000 1.4167 0.2500 kelloggi 0.9200 0.7600 1.2105 0.1600 latruncula 0.7556 0.6000 1.2593 0.1556 leucocoma 0.9231 0.6154 1.5000 0.3077 luna 0.8222 0.7333 ee 0.0889 mortensoni 0.9592 0.8163 1.1750 0.1429 nemoralis 1.0638 0.8723 1.2195 0.1915 neoinflata 0.8654 0.6154 1.4063 0.2500 ordwayae 0.7750 0.6000 1.2917 0.1750 pallidula 0.8723 0.7021 1.2424 0.1702 parkeri 0.7949 0.6410 1.2400 0.1538 ——— SSS VOLUME 108, NUMBER 1 Table 4. Continued. DD) | Species Sleye v/eye tv CDI peralta 0.7111 0.4444 1.6000 0.2667 pilosa 0.8000 0.6000 1.3333 0.2000 pinali 0.9200 0.7000 1.3143 0.2200 plena 0.9677 0.8065 1.2000 0.1613 prairiensis 1.0952 0.9524 1.1500 0.1429 producta 0.6923 0.5128 1.3500 0.1795 prolifica 0.6667 0.5333 1.2500 0.1333 rapax 0.8077 0.5769 1.4000 0.2308 setigera 0.7200 0.5000 1.4400 0.2200 similis 0.7031 0.5469 1.2857 0.1563 spiniventris 0.7419 0.5806 1.2778 0.1613 staminea 0.8571 0.6905 1.2414 0.1667 subcuprea 0.7442 0.5814 1.2800 0.1628 tabescens 0.7333 0.5556 1.3200 0.1778 tagax 0.7353 0.5588 ES IESS 0.1765 texana 0.9091 0.7636 1.1905 0.1455 tolandi 0.8333 0.7500 eee 0.0833 truncata 0.8864 0.7500 1.1818 0.1364 tuberculata 0.7632 0.5263 1.4500 0.2368 tucsoni 0.6341 0.4390 1.4444 0.1951 utahensis 0.8824 0.6863 1.2857 0.1961 varipes 0.9583 0.7917 1.2105 0.1667 vertebrata 0.6538 0.4615 1.4167 0.1923 wilcoxi 0.7586 0.6207 12222. 0.1379 willistoni 0.8077 0.5769 1.4000 0.2308 yermo 0.8000 0.7250 1.1034 0.0750 yuma 0.6250 0.5208 1.2000 0.1042 zonata 0.6667 0.4444 1.5000 0.2222 face/vertex and CDI scores for females fall- ing completely within the upper range of male scores. The observed distribution for E. albibarbis may indicate a morphological shift in males toward larger vertex and eye widths related if there has been selective pressure that favors larger eyes in males (or smaller eyes in females). Results from this study suggest that the degree of sexual dif- ferences in eye morphology varies among populations of E. albibarbis, allowing the effect of various natural selective regimes on eye morphology to be examined. Habitat differences among populations of E. albibarbis have not been described, but the widespread distribution of this species, and high variation in eye morphology in fe- males and males relative to FE. aestuans, suggest that adaptation to local environ- ments is possible. Specimens of E. albibar- bis from BAJA have lower face/vertex and CDI means than for the MWUSA group, but extensive sampling is needed to char- acterize geographic groups and populations of this species. A simple mode of mate in- terception after short flights (similar to flight observed for prey capture) and the lack of courtship displays has been noted for E. albibarbis and most Efferia species (Lavigne 2002), and the same selective ad- vantage in mate acquisition assumed for holoptic males in other groups of Diptera can be invoked concerning the larger vertex and eye widths in E. albibarbis males. Op- timization of eye dimensions for successful prey capture and predator avoidance is like- ly to constrain levels of variation within and among robber fly species, with enhanced PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 69S0'0 + 8091°0 00910 LLOe 0 6810 0— LoIlO + 16sc'l vive I 0009'| r086 0 18 AONE 6810°0— F086 '0 10 ~DIdNtdaqul * rOeO'O + S860 IS6l 0 89€C 0 el9lO C9LO'0 + 7C6E I OOrT' I OOST I 8LLC I SO Bye[MoIoqn |, 88700 + ScSI'0 8eSl 0 OOSTc 0 Sc90'0 18800 + Sco | ICL c90r I 8690'| 6C BOUTUIEIS £6700 + L8SI0 OO9TO v9CC 0 91800 LSLO'O + 9LIC 1 SOIT | 8Sle I 9COI I LO SPIuIsog 80¢0'0 + 9LLI'0 OSLI0 80€Cc 0 teel O c6LO'O + O00CE'I L16c I vrrr I evi I 60 PIeULIe) 66700 + VCcEl O 98T1 0 O00cC'O e800 VSLO0 + eI8I'l OI8T | evle | IZOT'T 90 ephy v8c0O'0 + CLCILO cool’) LOOT O 6880 0 9LV0'0 + OI8I'I Crst | OOSC I CLEN| LO ByeuUOUuYy £6800 + 8col 0 ecol O LLOe 0 0000'0 CS810 + O89e'T LOIV I 0009'| 0000'| 60 siqreqtqTy cvv0 0 + IL810 90810 OOST 0 c9IlO c060'0 + LOE | LvOe | LOTH'I 6crl I 80 SUenSoOV ds = uray uPIPI| xPul Uru ds = urs urIpeW xPul uTu u dnoig «dds Id) XID A /99R4 60cI 0 + 1S$99°0 6£L9°0 £c96 0 O6er 0 ¥860'0 + 6SC8°0 19c8'0 cSo0 | OSc9'0 18 JEONG £c96 0 rere 0 10 DIAN AIOIU * O1SO0 + 8IIS 0 8cIS 0 90880 O06Eer 0 O00SO';0 + £OIL'O OOcL 0 GE Syeno) Ive9'0 SO BENOIT, OrITO + crOL 0 IcOL'O vcso 0 80cS'0 8LOL'O + L9S8'0 ILS8'°0 cS60'| OSc9'0) 6C BOUTIUEIS cIsO0 + 66¢L0 OLILO c918°0 £9890 O0970'0 + $8680 Less 0 coS6 0 SET8 0 LO SPIuIsod 8£90'0 + 79960 69LS 0 OOOL'0 S19V'0 £1900 + Orrl 0 crrl 0 O00S8'0 L999°() 60 BJBULIE) 00S0'0 + LOrL'O SSeEL0 SEc8'0 £9890 O870'0 + IeZL8'0 87880 00c6'0 6£08'0 90 epuyv €£90°0 + 6LOL'0 cool. 0 SO8ZL'0 0009°0 £090'0 + OSt8°0 cees 0 68880 evIL 0 LO IE) MAREN ONIN Y/ c9E1 0 + I9LS°0 6978 0 cee8 0 brrr 0 8660°0 + 06920 eSeLO leco0 8eS9 0 60 SIqIVqIq |W 8Z80°0 + LEe9'0 8S19°0 O18 0 cess 0 SOLO'O + 80C8'0 ees 0 cOe6 0 eceL 0 80 suenj{soV ds = uray URIpPo|\ xPul Uru ds = uropy UIP I XPul Uru u dnoip ‘dds oAq/XOU9 A, ohq/oory ‘sdnois sarsads 10J sd109S WINUITXPU IO WUMUITUTUU yuM pajuasoid ore Yidnisajul “[ JOY Sa1OdS,, ((UONRIAOP prepuRls [| Uva pue ‘URIpaW ‘WUNUUTXeU ‘WNUUTUTUT) sonsHeis AreuuNs Aq paquiosap se ‘viuaffq7 Jo sdnois satseads 10} (9961) XOSTIAA Ul poodar s}UdAIMsKaU WOT, paye[No[eo saiods [QD pur ‘xaplad/aovf{ ‘aka/Kajiaa ‘aka/aovf{ Ul UONeLIeA OYytoedsiaqUy *¢ aTqR VOLUME 108, NUMBER | 223 0,20 rs Pad a "@ Hi 4 s ra) O H rp : 4 H a : : t poi cages pot ala alle ngaa =e Soe, dosti —— -0,15 face/vertex Fig. 5. Correlation plot for face/eye and CDI scores, showing overlap between single exemplars of Efferia species (small squares, data in Table 4) and intraspecific variation in E. albibarbis and E. aestuans (see Fig. 2). Arrows indicate positions of Wilcox (1966) exemplars for E. albibarbis (lower left) and E. aestuans (upper right). 0,35 5 : ; : a 03 > a, ae : . ‘ H as 0,25 +4-- acl aa =-be- -- aa a ' 7 ‘ s a s ‘ ‘ Lil 0.27 Sao . are eather ae is i = La H = . = - ’ -_ "3 . rene : 5 [B191S) slecccosccsootcede -aoccoarcepedterenen secemngosas ebeuc marco COM eESS aa ae Le bsbineane +ihisdoo-esccsc me eet tegs nee epsetenanp ss fen seme tconnsnngsecnoneon: donee cheontte SemceaGanacnaraes sconce = 8 a" eal | : 2 = as 4 a ae H a e® ; i Pt Ope a em eww OR asa ee mc ee Pe ee a ni se a eee ee OF i : t t 1 | 0.95 1,05 145 1.25 1,85 145 1,55 1,65 0105; — face/vertex Fig. 6. Correlation plot of face/vertex and CDI scores for all Efferia group species (see Table 4) (triangle = E. albibarbis, diamond = E. aestuans). 224 mate acquisition promoting larger vertex and eye widths relative to face width in males of E. albibarbis. Interspecific differences.—Single speci- mens are insufficient for robust compari- sons between most Efferia species based on intraspecific variation in E. aestuans and E. albibarbis, but these scores suggest that certain species have notable eye morphol- ogies. As noted, face/vertex and CDI scores show that 7. interrupta and E. albibarbis are lower outliers and E. leucocoma and E. peralta are upper outliers among Efferia group species. The taxonomic status of Ef- feria species groups has not been confirmed or clarified by eye morphology variables, with scores resulting in continuous distri- butions. Group circumscriptions, however, have substantial ranges of interspecific var- iation in eye morphology based on Wilcox (1966) measurements. Efferia species ex- emplars can be distinguished by face/vertex and CDI scores although scores for nearly half of the Efferia species fall within the range of intraspecific variation in E. aes- tuans and do not allow 7. interrupta to be distinguished from E. albibarbis specimens. Furthermore, scores for EF. albibarbis and E. aestuans specimens vary considerably around the scores calculated for single ex- emplars of these species. Additional intra- specific sampling will reveal whether levels of variation for Efferia species are similar to E. albibarbis and E. aestuans, and should allow differences in eye morphology among Efferia species to be identified and com- pared. Eye morphology variables identified in this study can be used to investigate evo- lutionary changes within or among mono- phyletic groups and ecological guilds. Dif- ferences in habitat preferences between E. aestuans and E. albibarbis may explain the lack of sexual dimorphism in E. aestuans; E. albibarbis is found primarily on or near the ground and often in sandy areas where- as E. aestuans is primarily arboreal and found in forested habitats (Bullington and Lavigne 1984). Ecological classifications PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON for Asilidae proposed by Londt (1994) would place E. aestuans in Category 6, “trees,” and E. albibarbis in Category 1, ‘““sround,” and Efferia species have been recorded from these and other the four eco- logical categories. Further data on eye mor- phology may reveal correlations with “hunting areas” for species similar to cor- relations noted between oviposititor mor- phology and oviposition sites (Londt 1994). Two groups of Efferia species that are strongly supported as monophyletic, based on the laterally compressed mesonotum with a medial crest of macrosetae (the Car- inata Group) and tubercle-like projections from the abdominal sternites (the Tubercu- lata group), are not differentiable based on exemplar scores, but differences in eye morphology scores among species suggest that trends in eye morphology variation can be identified by variables defined in this study. The ecological diversity in Efferia provides an informative context for inves- tigating sexual dimorphism in E. albibarbis and testing evolutionary hypotheses for monophyletic groups in Efferia; these stud- ies would promote a deeper understanding of robber fly diversity. ACKNOWLEDGMENTS We thank Eric Fisher and Aubrey Scar- brough for improving the text of this paper as well as Donald Webb and the INHS for technical support during initial data collec- tion. LITERATURE CITED Artigas, J. N. and N. Papavero. 1997. The American genera of Asilidae (Diptera): keys for identifica- tion with an atlas of female spermathecae and oth- er morphological details. [X.2. Subfamily Asilinae Leach—Efferia group, with the proposal of five new genera and a catalogue of the neotropical spe- cies. Arquivos de Zoologia 34(3): 65—95. Bullington, S. W. and R. J. Lavigne. 1984. Description and habitat of Efferia kondratieffi sp. nov. with notes on Efferia aestuans (L.) (Diptera: Asilidae). Annals of the Entomological Society of America 77: 404-413. Buschbeck, E. K. and N. J. Strausfeld. 1996. Visual motion-detection circuits in flies: small-field retin- VOLUME 108, NUMBER 1 optic elements responding to motion are evolu- tionarily conserved across taxa. The Journal of Neuroscience 16(15): 4563—4578. . 1997. The relevance of neural architecture to visual performance: phylogenetic conservation and varation in dipteran visual systems. The Jour- nal of Comparative Neurology 383: 282—304. Fisher, E. M. and H. A. Hespenheide. 1992. Taxonomy and biology of Central American robber flies with an illustrated key to the genera (Diptera, Asilidae), pp. 610—631. /n Quintero Arias, D. and A. Aiello, eds. Insects of Panama and Meso-America: Se- lected Studies. Oxford University Press, Oxford. Hine, J. S. 1919. Robber flies of the genus Erax. An- nals of the Entomological Society of America 12(2): 103-157. Hull, RK M. 1962. Robber flies of the world. The genera of the family Asilidae. Bulletin of the United States National Museum 224(1, 2): 1-907. Lavigne, R. J. 2002. Evolution of courtship behavior among the Asilidae (Diptera), with a review of courtship and mating. Studia Dipterologica 9(2): 703-742. Lehr, P. A. 1988. Family Asilidae, pp. 197-326. In Soos, A. and L. Papp, eds. Catalogue of Palaearc- tic Diptera, Vol. 5, Akadémiai Kiad6, Budapest. 446 pp. Londt, J. G. H. 1994. Afrotropical Asilidae (Diptera) 26. Ethological observations, and a possible eco- 225 logical classification based on habits. Annals of the Natal Museum 35: 97-122. Majer, J. M. 1997. 2.38. European Asilidae, pp. 549— 566. Jn Papp, L. and B. Darvas, eds. Contribution to a Manual of Palaearctic Diptera (with Special Reference to Flies of Economic Importance), Vol. 2 [Nematocera and Lower Brachycera], Science Herald, Budapest, 566 pp. Martin, C. H. and N. Papavero. 1970. 35b. Family As- ilidae, pp. 1-139. In A Catalogue of the Diptera of the Americas South of United States. Museu de Zoologia, Universidade de Sao Paulo. McAlpine, J. EF 1981. Morphology and Terminology— Adults, pp. 9-63 In McAlpine, J. E, B.V. Peter- son, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds. Manual of Nearctic Diptera, Vol. 1. Monograph No. 27, Research Branch Ag- riculture Canada, Ottawa, 674 pp. Wilcox, J. 1966. Efferia Coquillett in America north of Mexico (Diptera: Asilidae). Proceedings of the California Academy of Science 34: 85-234. Wood, G. C. 1981. Asilidae, pp. 549-573. In Mc- Alpine, J. F, B.V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds. Manual of Nearctic Diptera, Vol. 1. Monograph No. 27, Research Branch Agriculture Canada, Ot- tawa, 674 pp. Yeates, D. K. 1994. The cladistics and classification of the Bombyliidae (Diptera: Asilioidea). Bulletin of the American Museum of Natural History 219: 1— Oil, PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 226-236 SYSTEMATIC PLACEMENT OF LYCAENA COGINA SCHAUS (LEPIDOPTERA: LYCAENIDAE: POLYOMMATINAE), A BIOGEOGRAPHICALLY DISJUNCT NEW WORLD SPECIES ROBERT K. ROBBINS AND MARCELO DUARTE (RKR) Department of Entomology, P.O. Box 37012, NHB Stop 127, Smithsonian In- stitution, Washington, DC 20013-7012 U.S.A. (e-mail: RobbinsR @si.edu); (MD) Colegao de Lepidoptera, Museu de Zoologia, Universidade de Sao Paulo, Av. Nazaré 481, 04263-000 Sao Paulo SP Brasil (e-mail: mduartes @usp.br) Abstract.—Balint and Johnson described genus Elkalyce in the Lycaenopsis Section of the Polyommatini for the South American butterfly Lycaena cogina Schaus and suggested that it is closely related to the tropical Asian Oreolyce Toxopeus. This systematic place- ment was based on four characters that are phylogenetically uninformative or incorrect. A medial uncus without lateral hairy lobes, cephalad entry of the ductus ejaculatorius into the penis, and brief anastomosis of forewing veins Sc and R, further falsify this systematic placement. Elkalyce cogina is transferred to the Everes Section of the Polyommatini fol- lowing an unpublished hypothesis from the late John Eliot, where it is likely a close relative, perhaps a congener, of the primarily eastern Asian Tongeia Tutt. Elkalyce and Tongeia are the only genera with “false” alulae, which we characterize morphologically, on the male genitalia penis, but the position of the “false”? alulae in each genus is slightly different. A lectotype is designated to preserve stability of the name Lycaena cogina Schaus, and the distribution and habitat of E. cogina are summarized. Six cases are noted in which a New World lycaenid species, or species pair, is most closely related to an Old World lineage, but E. cogina is the only endemic South American lycaenid whose closest relative is in the Old World. Whether or not Elkalyce is congeneric with Tongeia, the relict distribution of E. cogina suggests extinction in the intervening areas. Key Words: Elkalyce, Everes Section, false alulae, Lycaenopsis Section, Polyommatini, Tongeia Schaus (1902) described Lycaena cogina from southern Brazil (Castro, Parana). Even is not congeneric with other New World genera, but suggested that it is most closely though Lycaena Fabricius was widely used at that time for species now placed in sub- family Polyommatinae (Draudt 1919-1921, Eliot 1973), Schaus did not suggest those polyommatine species to which L. cogina might be most closely related. Balint and Johnson (1996) described genus Elkalyce for L. cogina in the Lycaenopsis Section of the Polyommatini (Eliot 1973, Eliot and Kawazoé 1983). They noted that Elkalyce related to Oreolyce Toxopeus, a genus that occurs primarily in the tropical parts of Asia. Alternatively, John N. Eliot examined a male of Lycaena cogina, including its genitalia, and proposed that it is related to Tongeia Tutt in the Everes Section of the Polyommatini (unpublished letters to Rob- bins, January 1988), a genus that occurs primarily in temperate and subtropical parts of Asia. Eliot died in 2003 without publish- VOLUME 108, NUMBER 1 ing this hypothesis (Eliot and Barlow 2003). The first purpose of this paper is to pub- lish the evidence supporting Eliot’s hypoth- esis. Roger Vila and colleagues (Museum of Comparative Zoology, Harvard Univer- sity) are sequencing parts of the DNA of Polyommatini worldwide with special em- phasis on the Neotropical fauna. When the DNA of E. cogina is sequenced (it has not yet been sampled; Vila, personal commu- nication), phylogenetic analysis of the se- quence data can then be compared with the different morphological hypotheses of Eliot and of Balint and Johnson. The second purpose of this paper is to summarize and make available information about the poorly known E. cogina. We il- lustrate adults of E. cogina because they were not figured in the major compendia of Neotropical lycaenid butterflies (Draudt 1919-1921, D’Abrera 1995). We also des- ignate a lectotype for E. cogina and sum- marize information on its distribution and habitat. The third purpose of this paper is to summarize those cases in which a New World lycaenid species, or species pair, is most closely related to species in the Old World. This summary serves as_ back- ground information for assessing the bio- geographic significance of the distribution of E. cogina. MATERIALS AND METHODS The results in this paper are based pri- marily upon an examination of 23 speci- mens of FE. cogina in the National Museum of Natural History (USNM), Smithsonian Institution, Washington, DC, USA, and in the Museu de Zoologia (MZSP), Universi- dade de Sao Paulo, SP, Brazil (Appendix). The genitalia of two males of E. cogina, one male of Tongeia fischeri (Eversmann), and one male of Oreolyce quadriplaga (Snellen) were dissected using standard en- tomological techniques (Robbins 1991). Genitalic terminology follows Klots (1970) except for the terms uncus and uncus lobes, 227), which follows Eliot (1973) and Eliot and Kawazoé (1983). Venation was studied us- ing standard techniques (Robbins 1991) and illustrated using digital scanning. Vein ter- minology follows Eliot and Kawazoé (1983). The distribution of E. cogina was determined from 86 specimens in MZSP, USNM, and DZUP (Universidade Federal do Parana, Curitiba, Brazil) plus literature citations, as noted in the Appendix. Notes on the habitat of E. cogina are based on the literature and on fieldwork by the authors in the states of Sao Paulo, Minas Gerais, and Rio de Janeiro. RESULTS AND DISCUSSION Does Elkalyce cogina belong to the Lycaenopsis Section of the Polyommatini? Balint and Johnson (1996: 345) based their hypothesis that E. cogina belongs to the Lycaenopsis Section on four characters: (1) hindwing tailless, (2) male genitalia with uncus lobes produced, (3) vinculum with a pronounced subtriangular extension directed cephalad, and (4) caecum more or less developed, suprazonal portion short. The Lycaenopsis Section has been revised worldwide, including figures of adults and genitalia (Eliot and Kawazoé 1983), and we illustrate the male genitalia of E. cogina (Fig. 1) and Oreolyce quadriplaga (Fig. 2). Oreolyce is the genus suggested by Balint and Johnson (1996) to be a close relative of E. cogina in the Lycaenopsis Section. More than 20 of the 30 sections of the Polyommatini include tailless species (Eliot 1973), so the first character listed by Balint and Johnson (1996) provides little phylo- genetic information. The second character does not distinguish the Lycaenopsis Sec- tion from other sections (Eliot and Kawa- zoé 1983). Further, the full character de- scription is “male genitalia with uncus lobes usually produced, sometimes to a stout spike, and turned inwards and down- wards” (Eliot 1973: 449). Oreolyce quad- riplaga (Fig. 2) has uncus lobes that fit this description. Although the medial uncus of 228 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1 Fig. 1. Male genitalia of Elkalyce cogina. A, lateral aspect of genital capsule, arrow points to the uncus. B, lateral aspect of penis and valva, arrow points to false alula. C, dorsal aspect of genital capsule, arrow points to the uncus. Scale 0.5 mm. E. cogina is posteriorly produced (Fig. 1), there are no lobes that are produced inwards and downwards. The ‘“‘vinculum hump” character was defined as “a triangular or semicircular projection on the proximal side of the vinculum” (Eliot and Kawazoé 1983: 16). This structure is conspicuous in O. quadriplaga (Fig. 2, additional figures in Eliot and Kawazoé 1983). It is questionable whether the curved vinculum of EF. cogina (Fig. 1) fits this definition, but if so, it is significantly less pronounced than the Ly- caenopsis Section. The caecum of the Ly- caenopsis Section (Fig 2, additional. figures in Eliot and Kawazoé 1983) is lacking in FE. cogina (Fig. 1). In support of these re- sults, the illustrations of the male genitalia of E. cogina in Balint and Johnson (1996) (their figs. 3-5) do not show (1) uncus lobes turned inwards and downwards as they are in the Lycaenopsis Section, (2) a vinculum hump equivalent to that in the Ly- caenopsis Section, or (3) a caecum. Other character information from the male genitalia and wing venation also fal- sify the hypothesis of Balint and Johnson (1996). A comparison of the male genitalia uncus of Elkalyce and Oreolyce in dorsal aspect (Figs. 1C, 2C) shows little morpho- logical similarity. The uncus of E. cogina is a medial, posteriorly produced process (Fig. 1), but the uncus in the Lycaenopsis Section is a transverse band that is laterally extended into paired hairy lobes (Fig. 2, ~100 genitalic illustrations in Eliot and Ka- wazoé 1983). The ductus ejaculatorius en- ters the cephalad side of the penis in E. cog- ina (Fig. 1), but enters the dorsal side of the penis in the Lycaenopsis Section (Fig. 2, Eliot 1973). Forewing veins Sc and R, VOLUME 108, NUMBER 1 Fig. 2. the “‘vinculum hump,” arrow on top points to the uncus, arrow at right points to the lobes of the uncus. B, lateral aspect of penis, arrow points to the caecum. C, dorsal aspect of genital capsule, arrow points to the lobes of the uncus. Scale 0.5 mm. anastomose and separate in E. cogina (Fig. 4), but do not anastomose in the Lycaen- opsis Section (Eliot 1973, Eliot and Ka- wazoé 1983). In summary, there is no substantive mor- phological evidence to support the place- ment of Elkalyce cogina in the Lycaenopsis Section. Does Elkalyce cogina belong to the Everes Section of the Polyommatini? In contrast to the Lycaenopsis Section, taxonomy within the Everes Section of the Polyommatini has not been revised on a worldwide basis and is somewhat chaotic. For example, although Tutt (1908) differ- entiated Everes Htibner and Cupido Schrank, these names have recently been treated as subgenera without phylogenetic argument falsifying Tutt’s classification (Hesselbarth et al. 1995, De Prins and Iver- sen 1996). We quote from John Eliot’s letter to Robbins (14 January 1988) and present the supporting evidence for his hypothesis Male genitalia of Oreolyce quadriplaga. A, lateral aspect of genital capsule, arrow at left points to that E. cogina is closely related to Tongeia in the Everes Section. “The possession of a distinct uncus, narrowly bifid at the apex, must put it [E. cogina] into . the Everes Section ..., with which its other characters of venation, eyes, palpi, etc. are consistent. The genitalia are ... remark- ably similar to those of Tongeia Tutt, 1908, a primarily eastern Palaearctic Everid genus, even down to the false ‘“‘alulae”’ on the penis, of which Tutt (1908: 43) says: “‘the aedoea- gus itself ... has a marked raised zone ... where supported.”” This feature cannot be seen in his Plate II, fig. 3, wherein the pho- tographed genitalia are very distorted. How- ever, excellent figures of Tongeia genitalia can be found in Shirozu (1960) and Kawazoe & Wakabayshi (1976).” The male genitalia of E. cogina (Fig. 1) and Tongeia fischeri (the type species of Tongeia) (Fig. 3) are phenetically similar (also illustrated on page 170 in Kawazoé and Wakabayshi 1976), as noted by Eliot, and have “‘false’”’ alulae. Nabokov (1945: i) eS) io) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Male genitalia of Tongeia fischeri. A, lateral aspect of genital capsule. B, lateral aspect of penis and valva, arrow points to false alula. C, dorsal aspect of genital capsule. Scale 0.5 mm. 48) introduced the term alulae to describe *“‘out-turned flaps of subzonal sheath,’ and it was later defined in the glossary to Klots (1970: 221) as “‘paired flaps of manica, lat- Fig. 4. wing (digitally scanned). A, terminus of vein R,. B, Basal, anterior part of Elkalyce cogina fore- terminus of vein Sc. C, anastomosis of veins Sc and R,. erad of its attachment to the aedoeagus” (figs. 119, 121 in Eliot 1973). Alulae are sclerites of the manica and occur in the Eu- chrysops and Polyommatus Sections of the Polyommatini (Eliot 1973). The “‘false”’ al- ulae of Tongeia and Elkalyce are out- growths of the outer wall of the phallus, perhaps the same structure that Snodgrass (1935) termed the lepidopteran theca. Whatever the homology, they are not scler- ites of the manica, which is presumably why Eliot referred to them as “false” alu- lae. *‘False”’ alulae are known only in Ton- geia and Elkalyce. They are unrecorded in other genera of the Everes Section, specif- ically in Everes, Cupido, Bothrinia Chap- man (= Bothria Chapman), Shijimia Mat- sumura, Talicada Moore, and Binghamia Tutt (Chapman 1908, Tutt 1908, Shir6zu 1960, Elot 1973, Kawazoé and Wakabay- shi 1976). The male genitalia of Tiora Evans do not seem to have been illustrated, VOLUME 108, NUMBER 1 Fig. 5: Female forewing length 10.5 mm. but Tiora has been considered a junior sub- jective synonym of Everes (Hesselbarth et ale Hi995): As noted by Eliot, the morphology of E. cogina 1s consistent with the characteriza- tion of the Everes Section as outlined in Eliot (1973). Forewing veins Sc and R, anastomose briefly (Fig. 4), the hindwing is tailless (Fig. 5), battledore scales are absent (E. cogina and Tongeia males lack both an- droconia and blue scales dorsally), the eyes are smooth, the palpi are hairy, and the male genitalia are typical of the Everes Sec- tion (cf. fig. 116 in Eliot 1973). Although the evidence presented in the preceding paragraphs supports a close re- lationship between Tongeia and Elkalyce, we do not synonymize them. The “‘false al- ulae”’ of Tongeia (Fig. 3, male genitalic il- lustrations of other Tongeia species in Shi- rozu 1960) are more posterior than those of E. cogina (Fig. 1), casting some doubt on Male and female Elkalyce cogina. Female on top. Dorsal surface on left. Female is the lectotype. their homology. Characters of the female genitalia are likely to be an important line of evidence in the placement of Elkalyce, but documenting female genitalic structures in the Everes Section is beyond the scope of this small project. However, if new evi- dence supports the monophyly of Tongeia + Elkalyce, Elkalyce should be synony- mized because monotypic genera are of du- bious classificatory value (Farris 1976). Nomenclature.—To preserve stability of the name Lycaena cogina Schaus, we des- ignate as lectotype a female with a red type label corresponding with the original de- scription (Type No. 5920) and a handwrit- ten label (Lycaena cogina Schs type) from Castro, Parana (Figs. 5-6). A male and a female with the same red type label are from Sao Paulo and Rio de Janeiro states, respectively (Appendix), and lack the word *“*type”’ on the handwritten label. Since the original description noted only Castro, Pa- 232 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Lycaena cogina Schaus By: Robbins/Duarte 2006 Fig. 6. Labels on the lectotype. rand, Brazil, these two are not types. A fourth specimen has a “Castro, Parana”’ lo- cality label of the same kind as the lecto- type. Although it lacks a red type label, it could be a type (Appendix). Since the orig- inal description did not mention the number of specimens, our lectotype designation makes the name bearing type of this name clear. Habitat and distribution.—Brown and Mielke (1967: 151) noted that E. cogina is ‘“‘partial to marshy areas”’ in the central pla- teau of Brazil. In our limited experience with this species, it occurs not uncommonly in open areas, whether disturbed roadsides, cerrado vegetation, or grasslands, albeit, al- ways in the vicinity of wetlands. It is re- corded from 500 to 1,700 m elevation. Dates of capture include most months of the year, but no captures have been recorded in October and November. Elkalyce cogina occurs widely in the central plateau of Brazil (south of 10°S lat- itude), southern Brazil, and northeastern Argentina (Fig. 7, Appendix). Balint and Johnson (1996: 344) gave the distribution of E. cogina as “‘known only from Parana, Brazil at present.”” Anomalously, they ex- | 2 =no specific locality ice 72 300 O 300 Kilometers = | Distribution of Elkalyce cogina based on specimens and literature citations in the Appendix. VOLUME 108, NUMBER 1 amined specimens from Campos de [sic] Jordao, Sao Paulo, Brazil, and cite Hayward (1973) and Ebert (1969), who recorded E. cogina from Misiones, Argentina and Pogos de Caldas, Minas Gerais, Brazil, respective- ly. Further, Brown and Mielke (1967) had recorded it from many other Brazilian lo- calities. Other than Elkalyce, the only New World genus in the Everes Section is Ev- eres, Which occurs from Canada to the low- lands of Costa Rica and Panama (Godman and Salvin 1889, Robbins unpublished) and is more than 4,000 km distant from the known range of E. cogina. Disjunct New World Lycaenidae.—Al- though some lycaenid genera are Holotrop- ical (e.g., Leptotes Scudder, Zizula Chap- man) (Clench 1964, Eliot 1973) or Holarc- tic (e.g., Satyrium Scudder, Callophrys Bill- berg) (Clench 1961), there are six cases in which a New World lycaenid species (or non-sympatric species pair) is most closely related to an Old World lineage. Using the classification of Eliot (1973) and the recent overview of the Neotropical Lycaenidae (Lamas 2004a, b, c; Robbins 2004), these cases are (1) the North American Miletinae, Feniseca tarqunius (Fabricius), whose wing pattern, male genitalia, and pupal shape closely resemble Spalgis Moore in the Af- rican and Oriental Regions (Eliot 1973), (2) the North American Theclinae, Habrodais grunus (Boisduval) (including H. poodiae Brown & Faulkner in Baja California), which is probably most closely related to the /ratsume Sibatani & Ito in temperate Asia (Shir6zu and Yamamoto 1956), (3) the North American Theclinae, Hypaurotis cry- salus (Edwards), which is probably most closely related to the Palearctic Favonius Sibatani & Ito and Quercusia Verity (Shi- rozu and Yamamoto 1956), (4) the montane Guatemalan and Mexican Lycaeninae, /o- phanus pyrrhias (Godman and Salvin), whose wing pattern, male foreleg, and gen- italia resemble those of Melanolycaena Si- batani from montane New Guinea (Sibatani 1974), (5) the Polyommatinae genus Bre- phidium Scudder, which occurs in the An- 233, tillean Basin (Brephidium pseudofea {Mor- rison]), the Nearctic (B. exilis [Boisduval]), and South Africa (Eliot 1973), and (6) the South American Polyommatinae, Elkalyce cogina, which belongs to the primarily Pa- learctic Everes Section and may be conge- neric with the primarily temperate Asian Tongeia. There is currently a healthy controversy concerning disjunct butterfly taxa and whether current distributions are due to vi- cariance or dispersal (e.g., de Jong 2003, Hall et al. 2004, Braby et al. 2005). Elka- lyce cogina is the only endemic South American lycaenid whose closest relatives are in the Old World. It is the only case in the butterflies, of which we are aware, in which a species restricted to eastern South America is most closely related to lineages in temperate Asia. Whether or not E. cogina belongs to Tongeia, its relict distribution suggests extinction in the intervening areas, as was recently shown with an amber fossil and its extant relatives in the Riodinidae (Hall et al. 2004). ACKNOWLEDGMENTS This paper is dedicated to the memory of John Eliot, perhaps the butterfly world’s consummate amateur. We are indebted to our colleagues Gerardo Lamas and Olaf Mielke for sending data and bibliographic information and for other kindnesses too numerous to enumerate. We thank Nancy Vannucci for her efforts in finding speci- mens of E. cogina from Misiones, Roger Vila for inspiring us to write this paper, and Keith Brown for kindly sharing data on E. cogina. For the genitalic illustrations, we thank Vichai Malikul. For commenting on the manuscript, we are most grateful to Bob Busby, Jason Hall, Gerardo Lamas, and Roger Vila. The junior author has been fi- nancially supported by the State of Sao Paulo Research Foundation—FAPESP (02/ 13898-0 and 03/05895-4). FAPESP has also supported a research trip by RKR to Sao Paulo (process number 03/13985-3). 234 LITERATURE CITED Balint, Z. and K. Johnson. 1996. Systematic position and generic status of Lycaena cogina Schaus, 1902: An endemic Neotropical Lycaenopsina (Lepidoptera, Lycaenidae). Acta zoologica Aca- demiae Scientiarum Hungaricae 41(4): 343-347. Braby, M. FE, J. W. H. Trueman, and R. Eastwood. 2005. When and where did troidine butterflies (Lepidoptera: Papilionidae) evolve? Phylogenetic and biogeographic evidence suggests an origin in remnant Gondwana in the late Cretaceous. Inver- tebrate Systematics 19: 113-143. Brown, K. S. Jr. and O. H. H. Mielke. 1967. Lepidop- tera of the Central Brazil Plateau. I. Preliminary list of Rhopalocera (continued): Lycaenidae, Pier- idae, Papilionidae, Hesperiidae. Journal of the Lepidopterists’ Society 21(3): 145-168. Canals, G. R. 2003. Mariposas de Misiones/Butterflies of Misiones. Buenos Aires, L.O.L.A. Literature of Latin America, 476 pp. Chapman, T. A. 1908. Two new genera (and a new species) of Indian lycaenids. Proceedings of the Zoological Society of London 1908: 676-678. Clench, H. K. 1961. Tribe Theclini, pp. 177—220. In Ehrlich, P. R. and A. H. Ehrlich. How to Know the Butterflies, Brown Co., Dubuque, Iowa, 262 pp-., 525 figs. . 1964. A synopsis of the West Indian Lycaen- idae, with remarks on their zoogeography. Journal of Research on the Lepidoptera 2: 247-270. D’Abrera, B. L. 1995. Butterflies of the Neotropical Region. Part VII. Lycaenidae. Black Rock, Hill House, pp. 1-xi + 1098-1270. De Jong, R. 2003. Are there butterflies with Gond- wanan ancestry in the Australian region? Inver- tebrate Systematics 17: 143-156. De Prins, W. and E Iversen.1996. Family Lycaenidae, pp. 205-209. In Karsholt, O. and J. Razowski, eds. The Lepidoptera of Europe, A Distributional Checklist. Apollo Books. Stenstrup 380 pp. Draudt, M. W. K. 1919-1921. Familie: Lycaenidae. Jn Seitz, A., ed. Die Gross-Schmetterlinge der Erde. Stuttgart, Alfred Kernen 5: 744-831, 144-159 pls. Ebert, H. 1969. On the frequency of butterflies in East- ern Brazil, with a list of the butterfly fauna of Pogos de Caldas, Minas Gerais. Journal of the Lepidopterists’ Society 23 (Supplement 3): 1—48. Eliot, J. and H. Barlow. 2003. Lieutenant Colonel John Nevill Eliot. 29 August 1912-11 April 2003. Ma- layan Naturalist 57: 55—56. Eliot, J. N. 1973. The higher classification of the Ly- caenidae (Lepidoptera): a tentative arrangement. Bulletin of the British Museum of Natural History (Entomology) 28(6): 373-506. Eliot, J. N. and B. D’Abrera. 1992. The Butterflies of the Malay Peninsula. Fourth Edition (revised). Kuala Lumpur, Malayan Nature Society, 596 pp. Bliot, J. N. and A. Kawazoé. 1983. Blue Butterflies of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the Lycaenopsis group. London, British Museum (Natural History), 309 pp. Farris, J. S. 1976. Phylogenetic classification of fossils with recent species. Systematic Zoology, 25: 271— 282. Godman, E D. and O. Salvin. 1887. Biologia Centrali- Americana. Insecta. Lepidoptera-Rho-palocera. Fam. Lycaenidae. 2: 1-112. Hall, J. W., R. K. Robbins, and D. J. Harvey. 2004. Extinction and biogeography in the Caribbean: new evidence from a fossil riodinid butterfly in Dominican amber. Proceedings of the Royal So- ciety of London B 271: 797-801. Hayward, K. J. 1951. Catalogo sinonimico de los ro- paloceros argentinos excluyendo “Hesperiidae.” Acta Zoologica Lilloana 9: 85 281. . 1973. Catalogo de los ropaléceros argentinos. Opera Lilloana 23: 1-318. Hesselbarth, G. H., V. Oorschot, and S. Wagener. 1995. Die Tagfalter der Tiirkei: unter Beriicksichtigung der angrenzenden Linder. Bocholt. Vol. 1. Ger- many, 753 pp. Kawazoé, A. and M. Wakabayshi. 1976. Coloured Il- lustrations of the Butterflies of Japan. Hoikusha Publishing, Osaka, 422 pp. Klots, A.B. 1970. Lepidoptera, pp. 115-130. In Tuxen, S. L., ed., Taxonomist’s Glossary of Genitalia in Insects. Munksgaard, Copenhagen. Lamas, G. 2004a. Lycaenidae. Theclinae. Tribe The- clini, p. 137. Jn Lamas, G., ed. Checklist: Part 4A. Hesperioidea—Papilionoidea. /n Heppner, J. B., ed. Atlas of Neotropical Lepidoptera. Volume 5A. Gainesville, Association for Tropical Lepidoptera; Scientific Publishers. . 2004b. Lycaenidae. Lycaeninae, pp. 137-138. In Lamas, G., ed. Checklist: Part 4A. 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VOLUME 108, NUMBER 1 Gainesville, Association for Tropical Lepidoptera; Scientific Publishers. Schaus, W. 1902. Descriptions of new American but- terflies. Proceedings of the United States National Museum 24(1262): 383—460. Sibatani, A. 1974. A new genus for two new species of Lycaeninae (S. Str.) (Lepidoptera: Lycaenidae) from Papua New Guinea. Journal of the Australian Entomological Society 13: 95-110. Shir6zu, T. and H. Yamamoto. 1956. A generic revi- sion and the phylogeny of the tribe Theclini (Lep- idoptera: Lycaenidae). Sieboldia 1: 329—421. Shirézu, T. 1960. Butterflies of Formosa in Colour. Hoikusha Publishing. Osaka, 481 pp. Snodgrass, R. E. 1935. Principles of Insect Morphol- ogy. McGraw-Hill Book Co., New York, 667 pp. Tutt, J. W. 1908. A Natural History of the British Lep- idoptera, Their World-Wide Variation and Geo- graphical Distribution, Vol X. Swan Sonnenshein. London, 410 pp. APPENDIX Specimens examined and literature cita- tions for Elkalyce cogina (museum acro- nyms in Materials and Methods). If eleva- tion was not recorded on a specimen’s data label, we parenthetically note the elevation for that locality as it is listed in gazetteers. BRAZIL (north to south): DistRITo FED- ERAL. | d Southwest of Sobradinho (So- bradinho River), 1,025 m, 12 August 1965 (Brown and Mielke 1967: 151); 1 2 Bra- silia, Fazenda Agua Limpa [an ecological station of the University of Brasflia] (1,000—1,200 m), 22 May 1976 (USNM); 1 3 Brasilia, Ribeirao da Contagem (1,100 m), 23 February 1966 (DZUP); 1 ¢ same locality, 25 February 1966 (DZUP); 1 6 Brasilia, no specific locality, 23 February 1968 (DZUP); A number of ¢ and 2, Bra- silia, Jardim Zoologico de Brasilia, 1,020 m, 8 June 1966 (Brown and Mielke 1967: 151); 3 d and 4 2 Brasilia, Reserva Eco- l6gica do IBGE (1,000—1,200 m), 23 March [987° (DZUP): GotAs. | d Campinas [suburb of present- day Goidnia] (800 m), March 1930 (MZSP); A number of 6 and 2, 30 km north of Brasilia (Maranhao River), 700 m, 12 June 1966 (Brown and Mielke 1967: 151); | gd 30 km north of Brasilia (Mar- anhao River), 700 m, 17 August 1965 235 (Brown and Mielke 1967: 151); 1 ¢ Goids Velho [also known as “‘Goias,”’ city of the state of Golds, 144 km north of Goidnia, 15°57'S, 50°07'W] (500 m), 30 May 1976 genitalia dissection by J. N. Eliot] (USNM); | 3 same locality, 20 June 1976 [genitalia dissection by R. K. Robbins] (USNM); | & Viandpolis (1,000 m), March 1930 (MZSP). MINAS GERAIS. | 2 Paraopeba, 3 km east of BR-040 (Paraopeba Woods), 750 m, 7 June 1966 (Brown and Mielke 1967: 151); 5 6 Catas Altas, Caraga, 1,300—1,500 m, 1—5 February 1985 (DZUP); 1 ¢ same lo- cality, 1,300 m, 4-6 February 2003 (DZUP); 1 5 Carmo do Rio Claro (859 m), 20 February 1959 (DZUP); 1 3 Barbacena, 1,100 m, 20 July 1951 (DZUP); 1 ¢ same locality, 9 August 1951 (DZUP); 1 2 Var- ginha (600—-1,000 m), February 1972 (DZUP); Several specimens, Pocos de Cal- das, 1,000—1,500 m, February, April and May (Ebert 1969: 41); 1 2 15 km southeast of Itamonte, 22°21.8'S, 44°47.5'W, 1,450 m, 25 April 1994 (USNM). RIO DE JANEIRO: | 2 Teresopolis, Parque Nacional Serra dos Orgdos, 22°27'S, 43°00'W, 1,100 m, 16 February 1995 (USNM); | @ Itatiaia, Parque Nacional do Itatiaia, 22°27'S, 43°37'W, 1,100 m, 5 May 1995 (USNM); 2 ¢ Petrépolis, Sao José (800—900 m), 30 February 1954 (DZUP); 1 2 Petrdépolis (800 m), no date (USNM, from Schaus Collection, with a red label » Lype Nc; 5920 US NEM. * buts not.a type—see text). SAG) PAULO? IG Rio Claro; G00) m, 23 June 1963 (USNM); 1 ¢ Serra Negra (1,000 m), 24 September 1957 (DZUP); 1 2 same locality, 12 September 1957 (DZUP); 2 ¢ Campos do Jordao, 1,600 m, 26 January 1966. Cited in Balint and John- son (1996: 344); 6 5d and 1 2 same locality, Parque Estadual de Campos do Jordao, 1,600-1,700 m, 22-25 January 1992 (DZUP); 1 2 Itatiba (760—785 m), Decem- ber. 1935 "(MZSBP) 2" 2 "Sera “do Japi: 23°15'S, 46°54’'W, 1,100 m, 12 April 1991 (USNM); 1 2 Sao Paulo, ‘“‘“Matto do Gov- 236 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON erno”’ [presently known as “‘Parque do Es- tado”’ including both the Zoological and Botanical Parks] (760 m), February 1914 (MZSP); 1 o& same locality, May 1919 (MZSP); 1 d Sao Paulo, Santo Amaro [a neighborhood of the city of Sao Paulo] (760 m), 1 2 April 1945 (MZSP); 1 2 same lo- cality, 24 December 1958 (MZSP); 1 6 same locality, 26 December 1958 (MZSP); 1 6 Sao Bernardo do Campo (762 m), 29 April 1927 (MZSP); 1 ¢ Batatais (733 m), 25 December 1968 (DZUP); 1 2 Sao Pau- lo, no specific locality, no date (MZSP); 1 3 Sao Paulo, no specific locality, no date (USNM, from Schaus Collection, with a red label“ iype: No: 5920 (U:S2N-M..”” ‘but is not a type—see text). PARANA. | @ Castro (999 m), no date (USNM, from Schaus Collection, with a red label ‘““Type No. 5920 U.S.N.M.,”’ desig- nated lectotype); Sex undetermined (no ab- domen or forelegs), same locality, no date (USNM); 2 3 same locality, Ribeira, 530 m, 20 December 2002 (DZUP); 2 d Ponta Grossa, no specific locality, May 1947 (DZUP); 1 2 same locality, Olaria (700 m), no date (DZUP); 2 ¢ and 1 2 same local- ity, Piriquitos. (900 m),.3:cApril) 1971 (DZUP); 1 3 same locality, Rio Bonito (900 m), March 1947 (DZUP); 1 2 Cam- pina Grande do Sul, Jaguatirica, 1,000 m, 27 February 2003 (DZUP); | 6 Jaguariaiva (850 m), April 1951 (DZUP); 8 ¢ and 4 2 Campo Largo, Trés Corregos, 700 m, 7 March 1998 (DZUP); 1 6 same locality, 30 km north of Bateias (880 m), 4 March 2000 (DZUP); 1 2 Balsa Nova, Sao Luiz do Pu- runa, 900—1,000 m, 8 March 1980 (DZUP); 1 2 same locality, 25 February 1984 (DZUP); 2 2 same locality, 12 April 1986 (DZUP). Rio GRANDE DO SUL. | 2 no specific lo- cality, no date (MZSP). ARGENTINA: Misiones, no specific lo- cality (Hayward 1951: 142). Canals (2003) added no new information on E. cogina in Misiones. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 237—239 NOTE Adventive Hylaeus (Spatulariella Popov) in the New World (Hymenoptera: Apoidea: Colletidae) The bee subgenus Hylaeus (Spatulariella Popov) includes about 18 species native to the Palearctic (Michener 2000). Six species are known from Europe (Dathe 1980), of which two have been recorded as adventive in the New World, H. punctatus (Brullé) and H. hyalinatus Smith. The first known collection of subgenus Spatulariella in the Western Hemisphere was of H. punctatus at Playa del Rey, Los Angeles County, south- ern California, in 1981 (Snelling 1983). Subsequently, H. punctatus has been found elsewhere in California (Ascher 2001) and in the city of Santiago in central Chile (Toro et al. 1989). A second species of Spatular- iella, H. hyalinatus, has recently been re- corded from the Fingerlakes Region of cen- tral New York State (Ascher 2001). The purpose of this note is to further document the establishment and spread of these two adventive bee species in the New World. Information about the identification of the two adventive Hylaeus (Spatulariella), including illustrations and keys, can be found in Dathe (1980), Snelling (1983), Toro et al. (1989), and Ascher (2001). Hy- laeus hyalinatus males can be recognized by the shape of the eighth sternum (S8); the distal spatulate process is connected to the base by an extremely narrow elongate stalk. In Hylaeus punctatus males, the distal spat- ulate process of S8 is broadly connected to the base. In addition, H. punctatus males have less extensive face markings; the su- praclypeal area is usually black and lateral marks extend little, if at all, above the level of the epistomal sulcus. Hylaeus hyalinatus.—tThe first report of this species in the New World was based on specimens collected during 1998-2000 on and near the campus of Cornell University in Ithaca, Tompkins County, New York (Ascher 2001). Subsequently, numerous H. hyalinatus were collected in Ithaca, else- where in Tompkins County such as along Route 34 in the Town of Newfield, and in New York City and adjacent Westchester County. Records of specimens collected by P. Gambino in New York City and depos- ited in his collection include: Bronx Coun- ty: | 2, Bronx High School of Science, 10 August 1997; 1 @, Van Cortlandt Park, Gunhill Road, 6 July 2000; New York County (Manhattan): 4 ¢d, Hudson River at Rector/Liberty St., 29 July 1999; 1 2, Rob- ert Wagner Park, 8 July 1999. Specimens collected in urban gardens in Manhattan and the Bronx by Kevin M. Cox include: Bronx Co.: | ¢, Bronx Bathgate Garden, 25 June 2004; | 2, Fordham Bedford Lot- Busters Garden, 25 June 2004, yellow pan; 1 2, Krystal Garden, 25 June 2004; 1 6, Tremont Community Garden, 11 June 2005; New York Co.: 1 6, Harlem, Holy Rosary Garden, 27 May 2004. Additional specimens have been collected in Manhat- tan by J. S. Ascher in Central Park and Riv- erside Park (e.g., 1 d: 31 July 2002). Spec- imens collected during the Bronx River BioBlitz on 10 June 2005 by P. Gambino et al. are: Bronx County: 1 6, Drew Garden; 2 3, Vyse Avenue at 180th Street; 1 2, Prospect Avenue at 180th Street; Westch- ester County: 2 2, Bronx River at Cross County Parkway. Hylaeus hyalinatus has not yet been found in Orange or Putnam counties in New York, or Fairfield County in Connect- icut, which are all slightly to the north of Westchester County, despite intensive re- cent collecting in these areas. Records of flight in New York State now extend from May to October. The 1997 record from the 238 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Bronx is the earliest known collection of H. hyalinatus in the New World. Hylaeus punctatus.—The first records of this species from North America outside of California are of 15 2 and two ¢ speci- mens collected on the National Mall in the District of Columbia in 2004 by Sam Droe- ge. One male was collected from flowers in the National Botanical Garden on 6 August; the other male was collected 3 August in the garden along the National Museum of Natural History along with 3 females. An- other 12 females were collected in the Nat- ural History Museum garden at an undeter- mined date in August of the same year. Both locations contained an unusual diver- sity of plant species with the original stock coming from throughout North America, but not Europe. Despite extensive surveys of gardens and natural communities else- where in the downtown sections of Wash- ington and its suburbs, no other individuals of this species have been detected. The first records of H. punctatus from Chile outside of the city of Santiago are of two series found in the American Museum of Natural History collection and deter- mined by JSA. These series are from Re- gion V in Petorca Province, north of San- tiago, and from Region VI in Cachapoal Province, south of Santiago (in the Region Metropolitana: Santiago Province). The se- ries from Petorca Province is of 2 2 and 2 3d, collected at Cuesta El Melon in Novem- ber 1998 by R. Madariaga. The series from Cachapoal Province is of 24 2 and 25 ¢ collected at Lago Rapel in January 2001 by Madariaga. New Records from Santiago are of 2 2 and 1 6 collected at Cerro San Cris- t6bal on 9 February 1994 by J. M. Carpen- ter and A. Davidson. These records dem- onstrate that H. punctatus is not restricted to urban areas in Chile. Specimens of H. punctatus from H. Toro’s personal collec- tion cited in Toro et al. (1989) are now in the collection of the American Museum of Natural History. In both its native range and in the New World, H. punctatus is best known from Mediterranean-climate regions. However, records from central Europe (Dathe 1980) and the new record from Washington, D.C., suggest that this species has the potential to become established across much of temper- ate North and South America. Of the two adventive Spatulariella species, H. hyali- natus is the most likely to extend its range to the north, as this species is widely dis- tributed in Europe north to Finland. Discussion.—The disjunct distributions of Hylaeus hyalinatus and Hylaeus punc- tatus are consistent with human-assisted in- vasion processes, i.e., introduction and es- tablishment at isolated foci, from which populations locally disperse and colonize. Many records of both Spatulariella are from gardens and/or near greenhouses Where recently imported exotic plants are found. The potential role of botanical gar- dens, greenhouses, and nursery companies in spreading exotic bees both between and within countries should be assessed. Spe- cies of Hylaeus (Spatulariella) may be par- ticularly prone to long distance transport as their nests are small and concealed within a variety of natural and artificial cavities (Ascher 2001). Acknowledgments.—We thank Kevin M. Cox of Fordham University for making available specimens of H. hyalinatus from his ongoing studies of pollinators in New York City gardens. LITERATURE CITED Ascher, J. S. 2001. Hylaeus hyalinatus Smith, a Eu- ropean bee new to North America, with notes on other adventive bees (Hymenoptera: Apoidea). Proceedings of the Entomological Society of Washington 103: 184—190. Dathe, H. H. 1980. Die Arten der Gattung Hylaeus F in Europa (Hymenoptera: Apoidea, Colletidae). Mitteilungen Zoologisches Museum in Berlin 56: 207-294. Michener, C. D. 2000. The Bees of the World. The Johns Hopkins University Press, Baltimore and London, 913 pp. Snelling, R. R. 1983. Studies on North American bees of the genus Hylaeus 6. an adventive Palearctic species in southern California (Hymenoptera: Col- letidae). Bulletin of the Southern California Acad- my of Sciences 82: 12-16. VOLUME 108, NUMBER 1 Toro, H., Y. Frederick, and A. Henry. 1989. Hylaeinae (Hymenoptera: Colletidae), a new sub-family of bees for the Chilean fauna. Acta Entomologica Chilena 15: 201—204. John S. Ascher, Parker Gambino, and Sam Droege. (JSA, PG) Division of Invertebrate 239 Zoology, American Museum of Natural His- tory, New York, NY 10024-5102, U.S.A. (e- mail: ascher@amnh.org); (SD) USGS, Pa- tuxent Wildlife Research Center, BARC- East, Building 308, Room 124, 10300 Bal- timore Ave., Beltsville, MD 20705, U.S.A. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, p. 240 NOTE Nesidiocheilus hawaiiensis Kirkaldy, 1902, a Synonym of Acanthia campestris Fabricius, 1794 (Hemiptera: Heteroptera: Lyctocoridae) Acanthia campestris Fabricius, 1794, has been known as Lyctocoris campestris (Fa- bricius) for many years (Péricart 1972, 1996). Described from Europe, it has been introduced into many countries around the world, usually via stored products (Cham- pion 1900, Hiura 1966, Chu 1969, Ford 1979, Henry 1988), including the island of Hawaii (Beardsley 1981). Nesidiocheilus hawaiiensis was described from Maui, Ha- leakala, 7,000-10,000 feet by Kirkaldy (1902). Zimmerman (1948) included an il- lustration of the type in the British Museum (Natural History) (as Lyctocoris hawatien- sis). Zimmerman indicated that W.E. China of the British Museum stated that it was likely only a “‘form” of Lyctocoris cam- pestris. Beardsley (1981) reported speci- mens of Lyctocoris campestris collected on the island of Hawaii, on Mauna Kea, at 7,800 feet. These specimens were sent to J.L. Herring of the U.S. Department of Ag- riculture, Systematic Entomology Labora- tory, Washington, D.C. Herring stated “‘As far as the Lyctocoris is concerned, there is no doubt in my mind that it is campestris.” Both China and Herring were very experi- enced hemipterists with enormous collec- tions at their disposal and their consider- ations deserve attention. The illustration of Kirkaldy’s type by Zimmerman represents a specimen of Lyctocoris campestris. Based upon this evidence, I considered Nesidioch- eilus hawatiensis Kirkaldy 1901 to be a synonym of Acanthia campestris Fabricius 1794 [now known as Lyctocoris campestris (Fabricius)], New synonymy. The collec- tion of this species at higher elevations on Hawai and Maui reflects the distribution of a temperate species. The specimens from Hawaii were taken with specimens of a mealy bug, a likely prey. The bug species should be looked for on other islands at higher elevations. I have placed an adult of Lyctocoris campestris for reference in the collections of the Bishop Museum. Acknowledgments.—My thanks to A. Ramsdale, Bishop Museum for loan of specimens under this direction, a reviewer and to L. Parks for manuscript preparation. LITERATURE CITED Beardsley, J. W. 1981. Lyctocoris campestris reported from Mauna Kea, Hawaii 7,800’. Proceedings, Hawaiian Entomological Society 23(3): 327. Champion, G. C. 1900. Insecta. Rhynchota. Hemip- tera: Heteroptera, Vol. II. /n Biologica Centrali - Americana. London, xvi + 416 pp. Chu, Y. I. 1969. On the bionomics of Lyctocoris be- neficus (Hiura) and Xylocoris galactinus (Fieber) (Anthocoridae, Heteroptera). Journal of the Fac- ulty of Agriculture, Kyushu University 15: 1-136. Fabricius, J. C. 1794. Entomologia systematica emen- data et aucta, secundum classes, ordines, genera, species adjectis synonymis, locus, observationi- bus, descriptionibus. Proft, Hafinae. 4:1-v, 1—472. Ford, L. J. 1979. The phylogeny and biogeography of the Cimicoidea (Insecta: Hemiptera). M.S. De- gree, University of Connecticut vii: 1-138 pp. Henry, T. J. 1988. Family Anthocoridae, pp. 12-28. In Henry, T. J. and R. C. Froeschner, eds. Catalog of the Heteropera, or true bugs, of Canada and the con- tinental United States. E.J. Brill, Leiden, 958 pp. Hiura, I. 1966. Contribution to the knowledge of An- thocoridae from Japan and its adjacent territories (Hemiptera—Heteroptera). 3. Bulletin of the Osa- ka Museum of Natural History No. 19: 29-37. Kirkaldy, G. W. 1902. Hemiptera. Fauna Hawaiiensis Ill (1): 93-174. Pl. 4, 5. Péricart, J. 1972. Hémiptéres Anthocoridae, Cimicidae et Microphysidae de I’ Ouest—paléarctique. Jn Faune de l'Europe et du bassin méditerranéen. 7: i-iv, 1-401. Masson et Cie Editeurs. Paris. . 1996. Family Anthocoridae Fieber, 1836. pp. 108-140. Jn Aukema, B. and C. Rieger, eds. Cat- alogue of the Heteroptera of the Palearctic Region, Vol. 2. Cimicomorpha 1. 361 pp. Zimmerman, E. C. 1948. Insects of Hawaii, Vol. 3. Heteroptera. University of Hawaii Press, Hono- lulu, 255 pp. John D. Lattin, Department of Botany and Plant Pathology, Oregon State University, Corvallis, OR 97331-2902, U.S.A. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 241-242 NOTE Tetraphleps uniformis Parshley, 1920, a Synonym of Tetraphleps canadensis Provancher, 1886, and Tetraphleps furvus Van Duzee, 1921, Restored to Species Status (Hemiptera: Heteroptera: Anthocoridae) The anthocorid genus Tetraphleps Fieber is Holarctic with six species in the Palearc- tic and five species in the Nearctic Region @enicart 1972 1996: Henry’ 1988). ‘All known species are associated with conifers. Tetraphleps canadensis was described by Provancher (1886), based on a female from Cap Rouge, Quebec. The original descrip- tion was brief, based largely on color. The precise status of this species remained un- certain for many years (Parshley 1920, Van Duzee 1921, Blatchley 1926, Torre-Bueno 1930). Drake and Harris (1928) redescribed the type of 7. canadensis and indicated that the rostrum extended to the middle of the mesosternum, but also stated that the ros- trum of the holotype was missing, appar- ently basing their observation on the holo- type female of Tetraphleps americana Par- shley from Peaks Island, Maine. This agrees with Parshley’s (1920) statement that the rostrum of his species ‘“‘extended well beyond front coxa. ...’’ Drake and Harris (1928) also erred in considering a male of T. canadensis from Nordegg, Alberta, col- lected by J. McDunnough on larch on Au- gust 4, 1921—long after the date of the original description as the allotype. Knight (1925) had stated that specimens of Tetra- phleps americana trom Nordegg agreed perfectly with the original description by Parshley except for very small difference in length of antennal segment II. Based on this information, Drake and Harris (1928) syn- onymized 7. americana with T. canadensis. Subsequently, Kelton and Anderson (1962) placed 7. osborni Drake, 1923 from New York and 7. edacis Drake and Harris, 1926 from New York as synonyms of Tetra- phleps canadensis. Tetraphleps uniformis Parshley (1920) was described from Mt. Washington, New Hampshire, based upon the unique female holotype. The original description followed that of Tetraphleps americana and was quite brief, based chiefly on slight color dif- ferences of the dorsum. It was virtually identical in length to 7. americana (3.68 mm to 3.67 mm). For reasons not stated, T. uniformis was recognized as a distinct spe- cies over the following years by Van Duzee (1921), Blatchley (1926), and Torre-Bueno (1930). All expressed concern about distin- guishing this species from 7. canadensis. Kelton and Anderson (1962), Kelton (1966) and Henry (1988) continued to regard 7. uniformis as a distinct species without com- ment. All of the synonyms of 7. canadensis occur within a short distance of the holo- type locality. Tetraphleps uniformis Par- shley, 1920 is considered a synonym of Te- traphleps canadensis Provancher, 1886, new synonymy. Tetraphleps furvus Van Duzee, 1921 was described from Ward, Colorado. Kelton and Anderson (1962) regarded it as a valid spe- cies, but later Kelton (1966) placed T. fur- vus into synonymy with 7. uniformis Par- shley, 1920, here considered a synonym of Tetraphleps canadensis. My study of the holotpe indicates that 7. furvus is distinct from T. canadensis. The rostrum of T. fur- vus reaches only the anterior coxae, where- as the rostrum reaches at least the middle of the mesosternum or beyond in 7. unifor- mis. Additional characters include the trans- verse, nearly rectilinear pronotum, the scarcely elevated, closely, obscurely punc- tate callosities; anteriorly dark pronotum, the highly polished hemelytra, with fine dis- tant, obscure, testaceous brown punctures, paler embolium, and membrane with trans- 242 verse row of pale marks at apex of corium. As a consequence, 7. furvus is restored to species status, new status. Acknowledgments.—My thanks to R.W. Baumann of Brigham Young University for the loan of critical specimens of Tetra- phleps a careful reviewer, and L. Parks for careful attention to the manuscript. LITERATURE CITED Blatchley, W. S. 1926. Heteroptera of eastern North America. Nature Publishing Company, Indianap- olis, 1,116 pp. Drake, C. J. 1923. Heteroptera in the vicinity of Cran- berry Lake. New York State College of Forestry, Syracuse University. Technical Publication No. 16. 22: 54-86. Drake, C. J. and H. M. Harris. 1926. Notes on Amer- ican Anthocoridae with description of new forms. Proceedings of the Biological Society of Wash- ington 39: 33—46. Drake, C. J. and H. M. Harris. 1928. Tetraphleps can- adensis Provancher, a true Tetraphleps (Hemip.). The Canadian Entomologist 60: 50. Henry, T. J. 1988. Family Anthocoridae Fieber, 1837, pp. 22-28. Jn Henry, T. J. and R. C. Froeschner, eds. Catalog of the Hetereoptera, or true bugs, of Canada and the continental United States. Brill, Leiden, 958 pp. Kelton, L. A. 1966. Synopsis of the genus Tetraphleps Fieber in North America (Hemiptera: Anthocori- dae). The Canadian Entomologist 98: 199-204. Kelton, L. A. and N. H. Anderson. 1962. New Antho- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON coridae from North America, with notes on the status of some genera and species (Heteroptera). The Canadian Entomologist 94: 1302-1309. Knight, H. H. 1925. A list of Miridae and Anthocori- dae from Alberta, Canada (Hemiptera). The Ca- nadian Entomologist 57: 181—182. Parshley, H. M. 1920. Hemiptera from Peaks Island, Maine, collected by Mr. G. A. Moore. The Ca- nadian Entomologist 52: 80-87. Péricart, J. 1972. Hémiptéres. Anthocoridae, Cimicidae et Microphysidae de |’ OQuest—Paléarctique, Faune de |’Europe et du Bassin Méditerranéen. Mason et Cie, Paris. 7: 1—402. . 1996. Family Anthocoridae, pp. 108-140. In Aukema, B. and C. Reiger, eds. Catalogue of the Heteroptera of the Palearctic Region, Vol. 2. Cim- icomorph 1. The Netherlands Entomological So- ciety, Amsterdam. Provancher, L. 1885-1890. Petite Faune Entomolo- gique du Canada et Particulierement de la Prov- ince de Quebec, Vol. 3. Cinquieme Ordre les Hémipteres. Naturaliste Canadien, 1885, 1—64; 1886, 65-112; 1887, 113-184; 1888, 185-204; 1889, 205-282; 1890, 283-354, plates 1—5. Torre-Bueno, J. R. de la. 1930. Records of Anthocor- idae particularly from New York. Bulletin of the Brooklyn Entomological Society 25: 11—20. Van Duzee, E. P. 1921. Characters of eight new species of North American Anthocoridae or flower bugs. Proceedings of the California Academy of Scienc- es 11: 137-144. John D. Lattin, Department of Botany and Plant Pathology, Oregon State Univer- sity, Corvallis, OR 97331-2902, U.S.A. PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 243-247 NOTE Introduced Leaf Beetles of the Maritime Provinces, |: Sphaeroderma testaceum (F.) (Coleoptera: Chrysomelidae) Sphaeroderma testaceum (FE), reported from three localities in Nova Scotia by Hoebeke and Wheeler (2003), is the most recent of a long series of Old World beetles that have been introduced to North America through Nova Scotia and Atlantic Canada. Seaports offer many opportunities for such introductions and this region has a long his- tory of commercial traffic. Some introduced species have spread little, whereas others have used their foothold in Atlantic Canada as a springboard to spread to other portions of the continent. In some cases it is clear Where and when the introductions were made; in other instances, specific evidence is lacking or is speculative (Majka and Kli- maszewski 2004). In 1997 (August 1, August 26, and Sep- tember 17) while sweep-netting in an old field adjacent to the Nova Scotia Depart- ment of Natural Resources Insectary in Shubenacadie, Colchester County, Nova Scotia, Jeffrey Ogden collected 10 speci- mens of what proved on subsequent ex- amination to be Sphaeroderma testaceum (Nova Scotia Department of Natural Re- sources Insectary, Shubenacadie, Nova Scotia, Canada) (Fig. 1). Hoebeke and Wheeler (2003) reported the species on Canada thistle (Cirsium arvense (L.) Sco- poli) from collections made in 2001 and 2003 at Bible Hill, Antigonish, and New Glasgow in Nova Scotia (Cornell Univer- sity Insect Collection, Ithaca, New York, USA). C. G. Majka subsequently collected S. testaceum along Highway 104, 3 km southeast of Kemptown, Colchester County (August 20, 2002) and near Pratts Lake, Colchester County (July 23, 2004). In 2005 more extensive sweep-netting surveys were undertaken and specimens were collected from Milford Station (July 26, 2005), Elms- dale (August 22, 2005), and Enfield (Au- gust 22, 2005) all in Hants County, and from Bedford, Halifax County (August 22, 2005) (C. G. Majka Collection, Halifax, Nova Scotia, Canada). Thistle stands at oth- er sites in Nova Scotia, New Brunswick, and Prince Edward Island were also sur- veyed but no specimens of S. testaceum were found (Fig. 2). All the specimens collected have been on Canada thistle. Stands of Bull thistle (Cir- sium vulgare (Savi) Tenore) and Swamp thistle (C. muticum Michx.) were also swept but to date S. testaceum has not been found on either plant. Known host plants of Sphaeroderma are Carduus, Cirsium, On- opordum, Carlina, Serratula, and Cynara (Clark et al. 2004). Of these, Carduus cris- pus L, Cirsium vulgare, C. muticum, C. pal- ustre (L.) Scop., and C. arvense occur in Nova Scotia (Roland 1998). Individuals were observed moving on the stalks of Cir- sium. When disturbed, they would spring weakly, retract their legs, and feign death. The only other beetle found to be consis- tently associated with S. testaceum on this host was Cassida rubiginosa Miiller, anoth- er adventive chrysomelid which also feeds on Cirsium. At two sites (Enfield and Elmsdale) quantitative sampling was carried out to es- tablish an index of S. testaceum abundance. Areas of thistle stands were measured and carefully swept. While not all specimens present would necessarily have been cap- tured, the approach establishes a minimum level of abundance. At Enfield 0.24 indi- viduals/m? were found while at Elmsdale abundance was 0.22 individuals/m~?. The question arises as to how this species made its way to Nova Scotia. Cirsium ar- vense is Eurasian in origin and has been PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Rion Habitus photograph of Sphaeroderma testaceum. VOLUME 108, NUMBER 1 New Brunswick Fig. 2. found. Open circles indicate localities were thistle stands were examined for S. testaceum but no specimens were found. deliberately or inadvertently introduced to North America. Sphaeroderma testaceum would appear to have been introduced to the province in association with its impor- tation. Cirsium arvense is considered one of the world’s worst weeds (Holm et al. 1977, McClay et al. 2002) and is a serious pest in Nova Scotia where it is found throughout the province (Roland 1998). In this context, Hoebeke and Wheeler (2003) raised the possible biocontrol potential of S. testa- ceum. To date S. testaceum has only been found along major transportation corridors. The line of points plotted in Fig. 2 follows Highway 102 (the main arterial route lead- ing from Halifax as well as the Canadian National Railways line leading from the 245 oS 7 (3) Prince Edward Island Kilometres 50 ¢} SIGAIUE 50 100 Kilometres 4 Miles 30 ie) 30 ; 60 Miles j Distribution of Sphaeroderma testaceum. Closed circles indicate localities where the species has been port of Halifax) from Bedford Basin (at the head of Halifax Harbour) to Truro and thence westward along the Highway 104 (the trans-Canada Highway) to Antigonish. Every thistle stand examined in this corri- dor yielded specimens of S. festaceum. Eastward from Truro along Highway 104, and in other parts of Nova Scotia, New Brunswick, and Prince Edward Island, no specimens have been found. At present S. testaceum would appear to be confined to an area in the north-central mainland of the province. The pattern of distribution might be indicative of an introduction through the port of Halifax with a subsequent spread along the Highway 102 and 104 transpor- tation corridors. Majka and Klimaszewski (2004) discuss eleven species of adventive 246 beetles known to have been introduced to North America through the port of Halifax. Species such as Meligethes viridescens (F.) have subsequently spread from there to a large number of other localities in Nova Scotia and Prince Edward Island (Mason et al. 2003). This introduction appears to have taken place relatively recently. Since 1986 staff of the Nova Scotia Department of Natural Re- sources Insectary (based in Shubenacadie, NS) have regularly collected and monitored Coleoptera populations in Colchester, An- tigonish, Hants, and Pictou counties, in- cluding some of the locations where S. tes- taceum has been found, yet no specimens of S. testaceum were found prior to 1997. Thus, the recent appearance of this species and the current extent of its distribution in the province raise the possibility that it is rapidly expanding its range along traffic corridors where Cirsium arvense grows. Spheroderma testaceum is a recent ad- dition to the lengthy catalogue of species introduced to the continent through Atlantic Canada. The beetle fauna of Nova Scotia, in particular, includes 329 introduced spe- cies, or 14.6% of its total beetle fauna in- cluding 23 species of Chrysomelidae (C. Majka, unpublished data). While some, such as S. testaceum, Chrysolina hyperici (Forster), and Aphthona cyparissiae (Koch) are considered to be beneficial species, oth- ers such as Pyrrhalta viburni (Paykull), Lil- ioceris lilii (Scopoli), Crioceris duodecim- punctata (L.), C. asparagi (L.), Oulema melanopus (L.), and Meligethes viridescens (FE) are potentially serious pests. Although shipping and inspection practices now are more rigorous than they were prior to 1965 when quarantine restrictions on soil and plant material were imposed (Spence and Spence 1988), such introductions continue because exchanges and importations are considerably frequent than in the past. In Nova Scotia, recent programs by the Canadian Food Inspection Agency (CFIA) to exterminate the introduced Tetropium fuscum (FE) and Popillia japonica Newman PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON have cost millions of dollars, generated tre- mendous public controversy, and signifi- cantly disrupted forestry and horticultural practices (LeBlanc 2002, Majka and Kli- maszewski 2004). Continued vigilance 1s warranted lest other such inadvertent intro- ductions result in unplanned environmental consequences. Acknowledgments.—Sincere thanks to Peter Harris (Agriculture and Agri-Food Canada, Lethbridge), Jeffrey Ogden (Nova Scotia Department of Natural Resources), and M. Glen Sampson (Nova Scotia Agri- cultural College) for their assistance. Two anonymous reviewers read an earlier draft of paper and made many constructive sug- gestions. The first author thanks his col- leagues at the Nova Scotia Museum, Calum Ewing and Andrew Hebda, for continuing support and encouragement. This work has been assisted by a research grant from the Nova Scotia Museum of Natural History. LITERATURE CITED Clark, S: M., DIG, KeDoux, TN: Seeno; E> G2 Riley; A. L. Gilbert, and J. M. Sullivan. 2004. Host Plants of Leaf Beetle Species Occurring in the United States and Canada. Coleopterists Society, Special Publication No. 2. Athens, Georgia, 476 PPp- Hoebeke, E. R. and A. G. Wheeler Jr. 2003. Sphae- roderma testaceum (F.) (Coleoptera: Chrysomeli- dae), a Palearctic flea beetle new to North Amer- ica. Proceedings of the Entomological Society of Washington 105: 990—994. Holm, L. G., D. L. Plucknett, J. V. Pancho, and J. P. Herberger. 1977. The World’s Worst Weeds: Dis- tribution and Biology. University of Hawaii Press, Honolulu, 609 pp. LeBlanc, N. 2002. CFIA program to eradicate Japa- nese beetle in Halifax. Canadian Food Inspection Agency news release, March 25, 2002. Available from __ http://www.inspection.gc.ca/english/corpattr/ newcom/2002/20020325e.shtml (accessed 19 Au- gust 2005) Majka, C. G. and J. Klimaszewski. 2004. Phloeocharis subtilissima Mannerheim (Staphylinidae: Phloeo- charinae) and Cephennium gallicum Ganglbauer (Scydmaenidae) new to North America: a case study in the introduction of exotic Coleoptera to the port of Halifax, with new records of other spe- cies. Zootaxa 781: 1-15. Mason, P. G., O. Olfert, L. Sluchinski, R. M. Weiss, C. Boudreault, M. Grossrieder, and U. Kuhlmann. VOLUME 108, NUMBER 1 2003. Actual and potential distribution of an in- vasive canola pest, \ Meligethes viridescens (Co- leoptera: Nitidulidae), in Canada. Canadian En- tomologist 135: 405-413. McClay, A. S., R. S..Bourchier, R. A. Butts, and D. P. Peschken. 2002. Cirsium arvense (L.) Scopoli, Canada thistle (Asteraceae), pp. 318—330. Jn Ma- son, P. G. and J. T. Huber, eds. Biological Control Programmes in Canada, 1981—2000. CABI Pub- lications, Wallingford, U.K. Roland, A. E. 1998. Roland’s Flora of Nova Scotia (revised by M. Zinck). Nimbus Press and Nova Scotia Museum, Halifax, Nova Scotia, 1,297 pp. Spence, J. R. and D. H. Spence. 1988. Of ground- beetles and men: introduced species and the syn- 247 anthropic fauna of western Canada. Memoirs of the Entomological Society of Canada 144: 151— 168. Christopher G. Majka and Laurent Le- Sage. (CGM) Nova Scotia Museum of Nat- ural History, 1747 Summer Street, Halifax, Nova Scotia, Canada B3H 3A6 (e-mail: c.majka@ns.sympatico.ca); (LS) Agricul- ture and Agri-Food Canada, ECORC, K.W. Neatby Bldg., 960 Carling Ave., Ottawa, Ontario, Canada KIA OC6_ (e-mail: lesagel@agr.gc.ca) PROC. ENTOMOL. SOC. WASH. 108(1), 2006, p. 248 NOTE Rediscovery of Fallceon eatoni (Kimmins) (Ephemeroptera: Baetidae) Certain adult small minnow mayflies (Baetidae) from the 19 Century Godman and Salvin collections made in Mexico and Central America (now residing in the Brit- ish Museum) were one of three species re- ported by Eaton (1892) as new but un- named species of Baetis Leach. This small series from northern Sonora was restudied by Kimmins (1934) and described as B. ea- toni Kimmins. Waltz and McCafferty (1987) recombined this species with the ge- nus Fallceon Waltz and McCafferty based on its distinctive male genitalia. Although the F. eatoni had later been deemed syn- onymous with the ubiquitous F. quilleri (Dodds) (McCafferty and Waltz 1991), my subsequent examination of type material showed that the hindwing costal process was not hooked as in F. quilleri, and that the terminal segment of the genital forceps was more developed than in the latter spe- cies. Thus, F. eatoni was revalidated (McCafferty and Lugo-Ortiz 1994), but it’s continued existence has been questionable because it has not been known since its original collection well over 100 years ago. Among unidentified materials from Salt River Canyon in Arizona that were recently collected and donated by Will Reeves (At- lanta), | found a single specimen of Baeti- dae that proved to be F. eatoni (Arizona, Gila Co., Salt River Canyon, IV-18-2005, W. K. Reeves, deposited in the Purdue En- tomological Research Collection). The re- discovery of this species is significant in that it not only establishes the species’ con- tinued existence, but it provides first USA and Arizona records. Because of possible low numbers of populations and restricted distribution, it may be considered a species of environmental concern. LITERATURE CITED Eaton, A. E. 1982. Biologia Centrali-Americana: In- secta, Ephemeridae. Vol. 38. Bernard Quaritch, London. Kimmins, D. E. 1934. Notes on the Ephemeroptera of the Godman and Salvin Collection, with descrip- tions of two new species. Annals and Magazine of Natural History (10) 14: 338-353. McCafferty, W. P. and C. R. Lugo-Ortiz. 1994. Taxo- nomic status of three species of Fallceon (Ephem- eropta: Baetidae). Entomological News 105: 161— 163. McCafferty, W. P. and R. D. Waltz. 1991. Revisionary synopsis of the Baetidae (Ephemeroptera) of North and Midddle America. Transactions of the American Entomological Society 116: 769-799. Waltz, R. D. and W. P. McCafferty. 1987. New genera of Baetidae from some Nearctic species previous- ly included in Baetis Leach (Ephemeroptera). An- nals of the Entomological Society of America 80: 667-680. W. P. McCafferty, Department of Ento- mology, Purdue University, West Lafayette, IN 47907, U.S.A. (e-mail: meccaffer@ purdue.edu) PROC. ENTOMOL. SOC. WASH. 108(1), 2006, p. 249 NOTE A Replacement Name for Lepidostoma weaveri Bueno, Santiago, and Barba (Trichoptera: Lepidostomatidae) In a recent paper on Trichoptera collected in Mexico and Panama (Bueno-Soria et al. 2004:481), we described the new species Lepidostoma weaveri, unfortunately over- looking the fact that Harris (1986:36) ear- lier had used weaveri for a species of Lep- idostoma from Alabama. Lepidostoma weaveri Bueno, Santiago, and Barba, 2004, is thus a junior primary homonym and must be renamed. We here propose Lepidostoma textor Bueno, Santiago, and Barba, new name, for Lepidostoma weaveri Bueno, Santiago, ad Barba, 2004. The specific ep- ithet is Latin for weaver. It is a noun in apposition in honor of John Weaver. LITERATURE CITED Bueno-Soria, J., S. Santiago-Fragoso, and R. Barba- Alvarez. 2004. More new Trichoptera from Mex- ico and Panama. Transactions of the American En- tomlogical Society 130: 479-486. Harris, S. C. 1986. New species of caddisflies (Tri- choptera) from Alabama. Proceedings of the En- tomological Society of Washington 88: 30—41. Joaquin Bueno-Soria, Silvia Santiago- Fragoso, and R. Barba-Alvarez, Instituto de Biologia, UNAM, Cuidad Universitaria, Mexico, D.F. 04510, Mexico (e-mail: bueno@ servidor.unam.mx) PROC. ENTOMOL. SOC. WASH. 108(1), 2006, pp. 250-251 BOOK REVIEW Carpenter Ants of the United States and Canada. 2005. Laurel D. Hansen and John H. Klotz; Comstock Publishing As- sociates, Ithaca, NY. 204. pp. ISBN # 0- 8014-4262-1 Cost: $35.00. Ants are one of nature’s great evolution- ary success stories. Since emerging around 130 million years ago, ants have achieved unrivaled ecological dominance in most ter- restrial ecosystems—a dominance that has remained unabated since the Eocene. With nearly 12,000 described species (with an es- timated 20,000 species in the family), ants are a moderately species rich insect group. One particular interesting aspect of this di- versity is that the number of ant species is not evenly distributed across the family. In fact, analysis of species diversity across the nearly 300 ant genera reveals a stunning taxonomic lop-sidedness to generic-level species richness patterns. While the species richness of genera is quite variable, from monotypic (single species) to some contain- ing a few hundred species, only two ant genera could be characterized as hyperdiv- erse (sensu Wilson, 2003): Camponotus and Pheidole. These two genera combined ac- count for over 25% of all ant species, each containing about 13% of the world’s ant fauna. Camponotus and Pheidole are not only diverse, but they are also among the most ecologically dominant ants on earth. Anywhere ants are found, you will find Camponotus and Pheidole species. One morphological characteristic that both Camponotus and Pheidole share is that they are characterized by the presence of a major (sometimes called a soldier)/minor worker caste system. Camponotus is more properly described as polymorphic with typically several sizes from minor to major in between the two extremes in worker size. This may in part account for their tremen- dous species diversity (Wilson 2003). The New World Pheidole have recently been summarized by Wilson (2003), but there is as of yet, no comparably large synthetic work for Camponotus. This is unfortunate because the range of morphological, behav- ioral, and ecological variation among Cam- ponotus ants is truly impressive. One Cam- ponotus species, C. gigas of southeastern Asia, is among the largest ants on earth, with the major measuring up to 33 mm in total length. This species may in fact be the largest ant on earth, but just who exactly is the reigning giant among ants is debated by myrmecologists who have different ways of measuring such things. Camponotus are ex- pressed in a dazzling array of morphologi- cal shapes and colors, from flat-headed forms (e.g., subgenus Colobopsis) that plug-up nest entrances to stop invaders (usually other ants) to species with beauti- ful gold and silver gasters (e.g., C. detritus) that sparkle in sunlight. There are even some Camponotus (e.g., cylindricus com- plex) that literally explode upon being dis- turbed! It is against this backdrop that Laurel Hansen and John Klotz offer us a peek into the fascinating world of Camponotus ants. Although insect enthusiasts might have ap- preciated a more rounded survey of Cam- ponotus, the decision by Hansen and Klotz to focus on the Camponotus species that nest in wood (‘‘carpenter ants’’), which in- cidentally applies to about half of the known Camponotus from the U.S. and Can- ada (they list 24 out of 50 species as “‘car- penter ants’’), provides us with an extensive review of one of the most conspicuous and frequently encountered groups of North American ants. The volume joins a growing number of taxon-based ant books. Hansen and Klotz should be commended for their efforts because such books convey in an en- gaging style the world of ants to an audi- ence far larger then specialized research ar- ticles do. More taxon-based volumes, de- VOLUME 108, NUMBER | signed for the astute naturalist and specialist alike are much needed. Hansen and Klotz summarize the taxo- nomic status of Camponotus from the U.S. and Canada. Our taxonomic understanding of these particular Camponotus is far more complete than those from other regions of the world. An illustrated dichotomous key is provided for the species that are found “in and around structures.”’ A key that in- cluded all U.S. and Canadian Camponotus species would have been preferred, as this would have added value for people inter- ested in non-pestiferous species as well, but otherwise the key is well done. What fol- lows is an overview for each carpenter ant species, including a distribution map. The summary of ecology, life history and for- aging behavior of carpenter ants is compre- hensive and provides a good sense for the state of the science. That most of what is known comes’ from only a handful of spe- cies (C. modoc, for example is particularly well-documented) illustrates how little is known of the natural history and ecology of Camponotus ants, even among the eco- nomically important carpenter ants. The chapter on Camponotus morphology is especially well presented and valuable. Hansen and Klotz provide many well-illus- trated line drawings of both external and internal anatomy. The inclusion of male morphology is important, because this caste is often neglected in ant publications and excellent illustrations of the genitalia are provided. I have one small caveat. The au- i) Nn thors write that wing venation of Campon- otus ants can be used to separate them from other ant genera. They do not elaborate or provide a citation of this statement, and as far as | am aware Camponotus wing vena- tion is typical of most formicine ants and not useful for diagnostic purposes. Because the book is geared toward the “carpenter ants” of Camponotus, a chapter is provided dealing with the pestiferous as- pects and control of these ants. Hansen and Klotz provide the exterminator, pest man- ager, and concerned homeowner alike a readable overview of the methods that can be employed to control carpenter ants. My biggest criticism of the book stems from something I am not sure was even un- der the authors’ control. The text is richly illustrated with black and white photo- graphs, but the quality of many of these pic- tures could have been improved. Often- times the pictures are too dark to effectively see what is being portrayed. That criticism aside, Carpenter Ants of the United States and Canada will be a useful addition to any entomological library. LITERATURE CITED Wilson, E.O. 2003. Pheidole in the New World. Har- vard University Press, Cambridge, Massachusetts, 793 pp. John S. LaPolla, Department of Ento- mology, National Museum of Natural His- tory, Smithsonian Institution, P.O. Box 37012, MRC 188, Washington DC 20013- 7012, U.S.A. (e-mail: lapollaj@ si.edu) PROC. ENTOMOL. SOC. WASH. 108(1), 2006, p. 252 ENTOMOLOGICAL SOCIETY OF WASHINGTON SCHEDULE OF REGULAR MEETINGS FOR 2005-2006 October 6. R. Toby Schuh, American Museum of Natural History, NY. Exploring the Southern Hemisphere: Plant Bug Planetary Biodiversity Field Work in Australia and South Africa. November 3. Gary Hevel, Department of Entomology, Smithsonian Institution. Backyard Entomology. December |. Desmond Foley, NRC Research Associate, Walter Reed Biosystematics Unit. The Curious Case of the Ant & the Mosquito and Other Tales of Culicid Complexity. January 5. Jason P. W. Hall, Department of Entomology, Smithsonian Institution. Historical and Ecological Biogeography of the Napaeina Riodinid Butterflies. February 2. Diana Percy, National Zoological Park, Smithsonian Institution. Psyllids: Diversification in Islands, Host Plant Interactions and Acoustic Communication. March 2. Wayne FE Wehling. APHIS, PPQ. Systematist or Collector: APHIS and Permits for Live Arthropods. April 6. Charyn Micheli, Center for Systematic Entomology, University of Maryland. The Longhorned Woodboring Beetles of Puerto Rico and the Dominican Republic. May 4. Cole Gilbert, Department of Entomology, Cornell University. All the Better to See You with My Dear: Visually-Guided Aerial Pursuit of Females by Male Flesh Flies. Meetings at: 7:00 PM Cathy Kerby Seminar Room (CE-340) National Museum of Natural History 10th & Constitution Ave., Washington, DC Meetings Open to the Public Regular Pre-Meeting Dinner: R. Reagan Building Food Court, 5:30 PM 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. SERN ES eck ERS ies RED Bip LEE ae a ae ae, SRN PR AN A CO CON A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. 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CONTENTS wii NG KINKOROVA, JUDITA and MICHAELA MiCKOVA— Life history of the European tephritid fly Chaetostomella cylindrica (Diptera: Tephritidae) and its possible use as a biocontrol agent inthe USA: 9.0.0 ils seccences cecesetenecpnsueumenede soem che Be ae te Me carar Va ghana? se wae ancee 2 chapee Ts Reza Sil KIREJTSHUK, ALEXANDER G. and GEORGE POINAR, JR.—Haplochelidae, a new family of Cretaceous beetles (Coleoptera: Myxophaga) from Burmese amber ..............:0sseeeeeee 155 MAIER, CHRIS T. and CHRISTER HANSSON—Palearctic Sympiesis acalle and Sympiesis gordius (Hymenoptera: Eulophidae) in North America: Taxonomic changes and a review of Nearctic host: records), ..2..Ch cute ieccsstab ates ot peste aees Ane cainLn sue meu rN een e th besue Aa deanades duaseeansee tata ad 14 ROBBINS, ROBERT K. and MARCELO DUARTE—Systematic placement of Lycaena cogina Schaus (Lepidoptera: Lycaenidae: Polyommatinae), a biogeographically disjunct New World SPE CUES Here sua e le seurbir- tote pliog she teeen Care cams SP PER PI tc We ere Seba ith Mea, MEL Te Ae as. 226 RONG, BAO and XINLI WANG—Two new species of Myrmeleon Linnaeus, 1767 (Neuroptera: Myrmeleontidae) from China, with a key to Chinese Species .........:..ccsesesesesseseseesenersenteensesenes 125 SCARBROUGH, AUBREY G.—Two species of Clinopogon Bezzi (Diptera: Asilidae: Stenopogoninae) from Sabah, Malaysia ..............c::ccecesceeseeeseetsenseerecesecessenscesaesnesnaceseeesenreeasenas 92 SMITH, DAVID R.—Review of the cypress and juniper sawflies of the genus Susana Rohwer and Middleton (Hymenoptera: Tenthredinidae) .............:cccccesccesceeeeceneeeeeceseneneceeaseseersccesesessasnens 62 TOGASHI, ICHIJI—A new sawfly, Fagineura quercivora (Hymenoptera: Tenthredinidae) feeding on Quercus serrata and QO. mongolica crispula in Honshu, Japan ........:...scsseeesceseereenereeeesenes 169 WAN, XIA, XINLI WANG, and XINGKE YANG—Study on the genus Layahima Navas (Neuroptera: Myrmeleontidae) from China ...........c:cccccecceseceseeereeteeneetseesenseeeesessenssensersnensecsaans 35 WHEELER, A. G., JR. and THOMAS J. HENRY—Gampsocoris decorus (Uhler) and Metacanthus tenellus Stal (Hemiptera: Berytidae): Neotropical stilt bugs as colonists of an African grass, Urochloa mutica (Poaceae), in Florida, with a review of berytid-grass associations ..........-....+.. 1 ZACK, RICHARD S., CHRISTOPHOR N. LOONEY, and ROBERT D. GORDON—A new species of flightless, winter-active dung beetle, Stenotothorax woodleyi (Coleoptera: Scarabaeidae), associated with shrub-steppe habitat in Washington State ............:eeseeereeee 45 ZACK, RICHARD S., DAVID E. RUITER, DENNIS L. STRENGE, and PETER J. LANDOLT— Adult caddisfly (Trichoptera) phenology at the Hanford Reach National Monument, Washington State 12.2.5) Ne Bice ola. ca ad hae ite claconausoetee saadosmemsc gone hebhis 0s eten ins enate gaara Oa oxe setae a 131 ZHANG, YALIN, YINGLUN WANG, and YANLI CHE—A new genus of the subfamily Hemisphaeriinae (Hemiptera: Fulgoroidea: Issidae) from Chima ............:eeceescerseesceeseteseeeeeees 165 NOTES ASCHER, JOHN S., PARKER GAMBINO, and SAM DROEGE—Adventive Hylaeus (Spatulariella Popov) in the New World (Hymenotpera: Apoidea: Colletidae) BUENO-SORIA, JOAQUIN, SILVIA SANTIAGO-FRAGOS, and R. BARBA-ALVAREZ—A replacement name for Lepidostoma weaveri Bueno, Santiago, and Barba (Trichoptera: Lepidostomatidae) LATTIN, JOHN D.—Nesidiocheilus hawaiiensis Kirkaldy, 1902, a synonym of Acanthia cam- pestris Fabricius, 1794 (Hemiptera: Heteroptera: Lyctocoridae) LATTIN, JOHN D.—Tetraphleps uniformis Parshley, 1920, a synonym of Tetraphleps canadensis Provancher, 1886, and Tetraphleps furvus Van Duzee, 1921, restored to species status (Hemiptera: Heteroptera: Anthocoridae) MAJKA, CHRISTOPHER G. and LAURENT LESAGE—Introduced leaf beetles of the Maritime Provinces, 1: Sphaeroderma testaceum (F.) (Coleoptera: Chrysomelidae) McCAFFERTY, W. P.—Rediscovery of Fallceon eatoni (Kimmins) (Ephemeroptera: Baetidae) BOOK REVIEW LaPOLLA, JOHN S.—Carpenter Ants of the United States and Canada, by Laurel D. Hansen and John H. Klotz MISCELLANEOUS Regular Meetings for 2005—2006 VOL. 108 APRIL 2006 NO. 2> OL (ISSN 0013-8797) LfGl 0.2). When the moth rotates clockwise, it anchors its right foreleg, and when it rotates counterclockwise, it anchors its left foreleg. The leg that is used as an anchor moves very litthe compared to the other legs, only shifting it slightly when the moth makes large circles, changes direc- tion, or moves to another region of the leaf. In general, dancing takes place over the en- tire leaf, but at times can be localized to certain areas (Fig. 9). Each antenna is held PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON out laterally while the moth dances, and in one case, a moth was seen rotating slowly around a water droplet with its proboscis extended to the droplet (Fig. 10). When dis- turbed, the moth either stops dancing or flies from the leaf. The average time spent dancing was 71.14 seconds (SD = 36.7, n = 24) and dancing speed varied. The behavioral significance of dancing in Beltheca oni and other microlepidoptera largely remains unknown. The adult of some Brenthia sp. (Choreutidae) move in a jerky fashion after alighting (Fletcher 1920, Forbes 1923, Rota 2003, Aiello and Becker 2004), and the display behavior may mimic the movement of salticid jumping spiders (Robinson et al. 1994, Rota 2003, Aiello and Becker 2004). Dancing in Gelechtidae, Momphidae, Cosmopterigidae, and Tinei- dae is quite different, as the moth typically rotates rapidly without jerky movements. Observations of B. oni suggest that the adult dances during courtship. Typically only one moth was seen on a leaf, but there were three instances where two moths were dancing on the same leaf, two of which in- volved a male and a female. In one case, as soon as one of the sexes began dancing, the other danced also, and several seconds after one stopped, the other also stopped. This ‘““dance-stop”” scenario was repeated four times, and in each instance, dancing lasted 4-5 seconds before one of the moths stopped. Morrison (1968) described a sim- ilar scenario in Dryadaula pactoila Meyrick (Tineidae). He observed a male in captivity running around in circles beside a female while fanning his wings, and repeatedly dancing and stopping until the pair mated. Unfortunately we were able to witness dance-stop behavior between only two in- dividuals, and the pair flew away before the outcome of the scenario could be observed. Additional observations are necessary to conclusively determine that this dance-stop behavior is associated with courtship in mi- crolepidoptera. The moth also may dance to confuse a predator that uses visual cues for hunting. VOLUME 108, NUMBER 2 ss, ., \ 1 i] / / 7 7 We 7 7 / , / / ., ‘ ae Sy. mc ae S, She niece Fig. 10. its legs and rotates from point A to point F and back to point A to make one dancing rotation. Rotation can be clockwise or counterclockwise. At El Ceibo, similar dancing behaviors were observed in distantly related arthro- pods that have very different methods of courtship, suggesting that factors other than courtship may have selected independently for dancing in different lineages. A dancing pattern nearly identical to that of Beltheca oni was observed in a black psycodid fly and several Collembola on the surface of leaves from which B. oni were collected. Similarly, Robinson (1988:73) reported that, “In Sulawesi, [a] momphid and its dance are the model for, or mimic an iden- tical display by a similarly coloured collem- bolan—Lepidocyrtus sp. (Entomobryi- dae).”” We believe that microlepidoptera dance for courtship, but it may also be that moths and distantly related arthropods dance to avoid capture by confusing pred- ators such as salticid spiders that use visual cues to adjust for jumping distance. Future studies on dancing demand rigorous hy- pothesis-based behavioral experiments that ee A Z Z ee KF. LOPE au / yo" Ans ene a AS Rqrrn nena ., N. ‘. ~~ >. mre, Illustration showing how Beltheca oni dances while drinking from a drop of water. The moth shuffles test the ecological and evolutionary signif- icance of dancing in moths and other ar- thropods. ACKNOWLEDGMENTS We thank Klaus Sattler (formerly De- partment of Entomology, The Natural His- tory Museum, London, UK) for discussion on Beltheca. John Brown (Systematic En- tomology Laboratory [SEL], USDA, Wash- ington D.C., USA) and Ronald Hodges (formerly SEL, USDA, Washington D.C., USA) gave critical comments on an earlier version of the manuscript which greatly im- proved its quality. Christian Thompson (SEL, USDA, Washington D.C., USA) pro- vided nomenclatural suggestions on the specific epithet. Donald Davis (Department of Entomology, USNM, Washington D.C., USA), Keitaro Eda (Shizuoka, Japan), Jerry Powell (Essig Museum of Natural History, Berkeley, CA, USA), Gaden Robinson (De- partment of Entomology, The Natural His- 260 tory Museum, London, UK), and Jadranka Rota (University of Connecticut, Storrs, CT, USA), provided information on dancing be- havior. We also thank Robert Denno, Ste- phanie Johnson, Silvana-Marten Rodriguez, Jeffrey Sosa-Calvo, and Gina Wimp (Uni- versity of Maryland, College Park, MD, USA) who gave comments on the manu- script, and discussed the potential evolu- tionary and ecological significance of danc- ing. The first author thanks David Wagner (University of Connecticut, Storrs, CT, USA) for introducing him to the microlep- idoptera of Costa Rica in 2002. This study was made possible by funding provided to the first author through the Arthropods of La Selva (ALAS) ATBI research expedi- tions (2002-2004), The University of Maryland Graduate School, and the Orga- nization for Tropical Studies (OTS) 04-01 Tropical Ecology graduate course. LITERATURE CITED Aiello, A. and V. O. Becker. 2004. Display of the “Peacock Moth’: Brenthia spp. (Choreutidae: Brenthiinae). Journal of the Lepidopterists’ Soci- ety 58: 55-58. Busck, A. 1914. New genera and species of microlep- idoptera from Panama. Proceedings of the United States National Museum 47: 1—67. Clarke, J. EF G. 1969. Catalogue of the type specimens of microlepidoptera in the British Museum (Nat- ural History) described by Edward Meyrick, Vol. 6. Glyphipterigidae, Gelechiidae (A—C). Trustees of the British Museum (Natural History), London, 537 pp. Clemens, B. 1860. Contributions to American Lepi- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dopterology, No. 4. Proceedings of the Academy of Natural Sciences of Philadelphia [12]: 156— 174. Comstock, J. H. 1918. The wings of insects: an ex- position of the uniform terminology of the wing- veins of insects and a discussion of the more gen- eral characteristics of the wings of the several or- ders of insects. Comstock, Ithaca, 430 pp. Fletcher, T. B. 1920. Life-histories of Indian Insects— Microlepidoptera. Memoirs of the Department of Agriculture of India, Entomological Series 6: 1— DATE Forbes, W. T. M. 1923. The Lepidoptera of New York and neighboring states. Cornell University Agri- cultural Experiment Station, Memoir 68, 729 pp. Klots, A.B. 1970. Lepidoptera, pp. 115-130. Jn Tuxen, S. L. ed. Taxonomist’s Glossary of Genitalia in Insects. Munksgaard, Copenhagen, 359 pp. Meyrick, E. 1922. Descriptions of South American Mi- cro-Lepidoptera. Transactions of the Entomologi- cal Society of London 1922: 65-116. Morrison, B. 1968. A further record of Dryadaula pac- tolia Meyrick (Lep., Tineidae) in Britain with notes on its life history. Entomologist’s Gazette 19: 181-188. Plant, C. W. 1980. Unusual behavior of Mompha nod- icolella Fuchs (Lep.: Momphidae). The Entomol- ogist’s Record and Journal of Variation 92: 255— 256. Robinson, G. 1988. The systematic position of Ther- mocrates epischista Meyrick (Lepidoptera: Tinei- dae) and the biology of the Dryadaulinae. Nota Lepidopterologica 11: 70—79. Robinson, G. S., K. R. Tuck, and M. Schaffer. 1994. A Field Guide to the Smaller Moths of South-East Asia. Malaysian Nature Society, Kuala Lumpur, 309 pp. Rota, J. 2003. Choreutidae (Lepidoptera) of La Selva Biological Station (Costa Rica): taxonomy, ultra- structure of immature states, and life history. Mas- ter’s Thesis, University of Connecticut, Storrs, 160 Pp. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 261-266 A DESCRIPTION OF THE LARVA OF ARGIA FUNCKI (SELYS, 1854) (ODONATA: ZYGOPTERA: COENAGRIONIDAE) RODOLFO NOVELO-GUTIERREZ AND JOSE ANTONIO GOMEZ-ANAYA Instituto de Ecologia, A.C., Departamento de Entomologia. Apartado Postal 63, 91070 Xalapa, Veracruz, Mexico (e-mails: novelor@ecologia.edu.mx; anaya@ecologia.edu.mx) Abstract.—The last instar larva of Argia funcki (Selys) is described and illustrated. A comparison to its closest relative, Argia lugens (Hagen), is provided. The scalelike setae on sternite 8 and gonapophyses, and the bluntly-tipped gonapophyses easely separate A. funcki larva from that of A. /ugens. The larva of A. funcki belongs to the group of species with ligula very prominent and one palpal seta; it is the largest of all known larvae of the genus. Resumen.—Se describe e ilustra el ultimo estadio larval de Argia funcki (Selys), y se le compara con su pariente cercana Argia lugens (Hagen). Las sedas escamiformes en el esternito 8 y en las gonapofisis, asi como la punta roma de estas ultimas, diferencian facilmente a la larva de A. funcki de la de A. lugens. La larva de A. funcki pertenece al grupo de especies con ligula muy prominente y una seda en el palpo; es la mas grande de todas las larvas del género. Key Words: Mexico Argia funcki (Selys 1854) is a species re- stricted to Mexico and Guatemala (Paulson 1982) and the largest species of the genus. Together with its closest relative A. lugens (Hagen 1861), it belongs to the subgenus Hyponeura Selys. Recently, Paulson (2002) reported a population of A. funcki from Nayarit, Mexico, in which the adult males show an unusual, non-metallic bright red coloration. We observed the same colora- tion in the population from which we ob- tained the larvae for this study, located at Michoacan State, Mexico (Rio Pinolapa), as well as a dichromatism in females: blue and brown morphs. The larva of A. lugens was first described by Needham and Cockerell (1903), and par- tially illustrated by Needham (1904), and more recently by Novelo-Gutiérrez (1992). In this paper we describe, for the first time, Odonata, Zygoptera, Coenagrionidae, Argia funcki, larva, description, the last instar larva of A. funcki. Although we did not rear specimens until emergence, there is a reasonable certainty that they be- long to that species. Argia funcki (Selys) (Figs. 1-10) Material studied.—Four exuviae (¢d), | last instar larva (d¢), 2 penultimate instar larvae (d), 3 antepenultimate instar larvae (1 6,2 2). MEXICO: Hidalgo; Pemuxtitla, Rio Zacuala (900 m), 18 April 1998, R. Novelo leg., 3 ¢ (exuviae); Michoacan; Te- palcatepec, Rio Pinolapa (625 m), 16 May 2002, J.A. Gomez leg., | d (exuvia); same data but 12 March 2004, R. Novelo leg., 4 3,2 @. All material deposited in Coleccion Entomoloégica del Instituto de Ecologia, A.C. (EXA), Xalapa. Description.—Exuviae yellowish to yel- 262 lowish brown, larvae mottled brown on yel- low background on dorsum, yellow to red- dish yellow on venter; body robust and short, integument more spiny than setose; epiproct and paraprocts (caudal lamellae) obscurely mottled, swollen basally (Fig. la). Head: Color pattern as in Fig. 1, wider than long, dorsoventrally flattened, cephalic lobes large and bulging, covered with ro- bust spiniform setae on posterior and infe- rior surfaces. Medial margin of compound eyes straight. Labrum setose, dark brown with a large pale spot on each side of mid- line; anteclypeus dark brown, with a row of long setae on boundary with postclypeus, remainder of head light brown and quite smooth; occipital margin widely concave, with a thickened ridge at middle, becoming gradually thinner at sides. Antenna 7-seg- mented (Fig. 2), scape creamy, remainder of antenna reddish yellow, relative size of antennomeres: 0.25, 0.45, 1.0, 0.7, 0.5, 0.25, 0.15. Mandibles (Fig. 3) with 4 inci- sor cusps; right mandible with an extra cusp at base of largest ventral cusp, molar area with one small, blunt cusp; molar area of left mandible with two small blunt cusps; a dorsobasal emargination in both mandibles (see arrow on Fig. 3). Ventral pad of hy- popharynx more or less rectangular, with a tuft of 7—8 long setae located subapically to each anterolateral corner, and a transverse row of smaller setae on anterior margin (Fig. 4). Maxilla: Galeolacinia (Fig. 5) with 6 teeth, three dorsal teeth more or less of same size, three ventral teeth of different sizes, apical one longest, a row of stiff setae preceding both ventral and dorsal teeth. La- bium: Prementum-postmentum articulation reaching basal half of mesosternum; pre- mentum (Fig. 6a) longer than wide, colored ventrally as follows: a subrhomboidal, whitish, central area except tip of ligula red- dish yellow; a wide, dark brown band on each side of whitish area, converging ba- sally; lateral borders creamy pale; a row of 37—40 spiniform setae along lateral mar- gins, 13-18 basidorsal spiniform setae; lig- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ula very prominent, its apical margin finely serrulate. Palp bicolored, creamy yellow dorsally, dark brown ventrally, with one long seta and several spiniform setae along its dorsal border, ventral margin serrated (Fig. 6b); dorsal margin of dorsal end hook serrated (Fig. 6c); movable hook long and sharp. Thorax: Pronotal disc spiny at sides, posterior border widely convex and thick- ened. Synthorax robust, anterior wing sheaths extending to basal 0.6 of abdominal segment 5, posterior ones extending slightly beyond posterior margin of segment 5, yel- low brown with diffuse whitish spots at base, nodus, tips, and external borders. Legs yellow, with two dark rings on femora and tibiae (Fig. 1b); basal tarsomeres light brown, apical one brown; claws yellow; femora flattened laterally, with two rows of spiniform setae on dorsal and ventral bor- ders, profemora also with spiniform setae on anterior (external) surface; tibiae with one row of spiniform setae on anterior (ex- ternal) and posterior (internal) borders, as well as abundant, delicate, white, long setae on anterior surfaces; tarsi with two ventral rows of abundant, short, stout setae, either bi- or tridentate on apical tarsomeres, and long setae on dorsum. Abdomen: Cylindrical, tapering caudad gradually, with a pattern of alternating pale and dark longitudinal bands (Fig. 1), pale middorsal band with tufts of long, brown setae; posterior and lateral margins and dor- sal surface of tergites 5—10 with reddish- brown spiniform setae; posterior margin of tergite 10 deeply emarginated. Sternites 1— 5, and most of 6 quite smooth, with some spiniform setae on posterior margin of 6; posterior margins of sternites 7—8, and cen- tral area of 8 with scalelike setae (Fig. 7a); posterior margins of sternites. 9—10, and en- tire surface of 10 with spiniform setae (Fig. 7a). Male gonapophyses pyramidal, blunt- ly-tipped, in ventral view (Fig. 7a) slightly divergent and reaching posterior margin of sternite 10, with 23—24 scalelike setae on their ventral borders, and one scalelike seta VOLUME 108, NUMBER 2 263 Figs. 1-5. Morphology of Argia funcki larva. 1, Last instar larva: (a) dorsal view; (b) left hind leg, lateral view. 2, Right antenna, lateral view. 3, Mandibles, ventrointernal view: (a) right mandible; (b) left mandible (arrow indicates the emarginations). 4, Ventral pad of hypopharynx, ventral view. 5, Right galeolacinia, ven- trointernal view. 264 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 6-9. Details of the morphology of A. funcki larva. 6, Prementum: (a) dorsal view; (b) detail of left palp, dorsal view; (c) detail of left palp, frontolateral view. 7, Details of male abdomen: (a) sternites 8-10 showing gonapophyses and types of setae, ventral view; (b) gonapophyses, right lateral view. 8, Details of female abdomen: (a) sternites 9-10 showing gonapophyses, ventral view; (b) gonapophyses, left lateral view. 9, Male cerci: (a) laterodorsal view; (b) posterior (caudal) view; (c) dorsal view. VOLUME 108, NUMBER 2 Fig. 10. paraproct, lateral view; (b) epiproct, left lateral view. Caudal lamellae of A. funcki larva: (a) left on external surface (Fig. 7b). Female gon- apophyses (antepenultimate instar): laterals surpassing posterior margin of sternite 10, ending in long, parallel, blunt tips (Fig. 8a); ventral border with more or less three rows of 23-24 scalelike setae over basal 0.75, and one scalelike seta on external surface; central valvae smooth, shorter than laterals. Male cercus short, apically rounded (Fig. 9). Caudal lamellae: Paraproct (Fig. 10a) swollen at basal 0.7, laminar at apical 0.3, oblong, heavily mottled, ending in a short, apical filament, three times longer than its widest part, ventral margin roughly straight, dorsal margin more or less parallel to ven- tral one at basal 0.65, then curving down- ward gradually on apical 0.35; dorsal and ventral borders with small spiniform setae on basal 0.03, and white, delicate, hairlike setae on apical 0.2 and 0.3, respectively; a thick lateral carina on basal 0.3 beset with a patch of 4—5 rows of reddish-yellow spi- niform setae. Epiproct (Fig. 10b) swollen on basal 0.6, laminar on apical 0.4, heavily mottled, ending in a small tip, two times longer than its widest part, dorsal and ven- tral margins more or less straight and par- allel along basal 0.65, then gradually con- verging at apical 0.35, dorsal margin with spiniform setae at basal 0.01, and white, delicate, hairlike setae on apical 0.2; thick lateral carinae with 2—3 rows of reddish yel- low, spiniform setae on basal 0.07. Measurements (mm) (last instar larva only): total length (excluding caudal app.), 20.0; abdomen (ventral), 11.0; maximum width of head, 6.1; metafemur (dorsal mar- gin), 6.0; epiproct, 7.6; paraproct, 10.0. Remarks.— Argia funcki inhabits shal- low, rocky streams. Mature larvae were found close to the edge, in areas with abun- dant cobble and where the water flow is very slow or absent. The larve were very hard to find; it was necessary to pick up individual rocks to capture them. Mature larvae are apparently present only during a brief period of the dry season (late March to early April), at least in Rio Pinolapa (Mi- choacan); during this time, they are sepa- rated from each other in isolated, small pools formed by water evaporation at the stream edges. This isolation probably helps them to avoid predation by fish, which were very abundant at the central, mainstream, water flow. No A. funcki larvae were found latter microhabitat. Discussion.—The larva of Argia funcki belongs to the group of species with one palpal seta and very prominent ligula, and it is the largest of all known Argia larvae (although Novelo-Gutiérrez, 1992, Table 1, recorded a total length for males of A. lac- rimans of 21—21.9 mm, these measurements were made from alcoholic specimens with the abdomen relaxed, with several of the intersegmental membranes exposed). It shows a great resemblance to its closest rel- ative A. /Jugens. The main features that dis- tinguish both species at the larval stage are (A. lugens in parentheses [cf., Novelo-Gu- tiérrez (1992)]): a larger stature [total length excluding app. 20 mm] (smaller, 14.6—18.7 mm); a large size proportion between 3rd antennomere and scape and pedicel [1.0, 0.25, 0.45, respectively] (smaller size pro- portion, WO 0355 055, “tespectively) 7-4 smaller size proportion between 3rd and 4th antennomeres [1.0, 0.7, respectively] (larg- er size proportion, 1.0, 0.6, respectively); lateral margins of prementum with spini- 266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON form setae all the way along, excepting ex- treme base (spiniform setae at apical 0.60, only); scalelike setae on sternite 8 and gon- apophyses (spiniform setae on these struc- tures); male and female gonapophyses bluntly-tipped (sharply pointed); lateral ca- rina of paraproct short and rounded, and with spiniform setae only (larger and sharp, with spiniform setae intermingled with stiff, hairlike setae); epiproct and paraprocts densely mottled on pale background (usu- ally uniformly dark except for pale tip). ACKNOWLEDGMENTS We thank Dr. Sidney W. Dunkle (Plano, TX), Dr. Ken Tennessen (Wautoma, WI) and Dr. Thomas W. Donnelly (Binghamton, NY) for their comments and corrections on the final manuscript which improved it very much. Financial support was provided by a CONACYT grant (43091-Q). This work is part of the Doctoral dissertation of the ju- nior author, M. Sc. José Antonio Gomez- Anaya, of the Universidad Autonoma del Estado de Hidalgo. LITERATURE CITED Needham, J. G. 1904. New dragon-fly nymphs in the United States National Museum. Proceedings of the United States National Museum 27(1371): 685-720, 7 plates. Needham J. G. and T. D. A. Cockerell. 1903. Some hitherto unknown nymphs of Odonata from New Mexico. Psyche 10: 134-139. Novelo-Gutiérrez, R. 1992. Biosystematics of the lar- vae of the genus Argia in Mexico (Zygoptera: Coenagrionidae). Odonatologica 21(1): 39-71. Paulson, D. R. 1982. Odonata, pp. 249-277. In Hurl- bert, S. H. and A. Villalobos-Figueroa, eds. Aquatic Biota of Mexico, Central America and the West Indies. San Diego State University, San Di- ego, California. . 2002. Odonata records from Nayarit and Sin- aloa, Mexico, with comments on natural history and biogeography. Odonatologica 31(4): 359— 370. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 267-273 HYLURGOPS PALLIATUS (GYLLENHAL) (COLEOPTERA: CURCULIONIDAE: SCOLYTINAE), AN EURASIAN BARK BEETLE NEW TO NORTH AMERICA E. RICHARD HOEBEKE AND ROBERT E. ACCIAVATTI (ERH) Department of Entomology, Cornell University, Ithaca, NY 14853, U.S.A. (e- mail: erh2@cornell.edu); (REA) NA, Forest Health Protection, USDA Forest Service, Morgantown, WV 26505, U.S.A. (e-mail: racciavatti @fs.fed.us) Abstract.—Hylurgops palliatus (Gyllenhal), a widespread and abundant bark beetle in Europe and elsewhere in the Old World, is reported for the first time from North America, based on specimens collected between 2001 and 2004 from New York, Ohio, and Penn- sylvania. This bark beetle was collected from baited Lindgren funnel traps, part of a USDA, Forest Service and APHIS National Early Detection Pilot Project surveying ports for exotic Curculionidae: Scolytinae (bark and ambrosia beetles). The native distribution, biology, and habits of this newly arrived Palearctic species are summarized from the European literature. An existing key to North American species of Hylurgops LeConte is modified to include this newly detected immigrant. A diagnosis and habitus photographs of the adult are provided for recognition of this species. Key Words: species, new records Various exotic species unintentionally in- troduced into North America represent a se- rious threat to the integrity of North Amer- ican forest ecosystems. An Exotic Pest Rap- id Detection Team (EPRDT), under the joint auspices of the USDA Forest Service and USDA-APHIS, was established in 2001 to develop effective national strategies for the early detection and rapid response to ex- otic forest pests. The EPRDT promulgated national strategies for the early detection of exotic bark and ambrosia beetles (Curcu- lionidae: Scolytinae) and implemented pilot trapping surveys with a high degree of em- phasis placed on the expert identification skills of participating taxonomists. The National Early Detection and Rapid Response (EDRR) Pilot Project for 2001 in- volved trapping surveys using host volatiles and pheromones specific to bark beetles as baits placed on Lindgren funnel traps at Coleoptera, Curculionidae, Scolytinae, Hylurgops, North America, exotic nine ports in three regions (3 ports each in the Northeast, Southeast, and Western re- gions). As a result of the 2001 Northeastern regional survey, the Eurasian species, Hy- lurgops palliatus (Gyllenhal), was detected for the first time in North America. Species identity was confirmed by the senior author, based on the initial trapping of 11 speci- mens in a small planted stand of mixed co- nifers [Pinus sylvestris L., P. resinosa Ai- ton, and Picea abies (L.) Karsten] at an in- dustrial site in Erie, Pennsylvania, on 17 April 2001; 2 more specimens were trapped at the same site on 2 May 2001. In subse- quent years, EDRR provided a rapid re- sponse to this initial discovery through de- limiting surveys for H. palliatus in NY, OH, and PA at sites with the above coniferous hosts. Additional recoveries of this exotic bark beetle were made from other baited Lindgren funnel traps placed at several lo- 268 cations in Erie County, PA, in 2002 and from many traps placed in surrounding counties in NY, OH, and PA in 2003 and 2004. This paper reviews known and published information about H. palliatus. Herein, we provide: 1) a list of distributional records and a map for this newly detected bark bee- tle in the eastern United States; 2) a diag- nosis and revised species key for facilitating its recognition in North America; 3) a sum- mary of its native distribution, host tree preferences, biology and habits, and inter- ceptions; and 4) its response to lures used with the EDRR traps. Hylurgops palliatus (Gyllenhal) Hylesinus palliatus Gyllenhal 1813: 340. Hylastes palliatus: Thomson 1865: 349. Hylastes (Hylurgops) palliatus: Reitter 1894: 63. Hylurgops palliatus: Reitter 1913: 50. Diagnosis.—Hylurgops palliatus is su- perficially similar to some species of the ge- nus Hylastes Erichson, but can be differ- entiated by the following combination of characters: pronotum with a noticeable an- terior constriction (Fig. 1), usually with about equal numbers of larger and small punctures intermixed on the disc; third tar- sal segment broad, bilobed; elytra broad- ened posteriorly; elytral declivity usually with sparse, uniseriate rows of fine, erect, hairlike, interstrial setae; scalelike vestiture of elytra present from base to apex of de- clivity, more numerous on apical third; in- terstrial tubercles in a single row; small spe- cies, total length 2.5—3.2 mm (Wood 1982, Cavey et al. 1994). To accommodate H. palliatus, couplet #7 (pg. 83) of Wood’s (1982) key to the North and Central American species of Hylurgops should be modified as follows (alterations in boldface): 7. Scalelike elytral vestiture confined to de- clivity, or extending anteriorly to middle of elytral disc; interstrial raised tubercles con- fused, not uniseriate; total length > 3.2 mm PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON — Scalelike elytral vestiture present from base of elytra to apex of declivity, more nu- merous on apical third; interstrial tuber- cles in a single row; total length 2.5—3.2 mm; New York, Pennsylvania, and Ohio (introduced from Europe) ee Ceiernis rae See palliatus (Gyllenhal) 7a. Scalelike elytral vestiture confined to decliv- ity; elytral disc with only a few crenulations in uniseriate rows, these usually narrow and more nearly tuberculate, most interstrial punc- tures on middle half almost normal (not cren- ulate); setae on lateral and basal margins of pronotum very fine; pronotum reticulate; 3.6— 4.8 mm; California to Coastal British Colum- bia and Alaska rugipennis rugipennis (Mannerheim) — Scalelike elytral vestiture expanding anteri- orly to middle of elytral disc; elytral disc with most interstrial punctures crenulate or sub- crenulate, confused; setae on lateral and basal margins of pronotum coarse; pronotum smooth, shining; 3.7—5.2 mm; Utah and Col- orado to inland British Columbia and east through the coniferous forest to Nova Scotia and Alabama rugipennis pinifex (Fitch) Native distribution.—Hylurgops pallia- tus is widely distributed in the Old World, including Africa (Algeria), Asia (Heilong- jiang in China, Japan, Korea, Turkey, and Sakhalin Island, Siberia, and Ussuri in east- ern USSR), and Europe (Austria, Belgium, Bulgaria, Czechoslovakia, Denmark, Eng- land, Finland, France, Germany, Ireland, It- aly, Netherlands, Norway, Poland, Spain, Sweden, Switzerland, and Yugoslavia) (Wood and Bright 1992). Biology, hosts, and habits.—Hylurgops palliatus, considered a non-aggressive bark beetle (Volz 1988) or so-called ‘secondary’ species (Perttunen 1957), colonizes stems and crowns of dying, decayed, or damaged conifers (Escherich 1923; Byers 1992), with some preference for Norway spruce (Picea abies) in the initial stages of dete- rioration (Nuorteva 1956; Kohnle 2004). Host selection is largely governed by olfac- tory recognition of host-specific volatiles (terpene mix) and ethanol (Schroeder 1988, Volz 1988, Byers 1992, Kohnle 2004). A study by Perttunen (1957), however, showed that H. palliatus was strongly re- N VOLUME 108, NUMBER 269 Fig. 1. pelled by a high concentration of the ter- pene alpha-pinene, and only slightly re- pelled by a low concentration. Others (Kohnle 1985, Klimetzek et al. 1986, Volz 1988) have found that H. palliatus responds to ethanol alone, and that increasing ethanol concentrations actually enhances the re- sponse of adults to monoterpenes. Hylurgops palliatus. Lateral aspect (top) and dorsal aspect (bottom). Scale line = | mm. Female beetles produce a single, longi- tudinal egg gallery, 3—5 cm in length (Schwenke 1974; Brauns 1991). Larval feeding tunnels may be individual for their entire length, or may merge occasionally and again separate, forming a characteristic branched pattern and terminate in pupal cells (Browne 1968). Egg and larval galler- 270 ies are illustrated by Escherich (1923, Abb. 292, p. 568), Schwenke (1974, Abb. 156C, p. 396), and Brauns (1991, Abb. 482, p. 604). In Europe, most workers record 2 flight periods, one in March/April and an- other in July, thus indicating 2 generations annually (Escherich 1923; Braun 1991). Maturation feeding by the adults occurs in the bark of the hosts. The species is mo- nogamous. Adults are known to overwinter in the soil (Balachowsky 1949). Because this beetle is essentially a secondary invad- er of unhealthy trees, and frequently fol- lows an infestation by pine shoot beetle, Tomicus piniperda (L.), it is generally not considered an important pest species (Bal- achowsky 1949; Browne 1968). In its na- tive Eurasian range, its known host taxa are extensive, including the following conifers: Abies alba Miller, A. sibirica Ledebour, A. nordmanniana (Steven) Spach, Larix decid- ua Miller, L. gmelinii Ruprecht (Rupr. ex Kuzeneva), L. sibirica Ledebour, Picea abi- es, P. glehnii (Friedr. Schmidt), P. jezoensis (Siebold & Zuccarini) Carriere, P. koyamai Shirasawa, P. microsperma (Lind.) Carri- ére, P. obovata Ledebour, P. omorika (Pan- cic) Purkyne, P. orientalis (L.) Link, Pinus cembra L., P. leucodermis Antoine, P. mugo Turra, P. nigra Arnold (and its vari- ous subspecies), P. peuce Grisebach, P. pi- naster Aiton, P. pinea L., P. radiata D. Don, P. rotundata Link (Moor-Bergkiefer), P. sibirica (Du Tour) Krylov, P. strobus L., P. sylvestris, P. uncinata Miller ex Mirbel, and Cedrus spp. (Wood and Bright 1992, Lombardero 1994, Bright and Skidmore 1997): Interceptions.—Aylurgops palliatus was one of ten targeted bark and ambrosia bee- tles selected for the National Early Detec- tion and Rapid Response Pilot project, based on the frequency and number of in- terceptions of this species at U.S. ports-of- entry. It is the third most frequently inter- cepted bark beetle in the United States, with 295 interceptions from 20 countries for the period 1985—2000 (Haack 2001). In Cana- da, this bark beetle was intercepted from PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON imported cargoes on at least 5 occasions from April 1, 1997—March 31, 2000 (CFIA 2002). New North American records.—All rec- ords of H. palliatus listed below and mapped in Fig. 2 are based on specimens collected from baited Lindgren funnel traps set at selected sites from mid-March through late May, 2001—2004. Traps were fitted with collection cups partially filled with propylene glycol (low toxicity) and water (1:1, vol./vol.). Rain and snow often diluted this volume beyond the 1:1 ratio. Traps were generally checked every two weeks. The year each site was trapped, with the number of specimens collected in pa- rentheses, is noted below. Voucher speci- mens have been deposited in the Cornell University Insect Collection, Ithaca, NY; the National Museum of Natural History, Smithsonian Institution, Washington, DC; and the Carnegie Museum of Natural His- tory, Pittsburgh, PA. UNITED STATES: NEW YORK: Cat- taraugus Co., Allegany State Park, 2003 (1). Chautauqua Co., Panama, 2003 (1). OHIO: Ashtabula Co., Monroe Center, 2003 (50); Pymatuning State Park, 2003 (121). Columbiana Co., Beaver Creek State Park, 2004 (1); New Waterford, 2004 (1). Cuyahoga Co., Oakwood, 2004 (12). Geau- ga Co., Parkman, 2004 (55); Thompson, 2004 (19). Lake Co., Leroy Center, 2004 (9); North Madison, 2004 (16). Mahoning Co., Meander Reservoir, 2004 (10); North Lima, 2004 (9). Portage Co., Garretsville, 2004 (172); Mantua Corners, 2004 (17). Summit Co., Cuyahoga Valley NP, Horse- shoe Pond, 2004 (6); Cuyahoga Valey NP, Oak Hill, 2004 (1). Trumbull Co., Youngs- town State University Arboretum, 2003 (9). Tuscarawas Co., Beach City Dam, 2004 (1). PENNSYLVANIA: Allegheny Co., Beechwood Farms ANC, 2004 (4); Wex- ford, 2004 (1). Beaver Co., New Galilee, 2004 (1). Butler Co., Annandale, 2004 (1); Moraine State Park, 2004 (3). Crawford Co., Crossingville, 2003 (204); New Rich- mond, 2003 (83); Riceville, 2003 (25); VOLUME 108, NUMBER 2 Ashtabula ix® @ Geauga @ e Trumbull e ® 9 Summit id e Mahoning Ashland Coshocton Z i ‘ Washington a S Guernsey Muskingum Belmont Fig. 2. 271 New York = ~ Allegany Cattaraugus sani Cameron Jefferson Pennsylvania Clearfield Butler Armstrong Cambria ® Surveyed, found ©) Surveyed, not found AKS 1/05 Hylurgops palliatus survey in Ohio, Pennsylvania, and New York, 2001—2004. Solid circles = site surveyed, specimens trapped. Open circles = site surveyed, no specimens trapped. Table 1. Aylurgops palliatus trap catch by lure type and year. Total Lure Type Year Number Ethanol + alpha-pinene 2001 14 2002 78 2003 470 2004 369 Subtotal 931 Ethanol only 2001 | 2002 50 2003 229 2004 59 Subtotal 339 Chalcoprax® 2002 19 Subtotal 19 Ipslure® 2002 8 2003 15 2004 | Subtotal 24 Total hauls Shermansville, 2003 (138). Erie Co., Corry, 2003 (17); Erie (Asbury Nature Center), 2002, 2004 (175); Erie (Eriez Magnetics), 2001, 2002, 2004 (51); Erie (Riehl Nurs- ery), 2002 C9): racy 2008-07). Laywr- ence. Co.,. Edinbure,, 2003, 2004. (13): Mount Jackson, 2004 (8); Plain Grove, 2003 (3). Mercer Co., Camp Perry, 2003 (9); Mercer, 2003 (9). Venango Co., Polk, 2003 (2); Wallaceville, 2003 (3). Warren Co., Chandlers Valley, 2003 (1); Hearts Content, 2004 (1); Youngsville, 2003 (3). Washington Co., Hillman State Park, 2003 (1). Commercial lures as_ baits.—Lindgren traps used in this EDRR survey were baited with one of four commercial lures or lure combinations. The trap catches of H. pal- liatus to the lures are shown in Table 1. The lures, available from PheroTech, Inc. (Del- ta, BC) included: 1) Ultra-high release (UHR) ethanol only. This is a general attractant for wood- boring insects in deciduous hosts. Eth- anol is a degradation product from the phloem of weakened and dying trees. 2) UHR alpha-pinene and UHR ethanol to- gether. Alpha-pinene, a monoterpene, is a volatile from coniferous hosts and pro- vides strong attraction for many wood- boring insects, especially bark beetles. 3) Exotic Ipslure®. This is another attrac- tant (with the 3 components: cis-verben- ol, ipsdienol, and methyl butenol) spe- cific for conifer-feeding exotic bark bee- tles [such as the Palearctic species Jps typographus (L.) Ips sexdentatus (Boer- ner), Hylurgus ligniperda (EF), and Or- thotomicus erosus (Wollaston)]. 4) Chalcoprax®, a specific lure for Pityo- genes chalcographus (L.). ACKNOWLEDGMENTS We thank Helen Butalla and Ann Steke- tee (USDA Forest Service, Morgantown, WV) for providing a compilation of all lo- cality data for H. palliatus captured over the survey period and for mapping this lo- cality data, respectively. Kent Loeffler (De- partment of Plant Pathology, Cornell Uni- versity, Ithaca, NY) provided the habitus photographs of H. palliatus. We also are grateful to Robert J. Rabaglia (Maryland Department of Agriculture, Annapolis) and James R. LaBonte (Oregon Department of Agriculture, Salem) for providing helpful comments on a draft of the manuscript. LITERATURE CITED Balachowsky, A. 1949. Coleopteres Scolytides. Faune de France 50: 1-320. Brauns, A. 1991. Taschenbuch der Waldinsekten: Grundriss einer terrestrischen Bestandes- und Standort-Entomologie. G. Fischer Verlag, Stuut- gart, 860 pp. Bright, D. E. and R. E. Skidmore. 1997. A catalog of Scolytidae and Platypodidae (Coleoptera), Supple- ment | (1990-1994). NRC Research Press, Otta- wa, Ontario, 368 pp. Browne, FE. G. 1968. Pest and diseases of forest plan- tation trees: an annotated list of the principal spe- cies occurring in the British Commonwealth. Clar- endon Press, Oxford, UK, 1,330 pp. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Byers, J. A. 1992. Attraction of bark beetles, Tomicus piniperda, Hylurgops palliatus, and Trypodendron domesticum and other insects to short chain al- cohols and monoterpenes. Journal of Chemical Ecology 18: 2385-2402. Cavey, J., S. Passoa, and D. Kucera. 1994. Screening aids for exotic bark beetles in the northeastern United States. USDA, Forest Service, Northeast- ern Area. NA-TP-11-94. 19 pp. CFIA (Canadian Food Inspection Agency). 2002. In- tercepted plant pests, 1997-2000. http:// Www.inspection.gc.ca/english/sci/lab/cpqp/in- troe.shtml (accessed 12 February 2005). Escherich, K. 1923. Die Forstinsekten Mitteleuropas. Band 2, P. Parey, Berlin, pp. 519 532, 568-572. Gyllenhal, L. 1813. Insecta Svecica descripta. Claris I. Coleoptera Sive Eleuterata. Leverentz, Scaris, Vol. pts 0ipps Haack, R. A. 2001. Intercepted Scolytidae (Coleop- tera) at U.S. ports of entry: 1985—2000. Integrated Pest Management Reviews 6: 253-282. Klimetzek, D., J. Kohler, J. P. Vité, and U. Kohnle. 1986. Dosage response to ethanol mediates host selection by ‘secondary’ bark beetles. Naturwis- senschaften 73: 270. Kohnle, U. 1985. Untersuchungen tiber die Pheromon- systeme sekundaérer Borkenkafer (Col., Scolyti- dae). Zeitschrift fiir Angewandte Entomologie 100: 197-218. . 2004. Host and non-host odour signals gov- erning host selection by the pine shoot beetle, Tomicus piniperda and the spruce bark beetle, Hy- lurgops palliatus (Col., Scolytidae). Journal of Applied Entomology 128(9—10): 588-592. Nuorteva, M. 1956. Uber den Fichtenstamm-Bastkifer, Hylurgops palliatus Gyll., und seine Insektenfein- de. Acta Entomologica Fennica 13: 1-116. Lombardero, M. J. 1994. Estudio de los Scolytidae (O. Coleoptera) de Galicia. Departmento de Biologia Animal, Universidad de Santiago de Compostela, Santiago de Compostela. Perttunen, V. 1957. Reactions of two bark beetle spe- cies, Hylurgops palliatus Gyll. And Hylastes ater Payk. (Col., Scolytidae) to the terpene a-pinene. Annales Entomologici Fennici 23: 101-110. Reitter, E. 1894. Bestimmungs-Tabellen der Borken- kafer (Scolytidae) aus Europa und den angrenzen- den Liandern. Verhandlungen des Naturforschen- den Vereines in Briinn 33: 36—97. . 1913. Bestimmungs-Tabelle der Borkenkdfer (Scolytidae) aus Europa und den angrenzenden Landern. Wiener Entomologische Zeitung 32 (Beiheft): 1-116. Schroeder, L. M. 1988. Attraction of the bark beetle Tomicus piniperda and some other bark- and wood-living beetles to the host volatiles alpha-pi- nene and ethanol. Entomologia Experimentalis et Applicata 46: 203-210. VOLUME 108, NUMBER 2 Schwenke, W. 1974. Die Forstschadlinge Europas: ein Handbuch in fiinf Banden/unter Mitwirkung zahl- reicher Wissenschaftler. Zweiter Band (Kifer). P. Parey, Hamburg; 500 pp. Thomson, C. G. 1865. Skandinaviens Coleoptera, syn- optiskt bearbetade, af C. G. Thomson. Tom. VII. Tryckt Uti Lundbergska Boktryckeriet, Lund, 394 Pp. Volz, H.-A. 1988. Monoterpenes governing host selec- tion in the bark beetles Hylurgops palliatus and PS, Tomicus piniperda. Entomologia Experimentalis et Applicata 47: 31-35. Wood, S. L. 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolyti- dae), a taxonomic monograph. Great Basin Natu- ralist Memoirs 6: 1—1359. Wood, S. L. and D. E. Bright. 1992. A catalog of Scol- ytidae and Platypodidae (Coleoptera), Part 2: tax- onomic index. Great Basin Naturalist Memoirs 13: 1—1553. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 274-284 A NEW SPECIES OF APHIS L. (HEMIPTERA: APHIDIDAE: APHIDINA) LIVING ON URTICACEAE IN ARGENTINA M. P. MIER DURANTE, J. ORTEGO, AND J. M. NIETO NAFRiA (MPMD, JMNN) Departamento de Biologia Animal, Universidad de Leén, E-24071 Leon, Spain (e-mails: dbammd@unileon.es; dbajnn@unileon.es); (JO) INTA EEA Junin, Casilla de Correo 78, 5570 San Martin (Mendoza), Argentina (e-mail: jortego@junin. inta.gov.ar) Abstract.—A new Argentinean aphid species, Aphis mendocina, is described. It lives on the native Urticaceae, Urtica mollis. Descriptions of apterous and alatae viviparous females, oviparous females, and males, which are apterous, are presented. An identifica- tion key to the apterous viviparous females of Aphis species with large dorsal abdominal sclerotization recorded from South American is given. Resumen.—Se describe una nueva especie argentina, el pulg6n Aphis mendocina, que vive sobre la urticacea nativa Urtica mollis. Se exponen los caracteres de las hembras Viviparas apteras y aladas, de las hembras oviparas y de los machos, que son apteros. Se presenta una clave de identificaci6n de hembras viviparas apteras de las especies de Aphis con extensa esclerotizacion dorso-abdominal citadas de Sudamérica. Key Words: Aphis mendocina, Aphis, Aphididae, Aphidina, aphids, new species, Men- doza, Argentina, South America, identification key The total number of aphid species re- corded in Argentinean continental territory has risen from 133 to 200 between 1993 and 2003; moreover, the number of record- ed South American native species from Ar- gentina has also increased from 22.6% to 24.5%, including 22 new species described during this period (Nieto Nafria et al. 1994, Ortego et al. 2004), which belong to the subfamily Aphidinae, five of them to the tribe Macrosiphini and 17 to the tribe Aphi- dini and subtribe Aphidina, with 13 species included in the genus Aphis Linnaeus (Or- tego et al. 2004). Only two (Aulacorthum solani (Kalten- bach and Myzus persicae (Sulzer) (Macro- siphini)) of the 200 species have been re- corded on Urticaceae, specifically on Urtica urens L; another seven polyphagous species known from Argentina (Aphis craccivora Koch, A. fabae Scopoli, A. gossypii Glover, A. spiraecola Patch (Aphidini), Macrosi- phum euphorbiae (Thomas), Myzus asca- lonicus Doncaster and My. ornatus Laing (Macrosiphini)) could live on several spe- cies of Urticaceae; it is also possible that Microlophium carnosum (Buckton) (Ma- crosiphini) can be found in the country as it has been recorded from Chile (Smith and Cermeli 1979, Nieto Nafria et al. 1994, Seco Fernandez et al. 2000, Fuentes-Con- treras et al. 1997). Specimens were collected on Urtica mol- lis Steud. (Urticaceae) in Mendoza province (Argentina). When alive they were identi- fied as belonging to a species in the genus Aphis with apterous females similar to those of the polyphagous and well-known Aphis craccivora Koch. These specimens are described below as a new species. VOLUME 108, NUMBER 2 MATERIAL AND METHODS The techniques for catching, conserving, slide mounting and measuring aphids are the habitual ones in aphidology (Nieto Na- fria and Mier Durante 1998 provide a de- tailed exposure). The aphid terminology used is in general also the habitual (Miyazaki 1987; Foottit and Richards 1993), but we use the term papilla (plural: papillae) for the membra- nous and more or less hemispheric tubercle. Several aphid species have aestivating form of apterous viviparous females, which dif- fers from the “‘normal”’ form (from spring, early summer or autumn); it is named: (a) “summer dwarf” (“dwarf’’) if it is very small and diverse to “‘normal’’, or (b) “small summer” (“‘small’’) if it is smaller than “normal” but not very different in characters to it (Miyazaki 1987). We use the term “‘small” for the aestivating form of apterous viviparous females of the new spe- cies caught from January and February (the austral summer), which are at most 1.50 mm long and have an incomplete and brown discal thoracic-abdominal plate, and the term “‘big” for the apterous viviparous females from February to April, which are at least 1.50 mm long (exceptionally 1.40) and have a big and black discal thoracic- abdominal plate. Abbreviations used in the text and figure captions are as follow: AbdlI, AbdII, AbdIII, AbdIV, AbdV, AbdVI, AbdVII, AbdVIII are each abdominal segment I to VIII; Antl, Antlil, Antlll, AntIV, AntV, AntVIb, AntVIpt are antennal segments I to V plus base and processus terminalis of antennal segment VI, respectively; Ars is apical ros- tral segment; D is basal diameter of anten- nal segment III; and Ht2 is second segment of hind tarsus. Values in parenthesis are ex- ceptional values. Aphis mendocina Mier Durante, Ortego, and Nieto Nafria, new species (Figs. 1—5) Apterous viviparous female (n = 590; 34 measured) (Figs. 1, 2).—Body 1.07 to 2.13 DTS) mm. long, 6.1 to 10.8 times siphunculus. Field features: shiny black (big specimens) or dark green or dark brown (small speci- mens), with part of antenna and tibia yel- lowish and without waxy powder. Slide- mounted specimens: big specimens with dark brown dorsum (prothoracic arc, a com- plete discal plate from mesothorax to ab- dominal segment VI, a ring around the base of siphunculus excepted, and a bar on both abdominal segments VII and VIII), siphun- culi black, clypeus, distal part of rostrum and cauda brown and antennae and legs yel- lowish; small specimens with brown head, siphunculi and cauda, light brown to yel- lowish dorsum (a broad thoracic-abdominal patch and bars on prothorax and abdominal segments VII and VIII at most, to at least several disperse sclerites on thorax and ab- dominal segments I to VI) and zones of an- tenna and legs pigmented as dorsal scler- ites. Thoracic and abdominal dorsal cuticle delicately or inappreciably reticulated. Mar- ginal papillae on prothorax ogival and smaller than triommatidium; abdominal I and VII papillae similar in shape but larger than prothoracic ones; other abdominal pa- pillae absent. Front margin straight or moderately sin- uate. Dorsum of head and AntI smoky to dark brown. Setae on vertex fine, acute, 25— 45 wm long and 1.2—2.3 times D. Antenna 6 segmented, 0.50—1.13 mm long, 0.36— 0.67 times body length and without second- ary sensoria. AntlIII as pale as Antll, slight- ly imbricate on ventral side, 0.11—0.35 mm long (1.0—2.0 times AntVIpt) and with 3— 10 (rarely less than 5) setae, which are (13)18—28 pm long and 0.9-1.4 times D. Other segments of antennal flagellum well imbricated; AntIV as pale as AntlIII and 0.08—0.24 mm long; AntV_ progressively pigmented and (0.05)0.08—0.21 mm; AntVIpt smoky pigmented, 0.11—0.19 mm and |.3—1.7 (rarely more than 1.6) times AntVIb, which is 0.07—0.13 mm long and also smoky. Rostrum (0.36)0.42—0.56 mm long, reaching hind coxae and (0.23)0.25— 276 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. 0.36 times body length. Ars dark brown, 0.10—0.15 mm, 1.1—1.3 times Ht2, 1.2—1.6 times AntVIb, and narrow (2.3—3.1 times its basal width) and with slightly concave sides; it has 2 secondary setae. Coxae, trochanters, part of femora (up to the distal half on several small specimens, 0.5mm Aphis mendocina, apterous viviparous female “‘big’’. AntIII and AntVI are detailed. or to dorsal apical patch on several big specimens), apical zone of tibiae and tarsi dusky to brown. Inside seta of hind tro- chanter (33)40—55 wm, 0.8—-1.4 times di- ameter of trochanter-femoral joint. Longest dorsal setae on hind femur (15)25—45(50) wm and (1.2)1.5—2.9 times D. Outside setae VOLUME 108, NUMBER 2 25 Fig. 2. Aphis mendocina, apterous viviparous female “‘small’’. AntIII and AntVI are detailed. on middle length of hind tibia 25-50 pm long and 0.7—1.3 the diameter of the article at its insertion point. First tarsal segment with 3.3.2(3) setae. Ht2 0.09-0.12 mm long. Dorsal setae on AbdII to AbdIV fine and blunt, (23)30 to 45 pm long (1.2—2.3 times D). AbdVIII with 2—4 (very exceptionally 5) dorsal setae, which are fine, acute (as AbdVII ones), (33)40—60(65) pm and (1.4)2.2—3.3 times D. Siphunculus more or less cylindrical, Slightly tapering to apex, rough, (0.10)0.13—0.30 mm, 2.7—5.4 times its width at middle and 0.8—1.4 times cauda. Genital plate with 2—8 discal and 6—16 pos- terior setae. Cauda broad fingerlike, (0.13)0.15—0.24 mm and 1.2—1.7(2.1) times its basal width, with 5—13 setae (more fre- quently 8—11). Alate viviparous female (n = 13; 8 mea- sured) (Fig. 3).—Body 1.55—1.85 mm long. Field features: matt black with dusky an- tenna and brown legs. Slide-mounted spec- imens: antenna and legs more intensely and extensively pigmented than apterae. AbdII- AbdVI (very infrequently on AbdI) with marginal sclerites, AbdV-AbdVII (infre- quently also on AbdI-AbdIV) with spinal sclerites or bar (sometimes broken) and AbdVIII with a bar. Antenna 1.05—1.12 mm long and 0.59—0.68 times body length; 278 Fig. 3. AntlII 0.27-0.30 mm, 1.6—2.0 times AntVIpt and with (2)5—8 secondary senso- ria; AntlV 0.19—0.20 mm and exceptionally with | secondary sensorium; AntV 0.16— 0.19 mm; AntVIb 0.11—0.14 mm; AntVIpt 0.14—0.18 mm and 1.1—1.6 times AntVIb. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Aphis mendocina, alatae viviparous female. AntIII and AntVI are detailed. Other characters as those of apterous vivi- parae. Oviparous female (n = 98; 20 measured) (Fig. 4).—Body 1.47—1.83 mm long, 10.7— 16.4 times siphunculus. Field features: shiny black, with part of antenna and tibiae VOLUME 108, NUMBER 2 STR EE EN BEES “ oT ry i i q : 29 Fig. 4. Aphis mendocina, oviparous female. AntIII and AntVI are detailed. yellowish and without cereous powder. Slide-mounted specimens: head, Antl, AntVI, clypeus, distal part of rostrum, dor- sum of thorax, coxae, distal part of femora, apical % of tibia of anterior and intermedi- ate legs, hind tibia, tarsi, abdominal scler- ites, siphunculus, cauda, anal plate, and the periphery of genital plate dark brown to black; and AntlIlI, distal part of AntV, and trochanters dusky to brown. AbdI-AbdV normally with spinal-marginal bar partially coalesced between them, AbdVI with spinal sclerites, and AbdVII and AbdVIII with a more or less tenuous bar. Ars 0.11—0.13 mm, 1.0—1.4 Ht2, 0.8 to 1.2 times AntVIb. Hind tibia uniformly swollen, outside hairs at middle are 30—43 wm and 0.5—0.7 times diameter of article at its insertion point and with (40)80—150 scent plates (one specimen with only 21 and 25 on each leg). Siphun- culus 0.09—0.17 mm long, 2.0—3.3 times its width at middle and 0.6—1.1 times cauda. Postsiphuncular part of abdomen extended. Abd VIII with 9 to 13(16) dorsal setae. Gen- ital plate with 46—67 setae. Cauda more or less broadly triangular in shape, 0.14—0.17 mm long, 0.9—1.4 times its basal width, and with (8)10—14 setae. Other characters as those of apterous viviparae. Apterous male (n = 112; 15 measured) a te Pe eE. eo 4 0.5mm Fig. 5. (Fig. 5).—Body 1.12—1.30 mm. long, 10.3— 14.4 times siphunculus. Field features: shiny black with antenna and most of legs brown to dark brown. Slide-mounted spec- imens: dark brown dorsum, most of AbdVII-AbdVIII excepted; pigmented an- tenna (specially segments I, V and VI) and legs, (but most part of tibiae and basal part of femora yellowish), brown cauda, anal plate, and parameres, and black siphuncu- lus. Dorsal sclerotization very broad: arc on prothorax, complete or spinal-pleural bar and marginal patches on mesothorax and AbdI to AbdVI, and bar on both abdominal segments VII and VIII. Antenna 6 seg- mented, 0.87—1.18 mm and 0.71—0.90 times body length. AntIII with (11)13-17 (one specimen with O and 2 on each antenna), both AntIV and AntV with (1)6—13 and AntVIb habitually without (one specimen with | and another with 8, only on one an- tenna) secondary sensoria. Siphunculus smaller than those of apterous females: 0.08—0.13 mm, 1.9—3.1 times its width at middle and 0.7—1.0 times cauda. Cauda PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Aphis mendocina, male (apterous). AntIII and AntVI are detailed. 0.11—0.13 mm and 0.9—1.3 times its basal width. Other characters as those of apterous Viviparae. Type material—Holotype: apterous vi- viparous female (measured specimen num- ber 29) collected on Urtica mollis Steud. at Malargiie: Refugio del Club Andino (Men- doza province, Argentina, 35°24’S, 69°54'W, 2,183 m), 23-IV-1996, Ortego leg., in collection of the Universidad de Leon (Departamento de Biologia Animal). Paratypes: 589 apterous and 13 alate vivip- arous females, 98 oviparous females and 112 males found with the holotype and on the same host plant at: Malargiie: Refugio del Club Andino, 13-III-2003 (J. Ortego leg.), Malargtie: El Chihuido (35°42’S, 69°38'W, 1,900 m), 9-III-1997 (J. Ortego leg.), Malargtie: Los Molles (35°10’S, 69°57'W, 1,830 m), 5-II-2000 (Mier Du- rante, Nieto Nafria & Ortego leg.) and San Rafael: El Sosneado (Mendoza province, 35°45'S, 69°50'W, 2,000 m), 7-I-1994 (J. Ortego leg.), deposited in the collections of the Universidad de Ledén (Leon, Spain), VOLUME 108, NUMBER 2 INTA EEA Junin (Junin, Argentina), The Natural History Museum (London, United Kingdom), Muséum Nationale d’ Histoire Naturelle (Paris, France), and the National Museum of Natural History, Smithsonian Institution aphid collection (Beltsville, MD, USA). Etymology.—The specific name is an ad- jective, named for the inhabitants of Men- doza province; it is in the feminine gender as Aphis. Biology and distribution.—Aphis men- docina 1s monoecious and holocyclic on Urtica mollis (see discussion). Dense col- onies are formed on the stems and leaf pet- ioles, and large populations extend to the leaf limbs. The number of alatae and alate nymphs collected is very small, but we cannot be sure that such a low production of alatae (general or coinciding with this moment in time) is a usual characteristic of the species. Currently, it is only know from Mendoza province, but it is possibly distributed in ar- eas in South America where U. mollis lives, e.g., Andean provinces of Argentina from Jujuy to Santa Cruz, a part of Chile and Peru (Missouri Botanical Garden 1999). Discussion.—Aphis mendocina belongs to the nominotypical subgenus according to Eastop (1979a) and to the South American group of species according to Hille Ris Lambers (1974) and Remaudiére (1994). The taxonomic identity of the host plant is much-discussed (Missouri Botanical Gar- den 1999, Zuloaga and Morrone, 1999), for some authors it is a good species, whereas for others it is a variety from the Euroasiatic Urtica dioica. However, the authors (Ar- gentinean) of the latter opinion have no hes- itation in claiming that it is a variety from Argentina! The presence of a monoecious aphid (with apterous males!) could be an- other reason for considering them as differ- ent species as the specificity and taxonom- ical accuracy of the aphids is well-known (Eastop 1979b, 1998). If the hypothesis that the host plant is Urtica dioica were true and as Aphis mendocina belongs to the South 281 American species group in the genus Aphis, due to its characters, we would have to con- sider that it must live on an autochthonous species belonging to the genus Urtica or at least to the family Urticaceae. The search for this hypothetical plant has given no re- sults, but the territory to be prospected is enormous! Apterous viviparous females of Aphis mendocina usually have discal or spinal- pleural plate, sometimes with holes or bro- ken, on thorax and abdomen or only on ab- domen. A large dorsal sclerotization (from large segmental bands to discal plate) is also present in another fifteen species of ge- nus Aphis recorded from South America (Hille Ris Lambers 1974; Mier Durante and Ortego 1999, Mier Durante et al. 2003; Nie- to Nafria and Ortego 2002; Nieto Nafria et al. 1999; Ortego, 1998; Ortego and Mier Durante 1997; Remaudiére 1994): A. als- tromeriae Essig (Chile), A. berberidorum Ortego & Mier Durante (Argentina and Chile), A. cinerea Nieto Nafria & Ortego (Argentina), A. craccivora (introduced spe- cies; several South American countries), A. cytisorum Hartig (introduced species; Ar- gentina, Peru), A. danielae Remaudiére (Argentina), A. intrusa Ortego (Argentina), A. malalhuina Mier Durante, Nieto Nafria and Ortego (Argentina), A. marthae Essig (Chile), A. melosae Mier Durante and Or- tego (Argentina), A. mulini Hille Ris Lamb- ers (Argentina), A. Lambers (Argentina), A. papillosa Mier Durante, Nieto Nafria and Ortego (Argen- tina, Chile), A. roberti Nieto Nafria, Ortego and Mier Durante (Argentina, Chile), and A. senecionicoides Blanchard (Argentina). These species can be identified using the following key. Data from above mentioned authors and Garcia Prieto and Nieto Nafria (in press), plus our new observations. Note mulinicola Hille Ris that poorly-sclerotized or unsclerotized specimens are present in several species. Host plant and color when alive are given in square brackets. Apterous viviparous females with large dorsal sclerotization of the species of Aphis PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON KEY TO SPECIES recorded from South America ih N AbdII-AbdIV usually without marginal papil- lae, sometimes with | to 3 in all; if excep- tionally there are 4 to 6, siphunculus shorter than 0.5 times cauda, or if longer (0.7—1.2 times): Ars 0.09—0.13 mm and 1.0—1.3 times Ht2, and short setae (on AntlII, AbdII-AbdIV and AbdVIII respectively 10-14, 13-28 and QO=SS MMMM raed ote ternary oe ee eae 2 AbdII-AbdIV usually with at least 4 marginal papillae in all; if 1—3, exceptional “big” spec- imens living on Grindelia chiloensis .... . 1s Sclerotization dorsal on thorax and abdomen evidently segmental, with spinal-pleural bands, sometimes partially coalescent be- tween and/or to marginal small patches .... 3 Abdominal or thoracic-abdominal spinal- pleural to discal plate present, sometimes with intersegmental and/or spinal holes Ars 0.8—1.0 times Ht2. Cauda 0.21—0.31 mm. Setae on AbdVIII 22—33 ppm. Genital plate with 6—11 posterior setae. [On Lathyrus ma- cropus. White cereous powdered] ... A. cinerea Ars 1.0—1.1 times Ht2. Cauda 0.17—0.20 mm. Setae on AbdVIII 30—47 pm. Genital plate with 12—15 posterior setae. [On Alstroemeria. Possibly black] A. alstroemeriae Secondary sensoria present on AntIII. [On Se- necio subumbellatus. Shiny black] Secondary sensoria absent on AntlIII Siphunculus shorter than 0.5 times cauda. [On Senecio subumbellatus. Shiny dark brown to black] A. malalhuina Siphunculus longer than 0.7 times cauda... 6 AbdI-AbdVI with more than habitual 3 pairs of setae (2 marginal and | spinal). AbdVIII with 2(4) long (45—70 zm) setae, which have strong base and very fine apex. [On Quilajia saponaria. Shiny black] A. marthae AbdI-AbdVI with only habitual 3 pairs of se- tae (2 marginal and | spinal). Abd VIII setae if long in another shape A. intrusa Distal part of hind femur always paler than apical part of tibia and usually as pale as dis- tal part of other femora. AbdVIII with 3—5 setae. Ars 0.12—0.15 mm and 0.8—1.1 times Ht2. [On Lycium sp. Shiny black, sometimes poorly white cereous powdered] ... A. danielae Distal part of hind femur variably pigmented: but if it is pale and AbdVIII with 3-5 setae, Ars 0.09—0.12 mm and 1.1—1.3 times Ht2 .. 8 Ars at most 1.0 times Ht2. Usually incom- plete discal plate from metathorax to AbdVI (frequently broken spinal-pleural plate and AbdII-AbdIV marginal patches). AntVI 1.4— 2.0 times AntVIb (Ortego and Mier Durante 1997 wrotel.71—2.32, but it was a mistake). Siphunculus 0.15—0.31 mm, 2.7—5.3 times its width at middle and 0.8—1.4 times cauda. [On several species of Berberis. Shiny brown] SRN sacar Obs ol ih eaee ee ee A. berberidorum Ars 0.9—1.4 times Ht2; if shorter than 1.1, with other dorsal sclerotization, or other ratio AntVIpt/AntVIb, or other siphunculus lengths (absolute or relative) AntVIpt (1.3)1.6—3.5 times AntVIb; if shorter than 1.6 times, discal plate complete, siphun- culus 0.22—0.46 mm, |.2—2.0 times cauda and setae on Abd VIII 10—23 pm AntVIpt 1.1—2.2 times AntVIb; but if longer than 1.6 times, “dwarf” specimens living on Grindelia chiloensis with broken or incom- plete discal plate, siphunculus 1.0—1.4 times cauda and Ars 1.2—1.4 times Ht2 and setae on AbdVIII 25-38 pm . Shiny black without white cereous powder when alive. Ars 0.10—0.13 mm. [On many species of many families, mainly Fabaceae] eA Bg es ig eat WPS Dan SAE Seen ae A. craccivora White cereous powdered when alive, more or less shiny black in alcohol. Ars 0.11—0.16 mm (frequently up from 0.13 mm). [On arboreal or shrubby Fabaceae species] .... A. cytisorum . Femora mostly dark pigmented, with a pale basal portion. AntII as dark as Antl, and both are darker than AntIII. Setae on AbdII-AbdIV and AbdVIII 13—28 and 20-38 wm. [On Mu- linum spinosum. Shiny black] A. roberti Femora (specially front and middle) mostly pale, sometimes dusky to brown on a distal portion. AntII paler than Antl and nearly as pale as AntIII. Setae on AbdII-AbdIV and AbdVIII 22—45 and 30-65 pm ........ 12 . AntVIpt 1.3—1.7 times AntVIb. Cauda 0.13-— 0.24 mm and 1.2—2.1 times its basal width. [On Urtica mollis. Shiny black if “big” spec- imens, or dark green to dark brown if “‘small”’ specimens] A. mendocina AntVIpt 1.7—2.4 times AntVIb (exceptionally down to 1.4 in several “dwarf”? specimens). Cauda 0.08—0.14 mm and 0.9-1.3 times its basalkwidthivts.cc, peda cess sik A ee 14 . Marginal papillae large, except those on pos- terior abdominal segments; prothoracic papil- lae bigger than triommatidium. [On several species of Senecio. Opaque or shiny dark- brown to black] A. papillosa Marginal papillae thin, all of them similar in shape and volume or posterior ones are thin- ner; prothoracic papillae thinner than triom- matidium . AntVIpt 1.7—2.4 times AntVIb (exceptionally VOLUME 108, NUMBER 2 down to 1.4 in several “‘“dwarf’’ specimens). Setae on AntIIl 10—25 wm. Secondary sen- soria on AntIII mostly present in “big” spec- imens. [On Grindelia chiloensis. Shiny dark- brown to blackish-brown if “big” specimens, or opaque light-brown to dark-green if nGWatieetspecimens|=ea a mace ene oe A. melosae — AntVIpt 0.9-1.7 times AntVIb. Setae on AntIll variable in length, longer than 35 pm or shorter than 10 2m. Secondary sensoria al- WAY StADSCM Otis sesh Metin teaes whl athe 15 15. Setae on AntIll, AbdII-AbdIV and AbdVIII 7-8, 12-32, 22—45 pm. Siphunculus 0.4—0.9 times cauda. [On several species of Senecio. Opaque to shiny black] .... A. senecionicoides — Setae on Antlll, AbdII-AbdIV and AbdVIII longer than 35, 39 and 60 wm. Siphunculus aimleastaO:Sutimesicaldameryereaet ee ene ere 16 16. AntVIpt approximately 1.5 times AntVIb. Si- phunculus at least 1.3 times cauda. Cauda with 10—14 setae. [On Mulinum spinosum. Probably shiny dark brown to black] — AntVIpt approximately 1.0 times AntVIb. Si- phunculus at most 0.9 times cauda. Cauda with 8-10 setae. [On Mulinum spinosum. Probably shiny dark brown to black] ...... A. mulinicola eee en A. mulini ACKNOWLEDGMENTS We thank A. Dalmasso, E. Martinez, and E. Méndez, “Instituto Argentino de Inves- tigaciones de las Zonas Aridas”” (Mendoza, Argentina), for their help in identifying the host plant. We also thank two anonymous reviewers for comments, suggestions, and corrections. This work was funded by grants LE20/99 and LE45/02 from the re- gional government of Castilla y Leon (Spain). LITERATURE CITED Eastop, V. EK 1979a. Key to the genera of the subtribe Aphidina (Homoptera). Systematic Entomology 4: 379-388. . 1979b. Sternorrhyncha as Angiosperm Tax- onomists. Symbolae Botanicae Upsaliensis 22(4): 120-134. . 1998. Why do aphids do that?, pp 37—47. In Nieto Nafria, J. M. and A. E G. Dixon, eds. Aphids in Natural and Managed Ecosystems. Universidad de Leon. Leon. Foottit, R. G and W. R. Richards. 1993. The genera of the aphids of Canada, Homoptera: Aphidoidea and Phylloxeroidea. The Insects and Arachnids of Canada, 22. Agriculture Canada (Publication 1885). Ottawa. Fuentes-Contreras, E., R. Munoz and H. M. Niemeyer. 283 1997. Diversidad de afidos (Hemiptera: Aphidoi- dea) en Chile. Revista Chilena de Historia Natural 70: 531-542. Garcia Prieto, F and J. M. Nieto Nafria. In press. Gé- nero Aphis. In Nieto Nafria, J. M., M. PB. Mier Durante, F Garcia Prieto and N. Pérez Hidalgo: Hemiptera Aphididae III. Jn Ramos M. A. et al., eds. Fauna Ibérica, volumen 28. Museo Nacional de Ciencias Naturales (CSIC). Madrid. Hille Ris Lambers, D. 1974. On American aphids, with descriptions of a new genus and some new species (Homoptera: Aphididae). Tijdschrift voor Ento- mologie 117(4): 103-155. Mier Durante M. P. and J. Ortego. 1999. Two new species of Aphis L. (Hemiptera: Aphididae) from Argentina living on Asteraceae. Proceedings of the Entomological Society of Washington 101(2): 428-437. Mier Durante M. P., J. M. Nieto Nafria and J. Ortego. 2003. Aphidini (Hemiptera: Aphididae) living on Senecio (Asteraceae), with descriptions of a new genus and three new species. The Canadian En- tomologist 135: 187-212. Missouri Botanical Garden. 1999. w*TROPICOS. On line: http://mobot.mobot.org/W3T/Search/ vast.htIml. July 20 2004 consulted. Miyazaki M. 1987. Morphology of aphids: 1—25. In Minks, A. K. and P. Harrewijn, eds. Aphids, their Biology, Natural Enemies and Control, Volume 2A. In Helle W., ed. World Crop Pests. Elsevier. Amsterdam. Nieto Nafria, J. M. and M. P. Mier Durante. 1998. Hemiptera Aphididae I. /n Ramos M.A. et al., eds. Fauna Ibérica, volumen 11. Museo Nacional de Ciencias Naturales (CSIC). Madrid. Nieto Nafria, J. M. and J. Ortego. 2002 (1999). Aphis (A.) cinerea sp. nov. de Argentina, similar a la europea A. (A.) craccae (Hemiptera: Aphididae). Boletim da Sociedade Portuguesa de Entomologia suplemento 6: 177-183. Nieto Nafria, J. M., M. A. Delfino, and M. P. Mier Durante. 1994. La afidofauna de la Argentina, su conocimiento en 1992. Universidad de Le6n (Se- cretariado de Publicaciones). Le6n. 235 pp. Nieto Nafria, J. M., J. Ortego and M. P. Mier Durante. 1999. Three new species of Aphis (Hemiptera: Aphididae) living on Mulinum (Umbelliferae) in South America. The Canadian Entomologist 131(3): 283—292. Ortego J. 1998 (1997). Pulgones de la Patagonia Ar- gentina con la descripcion de Aphis intrusa sp. n. (Homoptera: Aphididae). Revista de la Facultad de Agronomia de La Plata 102(1): 59-80. Ortego, J. and M. P. Mier Durante. 1997. Les espéces sud-américaines d’Aphis inféodées au genre Ber- beris (Hemiptera: Aphididae). Annales de la So- ciété Entomologique de France (N.S.) 33(4): 41 1— 418. 284 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ortego, J.. M. E. Difabio and M. P. Mier Durante. 2004. Nuevos registros y actualizacion de la lista faunistica de los pulgones (Hemiptera: Aphididae) de la Argentina. Revista de la Sociedad Entomo- l6gica Argentina 63(1—2): 19-30. Remaudiére, 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. Seco Fernandez, M. V., J. Ortego, M. P. Mier Durante, and J. M. Nieto Nafria. 2000. Sobre cuatro espe- cies de la tribu Macrosiphini (Hemiptera Aphidi- dae: Aphidinae) de la Argentina. Boletin de la Asociacion Espanola de Entomologia 24(3—4): 157-164. Smith, C. EF and M. M. Cermeli. 1979. An Annotated List of Aphididae (Homoptera) of the Caribbean Islands and South and Central America. North Carolina Agricultural Research Service Technical Bulletin 259: 1-131. Zuloaga F O. and O. Morrone. 1999. Catalogo de las plantas vasculares de la Republica Argentina. II. On line: http://www.darwin.edu.ar. February 02 2005 consulted. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 285-288 REDESCRIPTION OF BETHULIA CHAMPIONELLA RAGONOT (LEPIDOPTERA: PYRALIDAE: PHYCITINAE) H. H. NEUNZIG AND M. A. SOLIS (HHN) Department of Entomology, North Carolina State University, Raleigh, NC 27695-7613, U.S.A.; (MAS) Systematic Entomology Laboratory, PSI, Agricultural Re- search Service, U.S. Department of Agriculture, % National Museum of Natural History, Smithsonian Institution, PO. Box 37012, MRC 168, Washington, DC 20013-7012, U.S.A. (e-mail: asolis@sel.barc.usda.gov) Abstract.—Based on a single female specimen, Ragonot (1888) described the genus Bethulia and its included species B. championella. Recently, additional specimens of both sexes of this small phycitine have been collected, chiefly in the Neotropics. Using this new material, the genus and species are redescribed, and the male genitalia are illustrated for the first time. Key Words: America Bethulia is a monotypic genus whose sole species B. championella was described by Ragonot in 1888 based on a single spec- imen collected in San Jaoquin, Verapaz, Guatemala. Subsequent descriptions of the species, all also based only on the type, were published by Druce (1896), Ragonot and Hampson (1901), and Heinrich (1956). Figures of B. championella were included in Druce (1896: table 64, fig. 10), and in Ragonot and Hampson (1901: plate xxxv, fig. 16), and illustrations of the wing ve- nation and genitalia were given by Heinrich (1956: figs. 123, 1120). Heinrich also es- tablished that the type was a female and not a male as stated by Ragonot (1888). Additional B. championella specimens recently have been collected in the south- western United States, Mexico, Central America, and northern South America. This material includes moths of both sexes, and it is now possible to provide a more com- plete account of the genus and species. Specimens studied came from the follow- Phycitinae, southwestern United States, Mexico, Central America, South ing sources: Instituto Nacional de Biodiv- ersidad, Santo Domingo, Costa Rica [IN- BIO]; The North Carolina State University Insect Collection, Raleigh, North Carolina, U.S.A. [NCSU]; the National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A. [USNM]; Collec- tion of Vitor O. Becker, Universidade do Brasilia, Brasil [VOB]; and The Natural History Museum, London, England [BMNH]. Bethulia Ragonot Bethulia Ragonot 1888: 37. Type species: Bethulia championella Ragonot, 1888, original designation. Description.—Head: Male antenna with basal half of shaft slightly swollen; female antenna simple; in both sexes frons with an- teriorly projecting cone of scales, vertex rough-scaled, labial palpus porrect to slight- ly oblique and extending beyond head for distance about equal to length of head, maxillary palpus short-scaled, haustellum PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Habitus of male Bethulia championella. moderately well developed and ocellus present. Forewing: Both sexes with 9 veins; R;,, completely fused with R;; M, from slightly posterior of anterodistal angle of cell; M, and M, completely fused; CuA, from posterodistal angle of cell; CuA, from before posterodistal angle of cell. Hind- wing. Both sexes with 6 veins (1A, 2A, and 3A together treated as one vein); Sc + R, completely fused with R;; M, and M, com- pletely fused with each other; CuA, from posterodistal angle of cell; CuA, from well before posterodistal angle of cell. Male ab- dominal segment 8 without scale tuft. Male genitalia (Figs 2, 3): Uncus subtriangular, broadly rounded apically; gnathos with me- dian, apical part strongly sclerotized, tonguelike; transtilla well developed with lateral arms forming a broad bridge poste- riorly; juxta U-shaped with fingerlike, seti- ferous, lateral, posteriorly-projecting lobes; valva moderatly elongate (when extending posteriorly projects beyond uncus); aedoea- gus simple, vesica unarmed, vinculum slightly longer than greatest width. Female genitalia (Fig. 4): Ostium bursae well sclerotized, broadly flaring, with convex posterior margin; ductus bursae lightly sclerotized with band of microspines pos- teriorly and scobinate anteriorly; corpus bursae oval, about as long as ostium bursae and ductus bursae combined, scobinate where it joins ductus bursae, and without signum (Heinrich’s 1956 figure 1120 shows a dark, slightly-curved, signum-like feature in the corpus bursae but close examination reveals this to be an ink smudge on one of the curved lines; also, Heinrich, in his de- scription of the female genitalia (p. 297) states “‘... there is no signum’’); ductus seminalis from about middle of corpus bur- sae. Comments.—Heinrich (1956) was of the opinion that Bethulia is closely related to Plodia Guénée and Ribua Heinrich. His as- sessment was based on similar wing vena- tion and female genitalia among the three genera. The appearance of the male geni- talia of Bethulia, in general, supports this view, particularly with regard to Plodia. Bethulia championella Ragonot (Figs. 1—4) Bethulia championella Ragonot 1888:37. Description.—Head: Frons mostly black with some white scales dorsally; vertex white with a few black scales on some VOLUME 108, NUMBER 2 specimens; labial palpus outwardly mostly, to half, white basally, black distally; max- illary palpus outwardly white to black ba- sally, black distally. Thorax: Dorsum of prothorax white with patches or spots of black. Forewing: Length 5.0—6.0 mm, up- per surface mostly white with scattered black scales; antemedial and postmedial lines white, concolorous with white of wing; small black patch at base of wing on anterior half; black, transverse line border- ing antemedial line, and a similar black line associated with postmedial line; series of black patches along distal margin of wing; discal spots black, separate; underside of wing suffused with black except for white streak along posterior margin. Hindwing: 287 Mostly white in male with brown along margins; grey in female with black along veins and wing margins; underside of both sexes suffused with black along costal mar- gin. Male and female genitalia: As de- scribed for genus. Material examined.—U.S.A.: 1 6, 1 @, Arizona, Coconino Co., Vail Lake Rd. 6,500’, 9 % mi SE Flagstaff, 11 July 1961, Ronald W. Hodges, genitalia slides 773 HHN, 774 HHN [USNM]. 3 <6, Arizona, Pima Co., Summerhaven, 7, 800’, July 2, 19985 Sept. 17,2001 Sune: 16.2002, RB. Nagle, genitalia slide 6334 HHN [USNM]. 1 3d, Arizona, Pima Co., Summerhaven, June 17, 2002, B. Nagle, genitalia slide 113, 035 JAL [USNM]. 4 3, 1 2, New 288 Mexico, Lincoln Co., Capitan Mts. near Nogel, I-VHI-1989, Wagner and Epstein, genitalia slides 3588 HHN, 3589 HHN [USNM]. MEXICO: 1 @, Chihuahua, Basa- seachic, 1,930 m, 28°10’57’N, 108°12'42’W, 6 VIII 2002, Balcazar y Solis [USNM]. 1 ?, Tamaulipas, Gomes Farias, 1,000 m, 29-31 VII 1988, Becker No. 69685, V. O. Becker and M. A. Solis, genitalia slide 6120 HHN [NCSU]. GUATEMALA: 1 2 (holotype), Verapaz, San Joaquin, 6,000’, Rogers [BMNH]. COSTA RICA: 1 2, Car- tago, Turrialba, 22-28-II-65, S. S. & W. D. Duckworth, genitalia slide 6337 HHN [USNM]. 1 36, 1 ¢&, Guanacaste Province, Estacion Cacao, Lado Suroeste del Volcan Cacao, 1,000—1,400 m, 25 Set.—11 Oct.— 1990, C. Chaves, LN323300, 375700, IN- BIO CRIOOO 590637, INBIO CRIOOO 576672, genitalia slides 4764 HHN, 4765 HHN [INBIO]. 1 3, Heredia, Refugio Vara Blanca, 6 km ENE Vara Blanca, 1,900 m, 10°11’N, 84°O7'W, 14.IV.2002, D. & M. Davis [USNM]. 1 6, Prov. Cartago, El Guarco, R. E Rio Macho, Macizo de la Muerte, Sector de la Esperanza, 2,600 m, Ape, 2002, R, Deleado; Tp de Luz, LN 185600 550000, # 67819 [INBIO]. PAN- AMA: 11 6, Cerro Campana, nr. Chica, 2- 5-IV-65, S. S. & W. D. Duckworth, geni- talia slides 6335, 6336 HHN [USNM; NCSU]. ECUADOR: 2 ¢, Santiago Mo- rona, Gualaquiza, 900 m, 19 XII 1992, V. O. Becker, genitalia slides 5864 HHN, 5933 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON HHN [VOB]. 1 °, Tungurahua, Rio Verde, 1,600 m, 26 XII 1992, Becker no. 103981, V. O. Becker, genitalia slide 5847 HHN [NCSU]. ACKNOWLEDGMENTS We thank G. S. Robinson, The Natural History Museum, London, England, for providing access to the type of Bethulia championella, and Ray Nash and Vitor Becker for making material from their col- lections available to us. We also thank R. Blinn; Ll. L. Deitz, and'D.L. Stephanijiot the Department of Entomology, North Car- olina State University, Raleigh, North Car- olina, M. G. Pogue, Systematic Entomolo- gy Laboratory, U.S. Department of Agri- culture, Washington, D. C., and J. C. Shaf- fer, George Mason University, Fairfax, Virginia, for critically reviewing drafts of the manuscript. LITERATURE CITED Druce, H. H. 1896. Lepidoptera-Heterocera, pp. 537— 592. In Godman, FE D. and O. Salvin. Biologia Centrali-Americana; or, Contributions to the knowledge of the Fauna and Flora of Mexico and Central America. Zoology: Insecta, Vol. 2. Heinrich, C. 1956. American moths of the subfamily Phycitinae. Bulletin of the United States National Museum 207: 1-581. Ragonot, E. L. 1888. Nouveaux genres et espéces de Phycitidae & Galleriidae. E. L. Ragonot, Paris. 52 Pp. Ragonot, E. L., and (completed by) G. E Hampson. 1901. Monographie des Phycitinae et des Galler- iinae. In Romanoff, N. M., ed. Mémoires sur les Lépidopteres. Paris. Vol. 8, 602 pp. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 289-296 TWO NEW ORIENTAL STEGELYTRINE LEAFHOPPER GENERA (HEMIPTERA: CICADELLIDAE) YALIN ZHANG, CONG WEI, AND M. D. WEBB (YZ, CW) Key Laboratory of Plant Protection Resources and Pest Management of Ministry of Education, Entomological Museum, Northwest A & F University, Yangling, Shaanxi, 712100 China (e-mails: yalinzh@cnipm.com or yalinzh@nwsuaf. edu. cn; Wei- cong215@163.net); (MDW) Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: M.Webb@nhm.ac.uk). Corresponding author: Cong Wei Abstract.—The following new stegelytrine leafhopper genera and species are described: Stenolora n. gen. with S. malayana n. sp. as its type species from peninsular Malaysia and S. abbreviata n. sp. from China; Platyvalvata n. gen. with P. longicornis n. sp. as its type species from Nepal and P. dalatiensis n. sp. from Vietnam. Key Words: Region Stegelytrinae Baker is a small leafhopper subfamily from the Palaearctic and Oriental regions. The group is distinguished by the following combination of characters: eyes encroaching onto pronotum laterally in dor- sal view (Figs. 1, 27, 29), lateral margin of face not or weakly incurved below eyes (Figs. 2, 26, 30), antennae arising low on face (Figs. 2, 26, 30); and forewing with crossvein between claval veins and between outer claval vein and claval suture (Figs. 3, 28, 31). By leafhopper standards, the Ori- ental stegelytrine genera are remarkably di- verse and were either only recently placed in the subfamily, being unassigned by Oman et al. (1990), or have been recently described or revised (see Webb 1999; Wei and Zhang 2003; Zhang and Wei 2002; Zhang et al. 2002, 2004). To help increase our knowledge of this little known group, we describe two new Oriental stegelytrine genera, Stenolora and Platyvalvata, together with two new species of each. The new genera form a distinct group with the other oriental stegelytrine genera, 1.e., Cyrta Melichar, Doda Distant, Hemiptera, Cicadellidae, Stegelytrinae, new genera, new species, Oriental Kunasia Distant, Placidus Distant, Placi- dellus Evans, Paraplacidellus Zhang, Wei, and Shen, and Temburocera Webb based on their very much longer antennae. They also share some other characters with some but not all Oriental genera i.e., clypellus ex- panded with lora narrow (Stenolora and Temburocera); clypellus with two apical stout setae (Stenolora, Doda, and Kunasia); lateral frontal suture extending well beyond corresponding ocellus (Platyvalvata, Cyrta, Placidus, Placidellus, and Paraplacidellus); pronotum with lateral carina strongly curved dorsally adjacent to eye (all except Temburocera); scutellum with one or two posteromedial longitudinal keels (all except Platyvalvata, Placidus, and Cyrta); fore- wing appendix well developed and hind fe- mur with extra subapical spines, elevated and mounted on strong bases (all except Temburocera). Unlike the Palaearctic genera (Stegelytra Mulsant and Rey and Wadkufia Linnavuori) that occur on oaks (Quercus), the biology and host plants of Oriental Stegelytrinae are unknown. However, the male specimen of 290 Stenolora abbreviata n. sp. (described here) and other male Stegelytrinae (Zhang in prep.) were caught on an exposed river bed. A similar environment is used by male Lep- idoptera and is connected to mineral uptake (Holloway 1984: 97), and in Cicadellidae the following observation (personal com- munication Chris Dietrich) was made in Peru: “large swarms, of leafhoppers were seen in a few places along streams and riv- ers; they were resting and feeding on the wet sand/mud. Every specimen was male and sweeping on the surrounding vegeta- tion yielded no females and very few ad- ditional males. Taxa were mainly xylem feeding Cicadellinae (including the rare Phereurhinini), commonly referred to as sharp-shooters, which were shooting (ex- creting) profusely.” Similarly, Stegelytrinae are rare in collections with many species known only from a single specimen, usually the male. Material examined is deposited in the in- stitutions abbreviated in the text as follows: BMNH (The Natural History Museum, London). BPBM (Bernice P. Bishop Museum, Hon- olulu, Hawaii). IRSNB (Institute Royal des Science Natu- relles de Belgique, Brussels). Stenolora Zhang, Wei, and Webb, new genus Type species.—Stenolora Zhang, Wei, and Webb, n. sp. Description.—Dark brown, head and tho- rax marked with yellow to whitish yellow; forewing with extensive hyaline patches. Head small, distinctly narrower than pronotum, front margin rounded; eyes en- croaching onto pronotum laterally. Vertex similar in length to basal width, sloping to front, shiny; coronal suture obscure. Face similar in length to width, shagreened; lat- eral margins very weakly insinuate below eyes; ocelli on anterior margin of vertex, situated approximately their own diameter from eye; clypeus narrow, with lateral mar- malayana PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gins subparallel below and above antennae; lateral frontal sutures extending to lateral margin of corresponding ocellus; transcly- peal suture distinct; clypellus broad, de- pressed apically, lateral margins convex ba- sally, broader basally than anteriorly, ante- rior margin slightly concave with a stout seta on each side of midline; rostrum ex- tending to apex of trochanter, labrum half length of labium; gena somewhat longitu- dinally sulcate under antennae, rugose lat- erally; lora narrow; antennal ledge distinct; antenna long, arising adjacent to lower cor- ner of eye. Pronotum about 2.5 median length, posterior margin slightly concave, lateral margin long with carina present, sharply curved to eye anteriorly. Scutellum long, basal width similar to width of head, distal two-thirds with a longitudinal medial keel. Forewing with five apical cells, mid- dle and outer subapical cells closed, inner subapical cell open; claval veins united by crossvein and a crossvein present between inner claval vein and claval suture; appen- dix moderately broad, extending to fourth apical cell; claval margin strongly elevated and crimped at apex. Legs densely setose; hind femur strongly broadened distally and slightly bowed with numerous apical setae, more distal setae elevated on strong bases; hind tibia moderately flattened and strongly bowed, several supernumeral setae present between anterodorsal and posterodorsal rows. Male pygofer side longer than broad, with several macrosetae distally, hyaline band from ventral to dorsal margin anteri- orly. Valve large, subquadrangular, articu- lated to pygofer. Xth segment short with a ventral process on each side. Subgenital plate short to moderately long, subtriangu- lar, with few short to moderately long fine setae laterally. Connective somewhat T- shaped; stem very long; arms short with weakly sclerotized fused apical extensions. Paramere with inner basal apophysis short, outer basal apophysis elongate; lateral lobe prominent, crenulate; apical process elon- gate, evenly tapered to acute apex distally; VOLUME 108, NUMBER 2 basal half swollen with a ventral heel and an inner tooth subbasally, crenulate ven- trally, with a few fine setae ventrobasally. Aedeagus simple, shaft cylindrical, elon- gate, slightly curved dorsally in lateral view with apex bifurcate; gonopore apical; basal apodeme moderately long. Dorsal connec- tive present between Xth segment and basal apodeme of aedeagus. Etymology.—Named after the very nar- row lora. The gender is feminine. Remarks.—This genus is similar to Tem- burocera in having the clypellus broad and lora narrow but differs from this genus and other Stegelytrinae genera by the ventral processes of the male Xth segment. Stenolora malayana Zhang, Wei, and Webb, new species (Figs. 1-15) Description.—Length: ¢ 5.8 mm (x1). Dark brown marked with yellow, including an irregular transverse band between ocelli on vertex and across midlength of scutel- lum; pronotum with pale irroration. Male genitalia with ventral processes of segment X falcate, swollen subapically and tapered to apex. Subgenital plate moderate- ly long. Aedeagal shaft slightly expanded apically in lateral view, evenly tapering to bifurcate apex in posterior view. Type material.—Holotype: d, Peninsular Malaysia, Pahang, Cameron Highlands, 24.vi.1936, H.M. Pendlebury (BMNH). Etymology.—Named after the type lo- cality. Remarks.—This species differs from S. abbreviata by its slightly smaller size, slightly different pale markings on the tho- rax dorsally, and by the shape of the male genitalia (see remarks under abbreviata). Stenolora abbreviata Zhang, Wei, and Webb, new species (Figs. 16—20) Description.—Length: 3, 6.5 mm (x1). Color similar to previous species but addi- tionally with three whitish spots near each eye on pronotum (medial spot largest) and 291 markings on scutellum forming discreet spots rather than a transverse band. Male genitalia with ventral processes of Xth segment falcate, evenly tapered to apex. Subgenital plate short with a group of apical moderately long setae. Style apical process extending to near apex of connec- tive, slightly swollen subbasally; preapical angle narrow. Aedeagal shaft expanded api- cally in lateral view, narrow throughout length and slightly expanded toward bifur- cate apex in posterior view. Type material.—Holotype: ¢, China, Nangling Ruyuan, Guangdong, stream, 1,500 m., 9.v.2004, P. Grootaert (IRSNB). Etymology.—Named after its short sub- genital plate. Remarks.—This species is similar to S. malayana but is slightly larger, has more discrete yellow markings on the thorax dor- sally, and in the male genitalia, the subgen- ital plate is shorter with a group of mod- erately long apical setae, the processes of the Xth segment are more tapered distally, and the aedeagal shaft is more expanded apically in lateral view and narrower with gonopore shorter in posterior view. Platyvalvata Zhang, Wei, and Webb, new genus Type species.—Platyvalvata longicornis Zhang, Wei, and Webb, n. sp. Description.—Yellow with reddish to brown markings. Head small, distinctly narrower than pronotum, front margin rounded; eyes en- croaching onto pronotum laterally; vertex slightly shorter medially than basal width; sloping to front, smooth, coronal suture ex- tending to anterior margin; ocelli on ante- rior margin of vertex, situated approximate- ly 3X their own diameter from correspond- ing eye; face similar in length to width, lat- eral margins weakly insinuate below eyes; clypeus smooth with lower area shagree- ned; lateral frontal suture extending well beyond lateral margin of corresponding ocellus; transclypeal suture distinct; clypel- lus strongly broadening anteriorly, lateral 292 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-15. Stenolora malayana. |, Head and thorax, dorsal view. 2, Face. 3, Forewing. 4, Head and thorax, lateral view. 5, Apex of hind femur, lateral view. 6, Apex of hind tibia and first hind tarsomere, lateral view. 7, Valve and left subgenital plate, ventral view. 8, Aedeagus, left lateral view. 9, Aedeagus, posterior view. 10, Hind wing. 11, Connective and right style, dorsal view. 12, Apex of right style, ventral view. 13, Male genital capsule and segment X, dorsal view. 14, Male genital capsule, left lateral view. 15, Male segment X, left lateral view. VOLUME 108, NUMBER 2 17 / 18 20 Figs. 16-20. Stenolora abbreviata. 16, Subgenital plate, ventral view. 17, Male segment X, lateral view. 18, Aedeagus, left lateral view. 19, Aedeagus, anterior view. 20, Aedeagus, posterior view. margins straight, anterior margin slightly concave medially; rostrum extending to apex of fore trochanter; labrum half length of labium; gena flat, rugose laterally; lora broad; antennal ledge absent; antenna long, extending beyond midlength of body, aris- ing adjacent lower corner of eye. Pronotum about 2.5 broader than median length, smooth; lateral margin long, lateral carina present, sharply curved to eye anteriorly. Scutellum moderately long; basal width about as broad as head and slightly longer than lateral margin, depressed medially. Forewing with five apical cells, middle and outer subapical cells closed, inner subapical cell open; claval veins united by crossvein and a crossvein present between inner cla- val vein and claval suture; appendix mod- erately broad extending to fourth apical cell; claval margin slightly elevated, crimped at apex. Legs densely setose; hind 293 femur strongly broadened distally and slightly bowed with numerous apical setae, more distal setae elevated on strong bases; hind tibia moderately flattened and strongly bowed, several supernumeral setae present between anterodorsal and posterodorsal rows. Male pygofer side longer than broad with several macrosetae distally; a lateral finger- like process dorsally, adjacent base of Xth segment. Xth segment large with a later- oventral folded area, crenulated distally. Valve very large, subquadrangular, articu- lated with pygofer. Subgenital plate mod- erately long, subtriangular, with a few short to moderately long stout setae ventrally. Connective large, T-shaped; stem sclero- tized medially, hyaline laterally; a ventral winglike expansion distally on each side; arms short. Paramere with inner basal apophysis very short, outer apophysis elon- gate; lateral lobe prominent with a few spinelike setae adjacent to apical process; apical process elongate, straight, tapering to acute apex; ventral margin crernulate over basal half and with a tooth at midlength. Aedeagus simple, shaft elongate, cylindri- cal, strongly curved dorsally and cephalad, anterodorsal margin with a series of blunt teeth over basal two-thirds; gonopore indis- tinct; basal apodeme short. Dorsal connec- tive present between Xth segment and basal apodeme of aedeagus. Etymology.—Named after the very broad male Xth sternite or valve. The gen- der is feminine. Remarks.—This genus forms a group with Cyrta, Placidus, Placidellus, and Par- aplacidellus, in having the lateral frontal sutures extending well beyond their corre- sponding ocellus. It differs from these and other stegelytrine genera in the dorsal pro- cesses of the male pygofer and lateral wing- like expansions of the connective. Platyvalvata longicornis Zhang, Wei, and Webb, new species (Figs. 21—28) Description.—Length: ¢ 6.8 mm (x1). Generally pale yellow; transclypeal suture, 294 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 24 Figs. 21-28. Platyvalvata longicornis. 21, Connective, paramere and aedeagus, in view of arrow in Fig. 22. 22, Connective, paramere and aedeagus, lateral view. 23, Genital capsule, lateral view. 24, Genital capsule and anal segment, dorsal view. 25, Valve and subgenital plate, ventral view. 26, Face. 27, Head and thorax, dorsal view. 28, Tegmen. VOLUME 108, NUMBER 2 295 Figs. 29-37. Genital capsule, lateral view. 33, Valve and subgenital plate, ventral view. 34, Genital capsule and anal segment, dorsal view. 35, Connective, pramere and aedeagus, dorsal view. 36, Aedeagus, dorsal view. 37, Connective and paramere, dorsal view. a medial spot subbasally on clypellus, gena and sublaterally on femur, marked with brown; vertex touched with red at base and apex of coronal suture and each side of cor- onal suture subbasally; clypeus with several indistinct reddish transverse bars and a dark yellowish spot subbasally centrally; prono- tum touched with red adjacent eyes and with two faint longitudinal bands medially and scutellum with a pair of reddish spots subbasally. Pygofer side elongate, dorsal process long and stout, bearing three macrosetae. Platyvalvata dalatiensis. 29, Head and thorax, dorsal view. 30, Face. 31, Right tegmen. 32, Aedeagal shaft elongate, expanded and bi- furcate apically in dorsal view, anterolateral margin strongly dentate over distal two- thirds. Type material.—Holotype: ¢, Nepal, De- partment of Agriculture. CIE. A. 3083 (BMNH). Etymology.—Named after the long py- gofer process. Remarks.—This species differs from P. dalatiensis in its paler color, longer pygofer process, and broader apex of the aedeagal shaft in dorsal view. The type is in poor 296 condition, and its markings may be paler than in fresh specimens. Platyvalvata dalatiensis Zhang, Wei, and Webb, new species (Figs. 29-37) Description.—Length: 6, 6.5 mm (x1). Generally sordid yellow; anterior margin of head with a nearly V-shaped whitish band between ocelli. Vertex with two brown spots subbasally; clypeus generally yellow- ish brown with a medial large dark brown patch ventrally and several indistinct red- dish transverse bars laterally. Pronotum touched with red laterally adjacent to eye and with a series of small blackish spots on midline basally. Scutellum yellowish brown with laterobasal area with a pair of small spots slightly basad of transverse suture dark brown. Venter generally yellowish brown with prosternum and femora heavily marked with dark brown. Fifth apical cell of left forewing indis- tinct. Pygofer process weakly developed and bearing a few fine setae. Aedeagal shaft in dorsal view slightly sinuate basally with apex slightly expanded and bifurcate in dor- sal view; apex flattened in lateral view, an- terolateral margin strongly dentate over dis- tal two-thirds. Type material—Holotype ¢, Vietnam, 1S km NW of Dalat, 1,850 m, 5.v.1960; L.W. Quate (BPBM). Etymology.—Named after its type local- ity. Remarks.—This species differs from P. longicornis in having the clypeus generally dark ochre with a nearly V-shaped white band basally and a large central blackish- ochre patch ventrally, the male pygofer pro- cess much shorter, and the aedeagal shaft in dorsal view slightly sinuate basally and less expanded apically. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ACKNOWLEDGMENTS We express our sincere thanks to the BPBM and the IRSNB for lending us the specimens on which this study is based. Thanks are also made to Patrick Grootaert, Isabella Van de Velde, and Jerome Constant for their help during a visit by MDW to IRSNB, made possible by a European Commission ABC grant. This study is sup- ported by the National Natural Science Foundation of China (grant no. 39870113), Northwest A & F University grants for out- standing faculty members, and the Excel- lent Competent Personnel Foundation. LITERATURE CITED Holloway, J. D. 1984. Notes on the Butterflies of the Gunong Mulu National Park, pp. 89-131. Jn Jermy, A. C. and K. P. Kavanagh, eds. Gunong Mulu Na- tional Park, Sarawak. An Account of its Environ- mental and Biota being the Results of The Royal Geographical Society/Sarawak Government Expe- dition and Survey 1977—1978 Part II. The Sarawak Museum Journal 30, Special Issue 2. Oman, P. W., W. J. Knight, and M. W. Nielson. 1990. Leafhoppers (Cicadellidae) A Bibliography, Ge- neric Check-list and Index to the World Literature 1956-1985. CAB International Institute of Ento- mology, Wallingford, England, 368 pp. Webb, M. D. 1999. Identity of Bythoscopus ignicans Walker, 1857 (Hemiptera: Auchenorrhyncha: Ci- cadomorpha: Cicadellidae: Stegelytrinae). Rei- chenbachia 33 (14): 111-114. Wei, C. and Y. Zhang. 2003. A new species of the genus Placidus (Homoptera: Cicadellidae: Stege- lytrinae) from Nepal. Entomotaxonomia 25(4): 91-94. Zhang, Y. and C. Wei. 2002. Study on the Oriental leafhopper genus Kunasia Distant (Homoptera: Cicadellidae). Entomotaxonomia 24(2): 83-88. Zhang, Y., M. D. Webb, and C. Wei. 2004. The Ori- ental leafhopper genus Doda Distant (Auchenor- rhyncha: Cicadellidae). Systematics and Biodiver- sity 1(3): 301-303. Zhang, Y., C. Wei, and L. Shen. 2002. A new species of Placidellus Evans and a related new genus (Ho- moptera: Cicadellidae). Entomotaxonomia 24(4): 239-244. Zhang, Y., C. Wei, and G. Sun. 2002. A systematic study on the genus Cyrta Melichar (Homoptera: Cicadellidae). Entomotaxonomia 24(1):27—44 (in Chinese with English summary). PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 297-305 AN UNUSUAL NEW BRACHYMYRMEX MAYR (HYMENOPTERA: FORMICIDAE) FROM COSTA RICA, WITH IMPLICATIONS FOR THE PHYLOGENY OF THE LASIINE TRIBE GROUP JOHN S. LAPOLLA AND JOHN T. LONGINO (JSL) Department of Entomology, National Museum of Natural History, Smithsonian Institution, RO. 37012, MRC 188, Washington, DC 20013-7012, U.S.A. (e-mail: lapollaj@ si.edu); (JTL) The Evergreen State College, Olympia WA 98505, U.S.A. (e-mail: longinoj @ evergreen.edu) Abstract.—Brachymyrmex Mayr is an exclusively New World ant genus that currently contains 38 described species. In this study, we describe Brachymyrmex nebulosus, new species from Costa Rica. The new species exhibits morphological characters suggestive of both Brachymyrmex and Myrmelachista Roger. Notes on the morphological characters that separate these two genera from each other are provided. Analysis of male genitalia suggests that Brachymyrmex is most closely related to Myrmelachista and Cladomyrma Wheeler, W.M. Previously, the African genera Aphomomyrmex Emery and Petalomyrmex Snelling were thought to be close relatives of Brachymyrmex as well, but our analysis, based on evidence from male genitalia, suggests this is not the case. The monotypic genus Pseudaphomomyrmex Wheeler, W.M, originally placed in the Formicinae and long thought to be a relative of Brachymyrmex, is transferred to the Dolichoderinae. Pseudaphomo- myrmex lacks the most obvious synapomorphy of the Formicinae, an acidopore. The species possesses a “‘dolichoderine habitus.’’ Other morphological characteristics suggest placement within the Dolichoderinae. For instance, the juncture where the mandibular masticatory margin rounds into the basal margin bears many small denticles, a mandibular feature characteristic of many dolichoderines. The structure of the petiole also suggests placement within the subfamily. Key Words: Pseudaphomomyrmex, Myrmelachista, new species, biogeography, Formi- cinae, Dolichoderinae Brachymyrmex Mayr ants are among the smallest ants in the New World. The genus contains 38 described species with dozens of additional “‘subspecies”’’, most from trop- ical America (Bolton 1995). One species, B. depilis Emery, is widespread in North America; a few are “tramp” species, wide- ly distributed by human commerce, with the remainder found in the Neotropics. In Neotropical forests, the common spe- cies of Brachymyrmex nest in a variety of small plant cavities, under epiphytes, or in the leaf litter. They seem quite generalized in choice of nest site, and the nests can be in relatively fragile or ephemeral substrates, suggesting frequent nest movement. Bra- chymyrmex species seem to feed mainly at carbohydrate sources, being common at ex- trafloral nectaries and at sugar water baits. Some species are known to tend Coccoidea (Hemiptera) in underground chambers (Wheeler 1910; Santschi 1923). Very little is known about the natural history for the vast majority of Brachymyrmex species. Taxonomic knowledge of Brachymyrmex is very limited and species boundaries are 298 not well established. The last taxonomic re- vision was by Santschi (1923). Perhaps fur- ther taxonomic work on Brachymyrmex has been discouraged by Creighton’s (1950) re- mark about the “‘miserable little genus” prone to taxonomic difficulties, with seem- ingly impossible morphological complexes of species. Emery (1925) placed Brachy- myrmex in the tribe Brachymyrmecini and there have since been several changes in the tribal classification. H6lldobler and Wilson (1990) considered Brachymyrmex part of an expanded Myrmelachistini that included Aphomomyrmex Emery, Brachymyrmex, Cladomyrma Wheeler, W.M., Myrmelach- ista Roger, Petalomyrmex Snelling, and Pseudaphomomyrmex Wheeler, W.M. Bol- ton (1995) considered a Brachymyrmecini that excluded Myrmelachista, returning Myrmelachista to is own tribe. More re- cently, however, Bolton (2003) synony- mized both the Brachymyrmecini and the Myrmelachistini into a vastly expanded Pla- giolepidini. The Plagiolepidini coupled with the Lasiini and Myrmoteratini form the lasiine tribe group (Bolton 2003). Here we describe a new Brachymyrmex from Costa Rica. In the course of discov- ering this new species, its exact generic as- signment came into question, for this pe- culiar species has morphological character- istics of both Brachymyrmex and Myrme- lachista. The size and shape of the mesosoma is very similar to Myrmelachista zeledoni Emery, a common species found sympatrically with the new Brachymyrmex, and the visual similarity in the field to Cre- matogaster Lund is shared with several montane Myrmelachista species. Nonethe- less, the antennae are 9-segmented and there is no antennal club, characters that suggested placement within Brachymyrmex. We therefore examine the generic defini- tions of both Brachymyrmex and Myrme- lachista and provide notes on their phylo- genetic relationship to each other and to other genera within the lasiine tribe group. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MATERIALS AND METHODS Specimens were examined from a num- ber of research collections and below fol- lows the list of the institutions and individ- uals’ collections that contributed to this study. BMNH: The Natural History Museum, London, United Kingdom. INBC: Instituto Nacional de Biodiver- sidad, Santo Domingo de Here- dia, Costa Rica. JTLG: J.T. Longino Collection, Ever- green State College, Olympia, WA, USA. LACM: Natural History Museum of Los Angeles County, Los Angeles, CA, USA: MCZC: Museum of Comparative Zoolo- gy, Harvard University, Cam- bridge, MA, USA. NHMB: _ Naturhistorisches Museum Ba- sel, Basel, Switzerland. USNM: National Museum of Natural History, Smithsonian Institution, Washington DC, USA. Examination and measurement of speci- mens were completed at various magnifi- cations using a light microscope (Leica Wild M10) and were recorded to the nearest 0.001 mm. All measurements are given in millimeters. Specimens were photographed using a JVC KY-F70B video camera mounted on a Leica M420 microscope and attached to an IBM Intellistation M Pro computer, on which composite images were assembled using Auto-Montage Version 3.04 software (Synoptics Ltd. 2003). Mor- phological terminology employed through- out follows Bolton (1994), with modifica- tions where noted. Anatomical abbrevia- tions are elaborated here: HW Total Length: HL+ML+GL. HL Head length: the length of the head proper, excluding the mandibles; mea- sured in full-face view from the mid- point of the anterior clypeal margin to a line drawn across the posterior mar- VOLUME 108, NUMBER 2 gin from its highest points (to accom- modate species where the posterior margin is concave). Head Width: the maximum width of the head in full-face view (excluding the portion of the eyes that extend pass the lateral sides of the head). Scape Length: the maximum straight line of the antennal scape excluding the condylar bulb. Mesosoma Length: the length of the mesosoma ( = alitrunk) in lateral view from the anteriormost point of the pronotum (including the “neck” of the pronotum) to the posteriormost point of the metapleuron. Gaster Length: the length of the gaster in lateral view from the anteriormost point of first gastral segment (third ab- dominal segment) to the posteriormost point of the acidopore. CI Cephalic Index: HW-100/HL. SI Scape Index: SL-100/HW. HW SL ML GL SYSTEMATIC TREATMENT Brachymyrmex nebulosus LaPolla and Longino, new species (Figs. 1A, B) Type material.—Holotype worker, COS- TA RICA: Puntarenas Prov.; 6 km south of Monteverde; 10°15'N, 84°49’W; 800 m; 22 June 1990 (J. Longino #4050) (LACM ENT 143546) (INBC); 1 paratype worker data same as holotype (INBC); 2 paratype workers COSTA RICA: Puntarenas Prov.; Ojo de Agua; 10°16’N, 84°50'W; 800 m; 5 July 1991 (J. Longino #2965) (INBI- OCRIO01279916) (MCZC) (USNM); 2 paratype workers COSTA RICA: Punta- renass brov-:’ Ojo sde Agua; 10°16'N, 84°50'W; 800 m; 28 July 1984 (INBI- OCRIO02281199) (BMNH) (LACM). Diagnosis.—Face smooth, with abundant erect setae; scapes surpass posterior margin of head by about length of first funicular seg- ment; metanotum deeply impressed, meso- soma hour-glass shaped; erect hairs on legs. Description.—Head dark brown, with 299 scapes and mandibles lighter brown to dusty yellow; smooth and shiny; abundant suberect to erect hairs throughout, with lon- gest along posterior margin and clypeus; posterior margin entire; antenna 9-segment- ed; scape with abundant, short suberect to erect hairs; scapes surpass posterior margin by length of first funicular segment; 3 small ocelli present, though lateral ocelli often difficult to distinguish; clypeus broad with median portion extended from margin forming a “‘lip’’; mandible with 5 teeth, api- cal and 4th (measured from apical) longest. Mesosoma dark brown; roughly hour-glass shaped; smooth and shiny, with erect hairs on pronotal dorsum; katepisternum and side of propodeum shagreened; declivity short and indistinct; legs dark brown with abun- dant suberect to erect hairs; tarsi lighter in color. Petiole short and inclined forward. Gaster dark brown, with appressed to erect hairs throughout. Measurements (from ho- lotype worker): TL: 2.90 mm; HL: 0.767 mm; HW: 0.704 mm; SL: 0.736 mm; ML: O:892;mme GL: 1-24 mm=Cl=92-" Siz 05: Etymology.—tThe specific epithet, nebu- losus, is Latin for misty or cloudy in ref- erence to the type locality being near Mon- teverde Cloud Forest Reserve and to the fact the generic designation of this species was in doubt for a time. Distribution and natural history.—This species has been collected only three times, from two nearby sites on the Pacific slope just below Monteverde Cloud Forest Re- serve in Costa Rica. Both sites are about 800 m elevation and are in the moist forest transition zone between cloud forest and lowland dry forest climate zones. Both ar- eas were mosaics of forest patches, pas- tures, scrubby vegetation, and road edges. All three collections have been of workers on vegetation in open scrubby vegetation. In the field, these ants look and behave re- markably like Crematogaster. DISCUSSION This species is easy to distinguish from other Brachymyrmex species in Costa Rica. 300 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig; 1: indicate diagnostic characters. A distinctly large Brachymyrmex, B. nebu- losus possesses the following diagnostic traits: blackish-brown and shiny, hour- glassed shaped mesonoma, and legs and scapes with erect hairs. The hour-glass shaped mesonoma could only be confused with B. santschii Menozzi, which possesses Brachymyrmex nebulosus, holotype worker. A, Head in full frontal view. B, Lateral view. Arrows a similarly shaped mesonoma, but it is not as well-defined as in B. nebulosus and the ant overall is smaller, much lighter in color (brownish yellow), and the cuticle is not shiny. Finally, B. santschii does not have erect hairs on the legs or scapes. Erect hairs on the legs and scapes is what Santschi VOLUME 108, NUMBER 2 (1923) used to define the subgenus Bryscha Santschi. The monophyly of Bryscha has been questioned (Brown 1973) and Bolton (2003) recently synonymized the subgenus under Brachymyrmex. While a phylogeny of Brachymyrmex is not within the scope of this study, it is worth noting that erect hairs on the legs and scapes are rarely observed in Brachymyrmex and may be of phyloge- netic importance. Only four other species have been reported with erect hairs on the legs and scapes (this has only been con- firmed by us for B. gaucho): B. antennatus Santschi, B. gaucho Santschi, B. microme- gas Emery, and B. pilipes Mayr. Based on worker morphology, the closest relative to B. nebulosus appears to be B. gaucho, which is known only from its type locality in Argentina. Like B. nebulosus, B. gaucho is a large, black, and shiny species. Unfortunately, the holotype of B. gaucho (and the only known specimen) has been badly damaged, with only the broken head and gaster remaining (holotype worker ex- amined [JSL], ARGENTINA: Cordoba: Unquillo, in NHMB). Nonetheless, com- parison of the holotype with B. nebulosus was possible. The main morphological dif- ference between the two species appears to be that the erect hairs on B. gaucho are much shorter than those observed on B. ne- bulosus, especially on the gaster. Brachymyrmex and Related Genera The genus Brachymyrmex is most likely to be confused with Myrmelachista. The most obvious distinction between these two genera is the presence of a 3 to 4-segment- ed antennal club in Myrmelachista. A\|- though Brachymyrmex usually possess in- crassate antennae, they never form a dis- tinct antennal club. Both genera include species with 9-segmented antennae, al- though Myrmelachista also includes some species with 10-segmented antennae (spe- cies in the synonymized subgenus Hinck- sidris [Snelling and Hunt 1975]). No Bra- chymyrmex have been recorded with 10- segmented antennae. Two other morpholog- 301 ical characteristics separate the two genera. The clypeus of Myrmelachista is typically subquadrate and compact, whereas Brachy- myrmex possess a broadly rounded (along the anterior margin) and wide clypeus. The shape of the mandible also differs between the two genera. In Myrmelachista, the mas- ticatory and basal margins form a right an- gle. Whereas in Brachymyrmex the masti- catory and basal margins form an obtuse angle (compare Figs. 1A and 2A). The uncertainty of the morphological boundaries of Brachymyrmex and Myrme- lachista underscores even greater uncertain- ty regarding their relationship to each other and other formicine genera. In the most cur- rent classification, Bolton (2003) places Brachymyrmex and Myrmelachista into the Plagiolepidini. While we do not present a phylogenetic analysis of the lasiine tribe group in this study, we do present notes on male genitalia that may reveal phylogentic relationships within the tribe group and pro- vide the basis for future morphological study. Examination of the male genitalia, in par- ticular the penis valves, reveals a potential synapomorphy for Brachymyrmex and Myr- melachista. Figs. 3A-D show that the apo- demal ridge in both Brachymyrmex and Myrmelachista species is dorsally placed, running along the dorsal margin of the penis valve. Further investigation of other poten- tially related genera found that only Clado- myrma, a genus restricted to southeastern Asia, possesses a dorsal apodemal ridge. Contrary to Bolton’s (2003) placement of Cladomyrma in the Lasiini, we hypothesize that Brachymyrmex, Cladomyrma, and Myr- melachista are closely related. Furthermore, molecular evidence from the D2 region of the 28S ribosomal RNA and EFl-alpha pro- tein-coding genes place Brachymyrmex and Cladomyrma together (Myrmelachista was not included in the analysis) (LaPolla et al. in preparation). Bolton (2003) stated that the Plagiolepidini and Lasiini might need to be combined in the future. Two other genera, Aphomomyrmex and Petalomyrmex, which are sister taxa (Snell- ing 1979; Chenuil and McKey 1996), have at various times been thought closely relat- ed to Brachymyrmex. Bolton (2003) placed them in the Plagiolepidini. Based on the placement of the apodemal ridge (Figs. 3A— H), however, the genera are not closely re- lated to Brachymyrmex, Cladomyrma, or Myrmelachista. This is interesting because earlier work based solely on worker mor- phology suggested that Myrmelachista at least was closely related to Aphomomyrmex and Petalomyrmex. For example, Snelling and Hunt (1975) stated, “There appears to be little difference between Myrmelachista PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Myrmelachista zeledoni, worker. A, Head in full frontal view. B, Lateral view. and Aphomomyrmex other than the presence of an antennal club.” The more medial placement of the apodemal ridge is difficult to assess across all other formicine genera, but the penis valve structure of Aphomo- myrmex and Petalomyrmex is very similar to that of Acropyga (Fig. 3H; LaPolla 2004), which Bolton (2003) placed in the Lasiini. In fact, molecular evidence from the D2 region of the 28S ribosomal RNA and EFl-alpha protein-coding genes place Acropyga as the sister to Petalomyrmex (Aphomomyrmex was not included in the analysis) (LaPolla et al. in preparation). VOLUME 108, NUMBER 2 100 um f——__| 100 um Fig. 3. Ectal (outer) view of penis valves. A, Brachymyrmex sp. 001. B, Brachymyrmex sp. 002. C, Myr- melachista sp. 001. D, Myrmelachista skwarrae. E, Cladomyrma maryatiae. F. Aphomomymyrmex afer. G Petalomyrmex phylax. H, Acropyga arnoldi. Arrows indicate the apodemal ridge. 304 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Pseudaphomomyrmex emeryi, holotype queen. A, Head in full frontal view. B, Lateral view. C, SEM image of head in full frontal view. VOLUME 108, NUMBER 2 Note on Pseudaphomomyrmex The monotypic genus Pseudaphomomyr- mex has been a source of confusion since its creation by Wheeler (1920). Its only spe- cies was originally described as Aphomo- myrmex emeryi by Ashmead (1905). Pseu- daphomomyrmex has been classified in var- ious tribes with little reference to the ho- lotype queen (holotype examined by JSL; label info: PHILIPPINES: Manila, coll. Robert Brown, in USNM), the only speci- men known for this species (see Bolton 2003 for a summary of various tribal place- ments of the genus). The species does not possess an acidopore (we examined the dis- articulated gaster with a scanning electron microscope) and in general the head has a *“‘dolichoderine habitus” (Figs. 4A—B). The juncture where the mandibular masticatory margin rounds into the basal margin bears many small denticles, a mandibular feature characteristic of many dolichoderines. The structure of the petiole, which is similar to the petiolar structure found in Tapinoma Foerster, also suggests placement within the Dolichoderinae. Therefore, we propose the transfer of Pseudaphomomyrmex from the Formicinae to the Dolichoderinae, subfam- ily transfer. It is possible that Pseuda- phomomyrmex 1s a synonym (junior or se- nior) of an already described genus within the Dolichoderinae; however, our prelimi- nary assessment is that Pseudaphomomyr- mex represents a distinct genus and there- fore no other changes in classification are proposed at this time. ACKNOWLEDGMENTS We thank Daniel Burckhardt (Naturhisto- risches Museum, Basel) and Roy Snelling (Natural History Museum of Los Angeles County) for specimen loans. Scott Whittaker (National Museum of Natural History) kindly provided assistance to JSL when working on the SEM. Phil Ward (University of Califor- nia, Davis) provided a helpful review that im- proved the manuscript. This work was sup- ported in part by National Science Founda- tion grant DEB-0072702 to JTL. LITERATURE CITED Ashmead, W. H. 1905. New Hymenoptera from the Philippines. Proceedings of the United States Na- tional Museum 29: 107-119. Bolton, B. 1994. Identification Guide to the Ant Gen- era of the World. Harvard University Press, Cam- bridge, Massachusetts, 222 pp. . 1995. A New General Catalogue of the Ants of the World. Harvard University Press, Cam- bridge, Massachusetts, 504 pp. . 2003. Synopsis and classification of Formi- cidae. Memoirs of the American Entomological Institute 71: 1-370. Brown, W. L. 1973. A comparison of the Hylean and Congo-West African rain forest ant faunas, pp. 161-185. In Meggers, B. J., E. S. Ayensu, and W. D. Duckworth, eds. Tropical Forest Ecosystems in Africa and South America: A Comparative Re- view. Smithsonian Institution Press, Washington, Di Ce vine SSOnpp: Chenuil, A. and D. B. McKey. 1996. Molecular phy- logenetic study of a myrmecophyte symbiosis: Leonardoxa/Ant Associations Diversify via cos- peciation? Molecular Phylogenetics and Evolution 6(2): 270-286. Creighton, W. S. 1950. The ants of North America. Bulletin of the Museum of Comparative Zoology 104: 1-585. Emery, C. 1925. Hymenoptera. Family Formicidae. Subfamily Formicinae. Genera Insectorum 183: 1-302. Ho6lldobler, B. and E. O. Wilson. 1990. The Ants. Har- vard University Press, Cambridge, Massachusetts, (B2=pp: LaPolla, J. S. 2004. Acropyga (Hymenoptera: Formi- cidae) of the World. Contributions of the Ameri- can Entomological Institute 33(3): 1—130. Santschi, EF 1923. Revue des fourmis du genre Bra- chymyrmex Mayr. Anales del Museo Nacional de Historia Natural de Buenos Aires (La Plata) 31: 650-678. Snelling, R. R. 1979. Aphomomyrmex and a related new genus of arboreal African ants. Contributions in Science Natural History Museum of Los An- geles County 316: 1-8. Snelling, R. R. and J. H. Hunt. 1975. The ants of Chile (Hymenoptera: Formicidae). Revista Chilena de Entomologia 9: 63-129. Synoptics Ltd. 2003. Auto-Montage Version 3.04 (computer software). United Kingdom: Cam- bridge. Wheeler, W. M. 1910. Ants: Their Structure, Devel- opment and Behavior. Columbia University Press, New York. . 1920. The subfamilies of Formicidae, and oth- er taxonomic notes. Psyche 27: 46—SS. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 306-311 TWO NEW SPECIES OF DENNYUS (COLLODENNYUS) CHEWING LICE (PHTHIRAPTERA: AMBLYCERA: MENOPONIDAE) FROM SWIFTLETS (APODIFORMES: APODIDAE) DALE H. CLAYTON, ROGER D. PRICE, AND KEVIN P. JOHNSON (DHC) Department of Biology, 257 South 1400 East, University of Utah, Salt Lake City, UT 84112-0840, U.S.A. (e-mail: clayton@biology.utah.edu); (RDP) 4202 Stanard Circle, Fort Smith, AR 72903-1906, U.S.A. (e-mail: rpricelice@aol.com); (KPJ) Illinois Natural History Survey, 607 East Peabody Drive, Champaign, IL 61820-6970, U.S.A. (e- mail: kjohnson @inhs.uiuc.edu) Abstract.—The new species Dennyus (Collodennyus) mimirogerorum from the Pap- uan swiftlet, Aerodramus papuensis (Rand), from Papua New Guinea and D. (C.) bartoni from the Philippine swiftlet, A. mearnsi (Oberholser), from the Philippines are described and illustrated. The genetic distinctiveness of these two species from other close relatives is also evaluated using mitochondrial DNA sequences. Key Words: Aerodramus Following a revision by Clayton et al. (1996) of the chewing lice in the meno- ponid genus Dennyus Neumann and sub- genus Collodennyus Ledger from swiftlets, we obtained two series of lice from swift- lets in Papua New Guinea and the Philip- pines. These lice represent new species and are described and illustrated herein. The characters associated with lice from Collodennyus, paraphrased from Clayton et al. (1996), are: Menoponid chewing lice with anterior head margin flattened, never smoothly rounded (Fig. 5). Prosternal plate well developed, with never more than total of 4 setae; first tarsi without claws; sternite I reduced, usually with only 2 setae. Female sternite VII fused with following sternites to form subgenital plate (Fig. 1); setae of VII typ- ically 2 + 4 + 2; each lateroposterior por- tion of subgenital plate with 4 medium to long submarginal setae (Figs. 2, 6). Male genitalia much as in Fig. 4. These features chewing lice, Collodennyus, Dennyus, Phthiraptera, Menoponidae, swiftlet, will not be repeated under the species de- scription. For the following descriptions, all mea- surements were made with an ocular mi- crometer and are given in millimeters. Ab- dominal tergal setal counts include the very long lateral setae and all posterior marginal setae between them. Abdominal sternal setae are given as either marginal or anterior, but do not include the 2 very short anterior setae on each side of sternite II and the setae in the brushes on sternites V-—VI. The female subgenital plate setae are limited only to those posterior to the portion attributed to sternite VII. Host classification below order follows that of Dickinson (2003). Partial sequences of the mitochondrial cytochrome oxidase I (COI) gene were ob- tained for both new species, and these se- quences are compared to those of close rel- atives. Methods for DNA sequencing are described in detail elsewhere (Johnson et al. 2001). VOLUME 108, NUMBER 2 Dennyus (Collodennyus) mimirogerorum Clayton, Price, and Johnson, new species (Figs. 1—5) Type host.—Aerodramus (Rand), the Papuan swiftlet. Female.—Head and thorax as in Fig. 5; abdomen as in Fig. 1. Tergal setae: I, 11— 12; If, 10-13; II, 12-14; IV_-V, 13-14; VI, 12-14; VII, 12; VIII, 12-13; median setae of tergite IV long. Marginal sternal setae: II, 12 (Fig. 3); Il, 12-15; IV, 13-16; V, 4— 6; VI, 3—4; VII, 8. Anterior sternal setae: II, 8—12; II, 7-12; IV, 11-20; V—VI, 0-2; VII, O—1. Setae in each brush on sternite V, 42-56; VI, 12—16; with posterior setae thin- ner and longer than others in brush. Total number of setae on sternite VI, 26—35. Sub- genital plate (Fig. 2) with total of 31-35 setae in portion posterior to VII; with 21— 22 marginal setae, not thicker than anterior setae and all in highly irregular double row. Anal ventral fringe of 52—57 setae, dorsal of 49-52 setae; total of 9-12 short setae anterior to ventral fringe. Dimensions: tem- ple width, 0.58—0.59; head length, 0.51— 0.52; prothorax width, 0.36—0.37; metatho- rax width, 0.61—0.62; abdomen width at segment IV, 0.83—0.90; abdomen length, 1.87—1.97; total length, 2.90—3.02; anus width, 0.27—-0.31. Male.—As in Fig. 5. Tergal setae: I, 10; II; 12-13; WI, 12-14; IV, 13-14; V—-VI, 14; VIt, 13-14; VIII, 12; median setae of ter- gite IV long. Marginal sternal setae: I-IV, 10-13; V—VI, 4—6; VII, 7-8; VIII, 6. An- terior sternal setae: H, 3—6; III, 4-8; IV, 4— 6; V—VIII, 0. Total number of anterior setae on sternites II-III, 7-14. Setae in each brush on sternite V, 37—40; VI, 11—14; with posterior setae thinner than others in brush. Genitalia (Fig. 4) with well-defined small triangular sac sclerite. Dimensions: temple width, 0.51—0.54; head length, 0.47—0.48; prothorax width, 0.32—0.33; metathorax width, 0.52; abdomen width at segment IV, 0.66; abdomen length, 1.35—1.41; total papuensis 307 length, 2.29—2.39; genitalia length, 0.68— 0.70. Type material—Holotype 2, ex Aerod- ramus papuensis, Papua New Guinea: Lo- savi Cave, 18 Nov. 2002, D. H. Clayton, SEA 396; in National Museum of Natural History, Washington, D.C. Paratypes: Ex A. papuensis, 1 3, same data as holotype; | 2, 1 3d, same except SEA 390; 1 2, same except SEA’ 391. Remarks.—This new species belongs to the distinctus species group and the elliotti species subgroup as defined in Clayton et al. (1996). The shape and chaetotaxy of the head as in Fig. 5, in conjunction with the chaetotaxy of the thorax and terminal ab- dominal segments, place D. mimirogerorum in the distinctus species group. It is clearly a member of the elliotti species subgroup on the basis of the female having a few short setae displaced anterior to the ventral anal fringe and abdominal tergites II-VI each with not more than 14 setae. The male does not show a clearcut separation, but this is not unexpected given the difficulties with male identification for other taxa of Den- nyus. The female of D. mimirogerorum is dis- tinguished from other taxa in the elliotti species subgroup by the combination of its large dimensions, the ventral anal fringe with 9-12 short setae displaced anterior to it (Fig. 2), sternum II with 8—12 anterior setae (Fig. 3), sternites V—VII with 0-2 me- dioanterior setae, the posterior subgenital plate setae in a very regular distribution and not stouter than the anterior setae (Fig. 2), and the ventral anal fringe of relatively long setae, especially laterally (Fig. 2). Ref- erence to the key to females given by Clay- ton et al. (1996) will show D. mimiroge- rorum either going to D. carljonesi Clayton et al. in the first part of couplet 8 or by- passing that and progressing to couplet 11, at which point it clearly comes out as being D. hahnae Clayton et al. The features men- tioned above will separate it from either of those options. The identification of the male is quite 308 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. 1-5, Dennyus mimirogerorum. 1, Dorsoventral female abdomen. 2, Ventral female terminalia. 3, Female sternite Il. 4, Male genitalia. 5, Dorsoventral male. 6, D. bartoni, ventral female terminalia. VOLUME 108, NUMBER 2 tenuous, with the best feature being the well-defined small triangular sclerite asso- ciated with the genital sac (Fig. 4). An at- tempt to identify the male of D. mimiro- gerorum in the key given by Clayton et al. (1996) again has it coming out either as D. carljonesi or D. hahnae. The length of the genitalia, the other dimensions, and the number of sternal setae will aid in recog- nition. An error in this key should be cor- rected. In the second half of couplet 5 for D. hahnae, it is erroneously given that the genitalia length is equal to or less than 0.62 when it should actually read as being equal to or more than 0.62. Etymology.—Dennyus mimirogerorum 1s named for the senior author’s children, Mir- iam (“‘Mimi’’) Erickson-Clayton and Roger Clayton, who assisted in the capture of the birds from which these lice were collected. Dennyus (Collodennyus) bartoni Clayton, Price, and Johnson, new species (Fig. 6) Type host.—Aerodramus mearnsi (Ob- erholser), the Philippine swiftlet. Female.—Head and thorax as in Fig. 5; abdomen, except for ventral terminalia, much as in Fig. 1. Tergal setae: I, 12; H- Ill, 12-13; IV, 14; V—VI, 13-14; VII, 12- 14; VIII, 11-12; median setae of tergite [V long. Marginal sternal setae: I, 9-11; II, OTIS; TY. Li=13:"V; 6; VI, 3-4; VIL.”8: Anterior sternal setae: I, 6—8; III, 4—6; IV, 5-13; V, O—2; VE-VII, 0. Setae in each brush on sternite V, 38—47; VI, 12—19; with posterior setae thinner and longer than oth- ers in brush. Total number of setae on ster- nite VI, 29-39. Subgenital plate (Fig. 6) with total of 25—27 setae in portion poste- rior to VII; with 14—17 marginal setae, not thicker than anterior setae and all in slightly irregular single row. Anal ventral fringe of 49-51 setae, dorsal of 39—42 setae; total of 7-8 short setae anterior to ventral fringe. Dimensions: temple width, 0.56—0.58; head length, 0.51—0.53; prothorax width, 0.34— 0.35; metathorax width, 0.58—0.60; abdo- 309 men width at segment IV, 0.75—0.85; ab- domen length, 1.81—1.88; total length, 2.79—2.92; anus width, 0.26—0.27. Male.—Unknown. Type material—Holotype 2°, ex Aerod- ramus mearnsi, Philippines: Mindanao, Gudkidnom Prov., 4 May 1999, D. H. Clay- ton, SEA 116; in National Museum of Nat- ural History, Washington, D.C. Paratypes: 2 2, same data as holotype. Remarks.—As with D. mimirogerorum, this new species belongs to the distinctus species group and the elliotti species sub- group as delimited in Clayton et al. (1996). The shape and chaetotaxy of the head as in Fig. 5, in conjunction with the chaetotaxy of the thorax and terminal abdominal seg- ments, place D. bartoni in the distinctus species group. It is clearly a member of the elliotti species subgroup on the basis of the female having a few short setae displaced anterior to the ventral anal fringe and ab- dominal tergites I-VI each with not more than 14 setae. The female of D. bartoni is separable from the other taxa in the elliotti species subgroup by the combination of its large dimensions, the ventral anal fringe with only 7—8 short setae displaced anteriorly to it (Fig. 6), sternite I] with 6—8 anterior se- tae, sternite III with 4—6 such setae, ster- nites VI—-VII without medioanterior setae, the posterior subgenital plate setae in a slightly irregular row of only 14—17 setae with longer setae medially (Fig. 6), and the anus with shorter ventral fringe setae (Fig. 6). The female identifies in Clayton et al. (1996) as for D. mimirogerorum. Etymology.—Dennyus bartoni is named for the senior author’s colleague Scott Bar- ton, Reid Park Zoo, Tucson, AZ, in recog- nition of his long-standing friendship, en- thusiasm, and support for research on birds and bird parasites. PHYLOGENETIC RELATIONSHIPS When Clayton et al. (1996) revised the species of Dennyus (Collodennyus), they placed the 23 louse taxa, all of which are 99 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON D. carljonesi forresteri ex A. francicus D. carljonesi forresteri ex A. francicus D. carljonesi fosteri ex A. elaphrus D. carljonesi fosteri ex A. elaphrus D. bartoni ex A. mearnsi Dennyus elliotti .D. bartoni ex A. mearnsi subgroup 91 D. mimirogerorum ex A. papuensis D. c. carljonesi ex A. fuciphagus vestitus D. c. carljonesi ex A. fuciphagus vestitus D. c. carljonesi ex A. fuciphagus vestitus D. kristinae ex A. s. spodiopygius D. kristinae ex A. s. spodiopygius D. kristinae ex A. s. spodiopygius Dennyus D. kristinae ex A. s. spodiopygius emersont subgroup 5 changes q Fig. 7. D. singhi ex A. spodiopygius assimilis D. singhi ex A. spodiopygius assimilis Phylogenetic tree resulting from unweighted parsimony analysis of a 379 bp. portion of the mito- chondrial cytochrome oxidase I (COI) gene for species of Dennyus in the elliotti subgroup. The tree is rooted on the D. emersoni subgroup as an outgroup, because phylogenetic analyses of morphological characters indicate that this subgroup is among the closest relatives of the e/liotti subgroup (Clayton et al. 1996). Numbers above branches indicate bootstrap support from 1,000 replicates. Branch lengths are proportional to the number of nucleotide changes reconstructed on that branch (scale indicated). D. = Dennyus; A. = Aerodramus. restricted to swiftlets (Apodidae: Apodi- nae), into 2 species groups: the distinctus group with 15 taxa in 3 subgroups and the thompsoni group with 8 taxa in 2 sub- groups. The 2 new species described above are Clearly members of the distinctus group, bringing the updated total of distinctus group louse taxa to 17. Based on morphol- ogy, D. mimirogerorum and D. bartoni can be further placed in the elliotti subgroup, bringing the total number of taxa in that subgroup to 8, all parasites of the genus Aerodramus. Analysis of a 379 base pair portion of the mitochondrial cytochrome oxidase I (COI) gene also supports the taxonomic and phy- logenetic findings based on morphology (GenBank Accession numbers DQ139292— DQ139307). Phylogenetic analysis of COI sequences supports inclusion of D. mimi- VOLUME 108, NUMBER 2 rogerorum and D. bartoni in the elliotti species subgroup (Fig. 7), and the mono- phyly of the e/liotti subgroup is supported by 91% of bootstrap replicates. The COI sequence from D. mimirogerorum is 7.4— 8.4% divergent (uncorrected pairwise di- vergence) from sequences of D. carljonesi. Sequences of D. bartoni are 7.1% divergent from D. mimirogerorum. These differences are as large as, or greater than, differences between other closely related Dennyus spe- cies. Material of D. hahnae from its type host, Aerodramus hirundinaceus (Strese- mann), is not available for DNA sequenc- ing, SO no comment can be made on the divergence of either D. mimirogerorum or D. bartoni from D. hahnae. Molecular phy- logenetic analyses of these new species (Fig. 7) also indicates that the subspecies of D. carljonesi do not form a monophyletic group, suggesting that consideration should be given to elevating these subspecies to full species. ACKNOWLEDGMENTS We thank Andy Mack, Avit Wako, Deb- orah Wright, and Ross Sinclair of the Wild- life Conservation Society for logistical as- sistance in PNG. We also thank Sarah Bush, University of Kansas, and Miriam Erick- son-Clayton and Roger Clayton for assis- tance with the fieldwork. Financial support was provided by NSF grant DEB-0107947 toy DELE DEB-O107891C tol KP, cand DEB-0118794 to D.H.C. and K.P.J. LITERATURE CITED Clayton, D. H., R. D. Price, and R. D. M. Page. 1996. Revision of Dennyus (Collodennyus) lice (Phthir- aptera: Menoponidae) from swiftlets, with de- scriptions of new taxa and a comparison of host- parasite relationships. Systematic Entomology 21: 179-204. Dickinson, E. C., ed. 2003. The Howard and Moore complete checklist of the birds of the world. 3rd edition. Princeton University Press, Princeton, New Jersey, 1,039 pp. Johnson, K. P, R. J. Adams, and D. H. Clayton. 2001. Molecular systematics of Goniodidae (Insecta: Phthiraptera). Journal of Parasitology 87: 862-— 869. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 312-334 STUDIES IN NEOTROPICAL NEOMIDA: A SYNOPSIS OF THE GENUS NEOMIDA (COLEOPTERA: TENEBRIONIDAE: DIAPERINI) FROM AMERICA NORTH OF COLOMBIA WITH NOTES ON OTHER WESTERN HEMISPHERE SPECIES CHARLES A. TRIPLEHORN Department of Entomology, Museum of Biological Diversity, The Ohio State Univer- sity, 1315 Kinnear Road, Columbus, OH 43212, U.S.A. (e-mail: ctriplhrn@ aol.com) Abstract.—The 27 species of the genus Neomida occurring in America north of Co- lombia are characterized, including synonymy, diagnosis, and distribution. A key to spe- cies and illustrations of each are presented. Neomida telecera, n. sp., from Costa Rica is described and notes on the remaining Neotropical species of the genus are discussed. All unrecognized names are listed and possible placements suggested. Four new synonymies are recognized: Hoplocephala testaceipes Pic is a synonym of N. picea (Laporte and Brullé), Neomida myllocnema Triplehorn is a synonym of N. occidentalis (Champion), Hoplocephala oblonga Chevrolat is a synonym of N. castanea (Bates), and Hoplocephala dytiscoides Chevrolat is a synonym of N. lateralis (Bates). New combinations recognized are: N. occidentalis (Champion), N. obsoleta (Champion), N. lateralis (Bates), N. hoff- manseggi (Laporte and Brullé), N. clavicornis (Champion), N. nigricornis (Champion), N. distans (Champion), and N. vitula (Chevrolat) Key Words: Neomida Latrielle has been a frustrating genus to study. Fortunately, few of them have been described from females only, but most of us working on the group ignored or were unaware of the tremendous intra- specific variation in cephalic armature in the males. This phenomenon is encountered in most “horned beetles,’ so it should come as no surprise to discover it in Neom- ida. In major males of most species of Neom- ida, the cephalic horns are rather spectac- ular structures but, in instances in which large series are available, the horns of some may be reduced to mere tubercles, and it is sometimes difficult to distinguish the sexes. The genitalia have proved to be diagnostic and in some minor males, examination of genitalia is the best means of determining the species. Tenebrionidae, Diaperini, Neomida, darkling beetles, Neotropical The females usually lack cephalic horns and often clypeal tubercles, but most of them have at least a trace of blunt tubercles at the inner margins of the eyes where the horns of the males are located. Selecting al- lotypes is risky since several species may inhabit the same fungus and the females may be very similar and difficult to asso- ciate with the males. The discovery of N. acera (Triplehorn 1994), was particularly disturbing since both males and females totally lack both ce- phalic horns and clypeal tubercles. External sexual dimorphism is likewise very subtle in N. paurocera (Triplehorn 1994), and N. obsoleta (Champion 1886). Many of the 42 species of New World Neomida described between 1776 (Fabri- cius) and 1965 (Triplehorn) were described VOLUME 108, NUMBER 2 from unique specimens or very small series. A certain amount of synonymy was inevi- table and my procrastination on this project has at least prevented me from creating more synonyms as I did with N. occiden- talis (Champion) (see below). Finally, the inaccessibility of many of the types contributed to my uncertainty and re- luctance to attempt a more extensive revi- sion at this time. I feel reasonably certain of the identity of the following species from America north of Colombia. For eight of the species recently described by me, only brief descriptive and distributional notes are presented. Institutions from which specimens were borrowed and acronyms used in the text are given in Acknowledgments. Neomida picea (Laporte and Brullé) @igs! 1; 23) Oplocephala picea Laporte and Brullé 1831:344(20). Arrhenoplita picea (Laporte and Brullé): Champion, 1886:179. Platydema piceum: Fleutiaux and Sallé 1889:424. Hoplocephala testaceipes Pic 1926:28. New synonymy. Neomida picea (Laporte and Brullé): Tri- plehorn 1965:375. The only other species of Neomida with a single median clypeal tubercle is N. suilla (Champion). The latter is smaller in size, has much smaller eyes and the pronotum has coalescent lateral punctures. The female of N. picea lacks clypeal tu- bercles, and the frontal horns (tubercles) are blunt but fairly well developed. The epi- pleura are entire, and the pronotum is broadly explanate as in the male. The type specimen of Oplocephala picea (Cartagena, Colombia) could not be located in MNHN. Champion (1886) had a male from the type locality upon which he based his identification, and I consider it correct. In the BMNH, there are two males and two females from San Geronimo, Guatemala, 313 and one male labeled “‘Cayenne.”’ The type of H. testaceipes Pic could not be located in MNHN. My above synonymy is based on the description alone which, although brief, is adequate, especially when the French Gui- ana type locality is considered. Specimens examined.—S53 from the fol- lowing localities: BRAZIL (Iguacu Falls; Nova Teutonia, Santa Catarina; Santarém). COSTA RICA (Guanacaste Prov., Volcan Miravalles; Alejuela, nr. Guayabal). FRENCH GUIANA (St. Laurent du Maroni). GUATEMALA (San Geronimo). PANAMA (Alhajuelo; Barro Colorado Island; Cerro Azul). SURINAME (Brokopondo Dist.). VE- NEZEULA (Suapure, Caura River). Hosts.—Host data on MCZC specimens from Barro Colorado Island include: Gan- oderma applanatum (Pers.)Pat., Ganoderma sp., and Trametes scabrosa (Pers.) G.Cunn. Neomida suilla (Champion) (Fig. 2) Arrhenoplita suilla Champion 1896:11. Neomida suilla (Champion): Triplehorn 1965:375; Marcuzzi 1984:87 This species differs from all other New World Neomida, except N. picea, in having a single prominent median clypeal tubercle. In N. suilla, the eyes are quite small, sep- arated ventrally by about 3X the diameter of one eye and extending medially to a point in line with the antennal base; in N. picea, the eyes are separated ventrally by about the diameter of one eye. Neomida pi- cea 18 approximately twice the size of N. suilla (2.0—2.2 mm). The type locality of this species is Kings- town, St. Vincent, West Indies. It was de- scribed from two males and one female (BMNH). In order to stabilize the name, I have selected and so labeled one of the males as lectotype. Specimens examined.—lIn addition to the types from St. Vincent, I have seen one male (of the three mentioned by Champion) from Guadeloupe (BMNH) and one male PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Neomida spp., lateral aspect of male head, 1. N. picea (Panama: Cerro Azul). scale bar = 1.0 mm. 2, N. suilla (Brazil: Recife). scale bar = 0.5 mm. 3, N. cioides (Mexico: S. L. P, El Salto Falls). scale bar = 0.5 mim. 4, N. occidentalis (Mexico: Baja California Sur, El Triunfo). scale bar = 1.0 mm. 5, N. paurocera (El Salvador, Quetzaltepec). scale bar = 1.0 mm. 6, N. obsoleta (Panama: Barro Colorado Is.). scale bar = 1.0 mm. labeled: Recife, Pernambuco, Brazil, IV- Neomida cioides (Champion): Triplehorn 1970, L. Xavierrillo (USNM). 1963-375: Neomida cioides (Champion) This species is extremely variable in re- (Figs. 3, 24) gard to cephalic armature. In the most high- Arrhenoplita cioides Champion 1886:180, ly developed males, the frontal horns are pl. 8, Fig. 9. long, forward-projecting, and curved down- VOLUME 108, NUMBER 2 ward apically, with a sharp bridge connect- ing the bases. The head is deeply excavate behind the bridge and there are usually two small but prominent acute tubercles on the bridge. The frons is almost vertical in lat- eral view. The clypeal tubercles are small, and located on the margins of the clypeus. The eyes are very small as in N. suilla and in fact, except for the armature of the head, N. cioides is very similar to that spe- cies. The pronotum is coarsely and densely, almost reticulately punctured, and the sides are broadly explanate (both ¢ and @). The antennae are usually bicolored, with the basal and three apical antennomeres dis- tinctly lighter. Specimens from the West Indies lack the two small horns on the bridge between the frontal horns, the dorsal punctation 1s much denser and the dorsal surface luster is dull- er. The two males from Brazil lack the bridge between the horns and the horns themselves are short (about half normal size). This species was described from a single male from Caldera in Chiriqui, Panama (BMNH). Specimens examined.—88 from the fol- lowing localities: BOLIVIA (Santa Cruz). BRAZIL (Maranhao, Aldeia Aracu; Para. 50 km e. of Candide; Santarém). COSTA RICA (3 km se Turrialba). DOMINICA (Clarke Hall). DOMINICAN REPUBLIC (Boca Chica; San Crist6bal) ECUADOR (Napo Limancocha). GUADELOUPE. MEXICO (Cordova; Fortin de las Flores; 9 km n. Tamazunchale; 3 km e. Xilitla; El Salto Falls, 12 km nw El Naranjo; 4 mi sw Veracruz; 2 mi. ne Bochil, Chiapas; Jct. Rts. 190 and 195, Chiapas; Landa de Matamo- ros, Queretaro). PANAMA (Barro Colorado Island; Caldera in Chiriqui; Fort Sherman; Madden Dam; Paraiso) PERU (Tambopala Prov., 15 km ne. Pto. Maldonado). SURI- NAME (Brokopondo Dist., Mazaroni Pla- teau, Brownsberg Natuurpark). TRINIDAD (Simla). Hosts.—Host data accompanying speci- mens collected by J. EK Lawrence include: S15 Auricularia polytricha (Mont.) Sacc., Gan- oderma applanatum (Pers.) Pat., Ganoder- ma sp., Polyporus concrescens Mont., P. tenuis Schwein., P. sector (Ehrenb.) Fr., P. lignosus Klotzsch, Pleurotus sp., Trametes corrugata (Pers.) Bres., and Agaricaceae. Neomida occidentalis (Champion), new combination (Figs. 4, 25) Arrhenoplita occidentalis Champion 1893: Se Neomida myllocnema Triplehorn 1965:386, pl. 6, Fig. 59. New synonymy. As Champion pointed out, “‘it is not very closely allied to any other members of the genus.” This is the only species known to me with abrupt basal pronotal angles, the entire epipleura is shared with only five oth- ers (see key), and the convex elytral inter- vals are likewise an unusual characteristic. Both males and females are armed with frontal horns; the horns of the male are much longer and are bowed inward apical- ly. Also, the frons of the male is distinctly excavate, shiny, and minutely punctured be- tween the horns. Examination of the unique type (BMNH) prompted the above synonymy. The holo- type is from Acapulco [Guerrero], Mexico and is not in very good condition, a fact mentioned by Champion(1893). Further- more, although I did not dissect it, the short, stout horns suggest that it is a female (not a male as indicated by Champion). When I described N. myllocnema, I had seen only eleven specimens, ten of which were from Baja California and the other from Mazatlan, Sinaloa. Since then, I have seen 25 more (see list below). Specimens examined.—25 from the fol- lowing localities: COSTA RICA (Guana- caste Province, Palo Verde Research Sta- tion, 24-VII-1975, J. Doyen, 2 3, CISC). HONDURAS (Francisco Morazan, Zamo- rano). MEXICO (Baja California, Sur, 7 mi nw El Triunfo, Summer, 1950, H.B. Leech, 2G 2 OSUE: 24) kinyesfodos Santos! 316 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-12. mm. 8, N. inermis (Costa Rica, n. slope Volcan Miravalles). scale bar = 0.5 mm. 9, N. bicornis (Delaware: Glasgow). scale bar = 1.0 mm. 10, N. aeneipennis (Costa Rica: Guanacaste Proy.). scale bar = 1.0 mm. 11, N. ferruginea (Georgia: Volusia County). scale bar = 1.0 mm. 12, N. punctatissima (Mexico: Jalisco). scale bar = 1.0 mm. Rancho La Burrera, 12-IX-1981, W.E. Steiner, 1 6, 2 2, WE. Steiner collection; Santa Rosa, holotype, allotypes and para- types of N. myllocnema, 3 3,3 2°, USNM); (Guerrero, Acapulco, holotype, 2°, BMNH); (Jalisco, Est. de Biologia Cha- a Neomida spp., lateral aspect of male head. 7, N. lateralis (Costa Rica: Heredia). scale bar = 1.0 mela, 28-V-1982, Atkinson and Equihua, 2 3,2 2, CEAM); (Puebla, 45 min Acatlan, 30-VII-1963, J. Doyen, 2 6, CISC); (Sin- aloa, Mazatlan, 4-IV-1918, Kusche, 3 6, USNM;; Mazatlan, 28-III-1918, 12, CISC) Host.—The series from El Triunfo was VOLUME 108, NUMBER 2 collected from Fomes applanatus (Pers.) Wallr. Neomida pentaphylloides (Champion) Arrhenoplita pentaphylloides Champion 1886:180. Neomida pentaphylloides (Champion): Tri- plehorn 1994:424 This differs from the other species of Neomida with entire epipleura in having a small barb near the base of each frontal horn, no trace of clypeal tubercles, and an excavation behind the frontal horns. It is most similar to N. deltocera (Triplehorn 1994) (see remarks under that species). There are four specimens from Capetillo and four from Cerro Zunil, Guatemala in the BMNH, all labeled syntypes. For no- menclatorial stability, I have selected and so labeled a male from Capetillo as lectotype. I have not seen any other specimens. Neomida deltocera Triplehorn Neomida deltocera Triplehorn 1994:423, Figs. 6, 14. This small species is similar to N. pen- taphylloides (Champion 1886), but does not have an excavation behind the frontal horns and it does have clypeal tubercles in the male (lacking in N. pentaphylloides). The type locality of this distinctive spe- cies is Barro Colorado Island, Canal Zone, Panama. It also occurs in Costa Rica, Su- riname, French Guiana, Brazil, and the Ca- ribbean island of Dominica. It has been col- lected on Ganoderma applanatum (Pers.) Pat., G. zonatum Murrill, G. nitidum Mur- rill, and Fomes sclerodermeus (Lev.) Sacc. I have seen several hundred specimens. Neomida acera Triplehorn Neomida acera Triplehorn 1994:426, Figs. 3S) 116: There is no external sexual dimorphism in this species; both males and females lack both frontal horns and clypeal tubercles. There is slight sexual dimorphism in the Si) following two species, both of which are similar to N. acera. The type locality of this species is Barro Colorado Island, Canal Zone, Panama. It also occurs in Costa Rica and most of the 34 specimens I have seen were collected on Fomes (= WNigrofomes) melanoporus (Mont.) Sacc. Neomida paurocera Triplehorn Neomida paurocera Triplehorn 1994:424, SS ey MOE This is similar to the following species, N. obsoleta (Champion), but is smaller, lacks clypeal tubercules in either sex, and has very tiny frontal tubercles, even in the male. No host data are available on this spe- cles. The type locality of this species is Copan Ruins, Honduras; it also occurs in El Sal- vador. Neomida obsoleta (Champion), new combination (Figs. 6, 26) Arrhenoplita obsoleta Champion 1886:178. In N. obsoleta, there is very little devel- opment of frontal armature in the male, only small tubercles on the inner margins of the eyes. The clypeal tubercles, on the other hand, are quite prominent. The female has only a slight indication of the frontal tubercles and completely lacks clypeal tu- bercles. This species was described from the State of Veracruz, Mexico (Cordoba, Jala- pa). Although I studied specimens from the type series (BMNH), I failed to select a lec- totype. Specimens examined.—133 from the fol- lowing localities: BELIZE (Toledo District, Blue Creek Village and Columbia Forest). COSTA RICA (Guanacaste and Turrialba). MEXICO (Cordoba; Jalapa; Oaxaca, 4.5 mi s. Valle Nacional; 4 km s Vera Cruz). PAN- AMA (Barro Colorado Island). 318 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 13-18. Neomida spp., lateral aspect of male head. 13, N. hoffmanseggi (Panama: Barro Colorado Is.). 14, N. castanea (Panama: Cocle Prov., El Valle). 15, N. nigricornis (Panama: Panama Prov., Las Cumbres). 16, N. lecontei (Honduras: Olancho, Muralla). 17, N. divergicornis (Mexico: Chiapas, Lagunas de Montebello). 18, N. distans (Panama: Barro Colorado Is.). Scale bars = 1.0 mm on all. Neomida heterocera Triplehorn Neomida heterocera Triplehorn 1994:421, Figs. 4, 12. The type locality of this species is Costa Rica, Guanacaste, W slope Volcan Miravalles (150 specimens in type series). One male specimen was collected at Las Cumbres, Pan- ama, 29 May, 1975 by Henk Wolda. This is the only species of Neomida known to me in which the frontal horns are asymmetrical (right horn longer and stouter than the left). VOLUME 108, NUMBER 2 319 Figs. 19-22. avalles). scale bar = 0.5 mm. 20, N. atricollis (Brazil: Rond6nia). scale bar = (Brazil: Rio de Janeiro). scaie bar = 1.0 mm. 22, N. vitula (Brazil: Rio de Janeiro). scale bar = 1.0 mm. Neomida lateralis (Bates), new combination (Figs: 7; 27) Hoplocephala lateralis Bates 1873:204. Hoplocephala dytiscoides Chevrolat 1877: 170. New synonymy. The most distinctive characteristics of this species are the large size (greater than 8 mm in length) and horizontal prosternal process. It is very similar to N. lawrencei and not greatly different from N. lecontei, both of which are among the largest in the genus. In the latter two, the frontal horns of the male are curved slightly backward api- cally whereas in N. lateralis they are per- fectly straight. Hoplocephala lateralis was described from a male and female labeled ““Colum- bia” (BMNH). For nomenclatorial stability Neomida spp., lateral aspect of male head. 19, N. telecera (Costa Rica: w slope Volcan Mir- 1.0 mm. 21, N. luteonotata I designate the male as the lectotype. The type of H. dytiscoides was not found in MNHN; the above synonymy is based on size and distribution. Specimens examined.—31 from the fol- lowing localities: COLOMBIA (d6 and @ types, BMNH). COSTA RICA (Vera Blan- ca, 7-VII-1928, 1,700 m, KF Nevermann, “in trochnem Holzigen Polyporus,” 2 d, 4 &, USNM;; Cocos Island, Wafer, II-1970, L. D. Gomez P, 2 2, USNM; Guanacaste Prov., Volcan Miravalles, 2 6, CISC; Puntarenas: Monteverde, 22—24-May-85 Gilbert, Sulli- van, Hovore, CDFA; 2 6, 3 ¢, Puntarenas, Rio Negro, 25 km ne La Union, 1,500 m, 20-Feb-1965, J. B. Karen (EMUS); 3 6, 3 2, Heredia, P. N. Braulio Carrillo, sector Cerro Cacho Negro, 15-VII-88, 2,136 m, A. Cachon (INBC). PANAMA (Chiriqui, 2.0 km w Cerro Punta, 17-VI-1973, 2,240 m, 320 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Erwin & Hevel, 3 6, 4 2, USNM; Cerro Punta, 11-I-1964, L. J. Bottimer, 1 dG, CNCI). VENEZUELA (Colonia Tovar, 3 3, 2 2 (MNHN); type series of N. dytiscoides; Caracas; Country label only, 3 2, HNC). Neomida pogonocera Triplehorn Neomida pogonocera Triplehorn 1994:422, Fies..o7 13: The most diagnostic features are the long setae at the apex of the frontal horns, the stout somewhat recurved clypeal tubercles, and the pale dorsal coloration. It is most similar to N. inermis (Champion). It has been collected on Ganoderma sp. The type locality of this species is Barro Colorado Island, Canal Zone, Panama. I have also seen specimens from Brazil, Bo- livia, Peru, and Venezuela, a total of 20 specimens. Neomida inermis (Champion) (Figs. 8, 28) Arrhenoplita inermis Champion 1886:179. Neomida inermis (Champion): Triplehorn 1965:37/5: This species is most like N. pogonocera, the only other small species of Neomida with confusedly punctured elytra in which the frontal horns of the male are long, slen- der, and convergent. It also resembles N. clavicornis in this latter respect, but that species is much larger and has punctate-stri- ate elytra. The description was based on six speci- mens collected by Champion at Zapote, Guatemala. Only three of these are still in the BMNH. I examined two of the three and both are females. I am convinced that Champion merely guessed that both sexes were represented in his small series, that he never saw a male, and presumed that both sexes lacked frontal horns, thus prompting the unfortunate specific epithet. A lectotype was not designated. Several sizeable series of this species were collected by J. E Lawrence on Hexa- gonia hydnoides (Sw.) M. Fidalgo in Pan- ama (Barro Colorado Island, Fort Sherman, and Madden Dam). Another large series (24 3, 59 2) was collected by J. T. Doyen in Guanacaste Province, Costa Rica, w. slope Volcan Miravalles, 3,000 ft. Both sexes were about equally represented. I initially thought this was an undescribed species un- til I saw the types. Since most of my taxonomic conclusions in Neomida are based on the male, I am supplying the following redescription of N. inermis, based on the description prepared when I thought it to be a new species. Male.—Small, elongate-oval, strongly convex, reddish brown, shining, head with two extremely long, slender, porrect, strongly convergent frontal horns in contact laterally with eyes and curving slightly for- ward, widely separated at base and con- spicuously hairy on apical half; entire fron- to-occipital area strongly concave, perfectly smooth and impunctate; clypeus smooth, impunctate, with two well-developed, flat- tened, porrect tubercles in contact with and smoothly continuous with anterior margin; genal shelf strongly rounded, margin re- flexed, very thin; eyes large, broadly oval and deeply emarginate anteriorly, no trace of postocular concavity; antennae yellow- ish, outer seven segments strongly trans- verse, forming rather compact club; termi- nal segment of maxillary palpus narrowly oval; mentum heavily bearded with fine, stiff, erect, pale setae. Pronotum strongly transverse, widest behind middle, all angles strongly rounded, base very feebly bisinu- ate, anterior margin strongly bisinuate with well defined median lobe; surface strongly, densely and uniformly punctured. Elytra slightly wider than pronotum, lateral mar- gins parallel, straight; surface with scat- tered, closely spaced but very shallow and inconspicuous punctures, each of which bears a very short, pale, more-or-less re- cumbent seta. Ventral surface reddish brown, legs paler; flanks of prothorax strongly, confluently punctured, prosternal process narrow between coxae, apex strongly deflexed; protibia not expanded; all VOLUME 108, NUMBER 2 321 Figs. 23-30. Aedeagi of Neomida spp., dorsal and lateral aspects. 23, N. picea. 24, N. cioides. 25, N. occidentalis. 26, N. obsoleta. 27, N. lateralis. 28, N. inermis. 29, N. punctatissima. 30, N. hoffmanseggi. tarsi clothed beneath with long, yellowish setae; metasternum strongly shining, prac- tically impunctate medially and with a well- defined median longitudinal incised line; abdominal sterna coarsely and densely punctured, densely pubescent; epipleura ab- breviated beyond last visible abdominal su- ture. Length: 2.0—2.7 mm.; width: 1.0—1.3 mm. Female.—Similar to male but without trace of cephalic armature. Clypeal suture well-defined, clypeus convex, sparsely punctate; interocular surface slightly and uniformly convex, coarsely and densely punctate; slightly swollen indications of tu- bercles at inner margins of eyes; genal shelf thicker than in male and not at all reflexed. Length: 2.2—2.9 mm.; width: 1.0—1.4 mm. Variation.—Some males have the ce- phalic armature drastically reduced. When the frontal horns are small, there is a cor- responding reduction in the size of the clyp- eal tubercles. In the most poorly developed males, the horns are merely conical tuber- cles and the clypeal tubercles are barely traceable. The fronto-occipital area is, how- ever, deeply concave and smooth as usual. The only variation in the females is in the development of the frontal tubercles. At best, these are merely slight swellings on the inner margins of the eyes but sometimes they are totally absent. 322 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Neomida bicornis (Fabricius) (Fig. 9) Hispa bicornis Fabricius 1776:215 (for complete synonymy, see Triplehorn 19652377): This is the only shining, metallic species of Neomida in America north of Mexico. It undoubtedly occurs in every state east of the Mississippi River and some of the states bordering it to the west (Texas, Kansas) and well into eastern Canada. It varies clinally in coloration; individuals from Canada and northern United States are almost invariably unicolorous green dorsally whereas those from extreme southern states almost always have the pronotum reddish. In the West In- dies, the dorsal coloration is usually blue rather than green. This phenomenon is dis- cussed in detail in my 1965 paper (pp. 378— 380). Neomida aeneipennis Triplehorn (Fig. 10) Neomida aeneipennis Triplehorn 1965:382, pl. 6, Figs. 62, 64. This species was originally determined as A. bicornis by Champion (1866). It differs from that species by its larger size, finely punctured and setose abdominal sterna, and short, obtuse clypeal tubercles. The aedea- gus is likewise diagnostic. The holotype male and allotype female from Rio Hondo, British Honduras (Belize), are deposited in BMNH. Neomida aeneipennis and N. bicornis are completely allopatric. In addition to the rec- ords already published: Mexico (San Luis Potosf, Tamaulipas, Veracruz, Yucatan), Guatemala (Capetillo, Guatemala City, Za- pote) and Nicaragua (Ometepe), I now have records of 73 more from the following lo- calities: Specimens examined.—COSTA RICA (Guanacaste Province, Taboga). EL SAL- VADOR (San Salvador; 10 mi n. Quetzal- tepeque). HONDURAS (Isla Roatan). MEXICO (Chiapas, Hidalgo, Oaxaca, Puebla, Queretaro, Quintana Roo). Neomida ferruginea (LeConte) (Fig. 11) Evoplus ferruginea LeConte 1866:128; Bates 1873:234; Horn 1870:366. Oplocephala castanea Motschulsky 1873: 467 (not Bates 1873:204). Arrhenoplita ferruginea (LeConte): Cham- pion 1886:176. Hoplocephala ferruginea: Gebien 1911: 368. Neomida ferruginea (LeConte): Triplehorn 1965:384; Marcuzzi 1984:88. This species may be recognized by the deep pits behind the eye and horn on either side of the head. Only the much larger N. lawrencei and the coarsely punctured N. punctatissima share this character. The ely- tral intervals are usually (except those from Jamaica) distinctly convex. The male holotype of E. ferruginea was described from Louisiana (MCZC type No. 4671). My remarks (1965:385), comparing N. ferruginea with N. lecontei were based on my misidentification of the latter. This was discovered when I examined the type. Specimens upon which I reported from Ja- maica, while slightly different in sculpture, punctation and luster, appear to be conspe- cific with N. ferruginea; the genitalia are identical. The name Oplocephala ferruginea Mot- schulsky was applied to a species from the East Indies. The resulting homonym was re- solved by Gebien (1910:369) who supplied the name Hoplocephala orientalis Gebien as a replacement for the East Indian species. Motschulsky is also responsible for an- other homonym involving the present spe- cies. In 1873, he used the name Oploce- phala castanea for a species described from New Orleans, Louisiana (by coincidence, Louisiana is the type locality of N. ferru- ginea LeConte 1866). There is no question about that synonymy. Also, in 1873, Bates used the name Hoplocephala castanea for a species he described from Colombia, thus that name became a homonym as well as a synonym. I have established the publication VOLUME 108, NUMBER 2 28 Figs. 31-38. Aedeagi of Neomida spp., dorsal and lateral aspects. 31, N. clavicornis. 32, N. castanea. 33, N. nigricornis. 34, N. distans. 35, N. telecera. 36, N. atricollis. 37, N. luteonotata. 38, N. vitula. dates for Bates (February) and Motschulsky (July) and, therefore, N. castanea Bates is the valid name for the Colombian species. Specimens examined.—As already re- ported (Triplehorn, 1965:386), this species occurs commonly in Alabama, Florida, Louisiana and Texas. I can now add Geor- gia to the list from the United States. South of the United States border, I have seen approximately 100 specimens from the fol- lowing localities: BELIZE (Manatee Dis- trict Toledo: District). \COSTA RICA (Guanacaste Province; Puntarenas Prov- ince; La Selva; 4 km w. Ouayabo; nr. Guayabal; Volcan Miravalles). CUBA (So- ledad). DOMINICAN REPUBLIC (Trina, near Morca). GUATEMALA (Tikal; Baja Vera Paz; Duenas). GUYANA (Augustine; Bartica). HAITI (Ennery). JAMAICA (Balaclava; Pt. Antonio). MEXICO (States of Colima; Hidalgo; Nayarit; Puebla; San Luis Potosi; Sinaloa; Tabasco; Tamaulipas; Veracruz). PANAMA (Alhajuelo; Barro Colorado Island; Bouquete; Cerro Cam- pana; Chiriqui; Fort Sherman; La Chorre- ro). Hosts.—Host data accompanying speci- mens collected in Panama and Costa Rica by J. E Lawrence include: Ganoderma ap- planatum (Pers.) Pat., G. zonatum Murrill, Ganoderma sp., and Fomes sclerodermeus (Lev.) Sacc. Neomida punctatissima (Champion) (Figs: 127 29) Arrhenoplita punctatissima Champion 18932537. Neomida_ punctatissima (Champion): Tri- plehorn 1994:422. In both the male and female of this spe- cies, the head, pronotum and elytra are very coarsely and densely, almost rugosely punc- tured. In the male, the frontal horns are widely separated (as in N. distans), broadly flattened, curved slightly inward apically and abruptly truncate; the clypeal tubercles are acute, prominent and rather widely sep- arated; another tubercle, almost equally prominent is situated on the antennary orbit between the clypeus and eye. This species was described from two males and a female from Acapulco Guer- rero, Mexico, collected by Hodge. For no- menclatorial stability, I have selected and so labeled one of the males as a lectotype. Specimens examined.—11 (6 6, 5 2)— MEXICO (Jalisco, Est. de Biol6gia Cha- mela, 28-V-1982, 100 m, Atkinson & Equi- hua, basidiocarpa de poliporo (OSUC and CEAM). Neomida lawrencei Triplehorn Neomida lawrencei Triplehorn 1994:419, Biase 2-010: This species closely resembles N. lecon- tei which has almost identical frontal horns but lacks the sharp bridge connecting the horns at the base. Also, N. lawrencei has deep postocular pits and the head and pro- notal punctures are very minute. The male aedeagus is quite diagnostic (Triplehorn 1994, Fig. 10). The type locality of this species is Osa Peninsula, 2.5 mi sw. Rincon, Puntarenas, Costa Rica. It has been collected several other places in Costa Rica as well as in Be- lize (Toledo District, Blue Creek Village), Panama (Barro Colorado Island) and Mex- ico (Oaxaca and Hidalgo). Fungus hosts in- clude Ganoderma nitidum Murrill, G. ap- planatum (Pers.) Pat., and Phellinus sp. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Neomida hoffmanseggi (Laporte and Brullé), new combination (Figs. 13, 30) Oplocephala Hoffmanseggi Laporte and Brullé 1831:346(23). This species lacks tubercles or horns on the clypeus in either sex. The anterior mar- gin of the clypeus is quadrate, narrowly re- flexed and continuous with the similarly re- flexed margins of the genae. The frontal horns of the male are thin, cylindrical and widely separated. The frontal tubercles of the female are more strongly developed than in most species of Neomida. The type specimen of this species de- scribed from French Guiana could not be located. I am basing my determination on specimens in BMNH determined, I believe, by Bates. There are 13 specimens from Ob- idos, Brazil, in BMNH. Specimens examined.—340 from the fol- lowing localities: BOLIVIA (La Paz; San José). BRAZIL (Chapada; Jatai/Salesépol- is; Santarém; Tapurucuard, Rio Negro; Amazonas, Rondonia, 63 km sw Arique- mes). COSTA RICA (25 mi sw. Rincén; Heredia). GUYANA (Bartica). MEXICO (Colima). PANAMA (Barro Colorado Is- land). PERU (Iquitos). TRINIDAD (Morne Bleu). VENEZUELA (Merida, 6 km s. Azulita; Aragua, Maracay). Hosts.—The large series (194) collected by Zetek on Barro Colorado Island, Pana- ma, were ““bred’’ from Polyperaceae; those collected by Lawrence on the same island (22) were on Ganoderma and Phellinus. Neomida clavicornis (Champion), new combination (Fig. 31) Arrhenoplita clavicornis Champion 1886: L/Gsypl., 8: JRig., 7. The abrupt six-segmented club is certain- ly distinctive. In addition, the pronotal punctation is coarse and dense but not to the degree seen in N. punctatissima. The most similar species is N. divergicornis (see remarks under that species). VOLUME 108, NUMBER 2 This species, known only from the type series (BMNH), was described from six specimens in the Sallé collection collected at Cordova, Veracruz, Mexico. I studied a male (the specimen figured by Champion) and a female from the type series and, for nomenclatorial stability, have designated and so labeled the former as lectotype. The male lectotype is in reasonably good condition except that the antennae are both broken off (the right with basal six and the left with four segments remaining). The fe- male antennae are intact and, as Champion described, with a six-segmented club. Both antennae consist of only ten segments. This is an anomaly I have observed in no other Neomida. Curiously, Champion’s illustra- tion of N. clavicornis also indicates only ten segments but the other two species illus- trated on the same page have the normal 11-segmented antennae. Champion does not mention this condition in his descrip- tion. The lectotype is precariously double- mounted and would probably fall apart if a dissection were attempted; therefore, I was unable to illustrate the genitalia. I have been perplexed by Champion’s statement following the description of JN. clavicornis, ““This is one of five allied spe- cies found by M. Sallé at Cordova.’ Does he mean that there are four more he chose not to describe or was he merely noting that five species of Neomida were collected at Cordova? Actually, he recorded six species from Cordova in the Sallé collection, four of which he described as new. Since I saw no unidentified Sallé Neomida at BMNH, I believe he was documenting that five spe- cies occur at Cordova. Neomida castanea (Bates) (Figs. 14, 32) Hoplocephala castanea Bates 1873:204 (not Motschulsky 1873:467). Hoplocephala oblonga Chevrolat 1878a: XCVII. New synonymy. Neomida oblonga Chevrolat: Maes and Merkl 1991:38. 325 The most distinctive characteristic of this species is the deep median excavation be- tween and behind the frontal horns of the male. The excavation is sharply defined be- hind in a broad are (often concealed when the head is retracted). The frontal horns, even in the most well-developed males, are relatively short, stout, and blunt as com- pared with other species of Neomida. The head is nearly impunctate (only a few wide- ly separated, fine punctures, especially on the clypeus); clypeal tubercles are small and blunt, located on the apical margin of the clypeus; cephalic horns short to moderately long, stout and blunt. The female has two large, broad tuber- cles in place of cephalic horns, the clypeal tubercles are absent and the head is finely and sparsely punctate. This is a fairly large, stout species aver- aging about 6.0 mm in length and almost 3.0 mm in width. It was described from a male and female (“‘New Grenada,”’ BMNH). For nomenclatorial stability, I have selected and so labeled the male as lectotype. The types (male and female) of H. ob- longa are glued to the same card (MNHN) and are in good condition; described from *“Parahyba (= Paraiba), Brazil. Champion (1886:176) reported this species from Chontales under that name. Specimens examined.—182 from the fol- lowing localities: BRAZIL (Paraiba; Teffe, Amazonas). COLOMBIA (Country only). COSTA RICA (Heredia Province, La Sel- va; Las Mercedes, Santa Clara; Guanacaste Prov., Volcan Miravalles; Turrialba; Puntar- enas Province, Los Alturas). ECUADOR (Rio Palenque, 47 km s. Santo Domingo). FRENCH GUIANA (St. Laurent du Maro- ni). GUYANA (Bartica; Kamakusa). NIC- ARAGUA (Chontales). PANAMA (Alha- juelo; Barro Colorado Island; Cerro Cam- pana; El Valle; Finca Lerida, near Boquete; Fort Sherman; Porto Bello). PARAGUAY (San Bernadino). PERU (Tingo Maria; Tambopata Reserve). SURINAME (Bro- kopondo Dist., Mazaroni Plateau, Browns- 326 berg Naturpark). VENEZUELA (San Esté- ban). Hosts.—Data accompanying specimens include: Fomes sclerodermeus (Lev.) Sacc., Ganoderma applanatum (Pers.) Sacc., and G. zonatum Murrill. Neomida nigricornis (Champion), new combination (Figs: 5:33) Arrhenoplita nigricornis Champion 1886: N72) I initially thought this to be a new species until I saw the type. Even the female is easy to identify since it also has bicolored anten- nae as in the male, has the clypeus sharply delineated (but lacking even tubercles) and strongly swollen. The male of this species has long clypeal horns (not merely tubercles), frontal horns are completely absent and antennal seg- ments 4—10 are black with the three basal and the apical segments light reddish brown. Neomida nigricornis is a small species, averaging about 4.0 mm in length. It is a tribute to Champion’s keen eye that he rec- ognized this as a distinct species. Females of this genus are very difficult to determine to species. This species was described from a unique female from San Gerénimo, Guatemala in BMNH. Specimens examined.—53 from the fol- lowing localities: BELIZE (Toledo Dist., 5- X-1906, Peck, 3 6, 2 2, Bowditch Collec- tion, MCZC). COSTA RICA (Puntarenas Prov., 2.5 mi sw Rinc6én, 1—7 March, 1967, 1 6, MCZC; Turrialba, 30-VHI-1966, Rob- in Andrews, | 6, MCZC; Guanacaste Prov- ince, Volcan Miravalles, 2 ¢, 3 2, CISC). GUATEMALA (San Geronimo, ¢, holo- type, BMNH; 6 mi e Esquintla, 11-VII- 1965, A. Raske, 1 6, CASC). PANAMA (Barro Colorado Island, 5-24-II-1968, J. F Lawrence, 8 6, 8 2, MCZC; same locality and collector, 11-VII-1969, 1 2, MCZC; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON same locality, IV-X-1943, J. Zetek, 1 6, 4 2, USNM;; Frijoles, 27-XI-1911, A. H. Jen- nings, 4 6, | 2, MCZC; same locality, E. A. Schwarz, 1 6,2 2, USNM; Porto Bello, 15-17-11-1911, E. A. Schwarz, 1 2, USNM; a. Campana, VIL-XIE 1938; “von ao USNM;; Las Cumbres, 21-V-1974, H. Wol- da, UV light trap, 1 6, OSUC; Madden Dam, 18-VII-1969, J. E Lawrence, 4 6, 3 2, MCZC; Fort Sherman, 2-IV-1967, J. EF Lawrence, 1 6, MCZC). Hosts.—All specimens with host data were collected in Hexagonia hynoides (Sw) M. Fidalgo by J. KE Lawrence in Panama, except those from Frijoles collected by Jen- nings from “‘flowers of Lingua de Vaca.” Neomida lecontei (Bates) (Fig. 16) Evoplus lecontei Bates 1873:233. Arrhenoplita lecontei (Bates): Champion 1886:177. Hoplocephala l\econtei (Bates): Gebien 1911:368. Neomida lecontei (Bates): Triplehorn, 1965: 375: This species was described from a single male labeled ““Colombia’”’ (in BMNH). The description by Bates is excellent. The fron- tal horns are slender, slightly divergent from base, curved backward and slightly convergent apically, with a sharp, abrupt bridge connecting the bases (Fig. 16). The postocular depression is very shallow (as compared to N. ferruginea, N. lawrencei, and N. lateralis). The pronotum is finely punctured. The female (unknown to Bates) lacks clypeal tubercles but has distinct promi- nences at the inner margins of the eyes (where horns of males are located). My comments (Triplehorn, 1965:315) comparing N. lecontei with N. ferruginea were made before I examined the type. The two are really very different: N. lecontei av- erages much larger in size, has, at most, shallow postocular pits, has the distinctive VOLUME 108, NUMBER 2 bridge uniting the bases of the frontal horns, has flat elytra intervals and the pron- otum is minutely punctured, the opposite characteristics prevailing in N. ferruginea. Specimens examined.—143 from the fol- lowing localities: BELIZE (Toledo Dist., Blue Creek Village). BRAZIL (Amapa, Porto Platon). COSTA RICA (Puntarenas, Osa Peninsula, 2.5 mi s. Rincon). DOMIN- ICAN REPUBLIC (Prov. Hato Mayor, Par. Nac. Los Haitises). GUADELOUPE. GUY- ANA (Kamakusa; Maroni; Bartica). HON- DURAS (La Muralla). JAMAICA (Balacla- va; Pt. Antonio; Mandeville). MEXICO (Cordoba, V. C.; Laguna de Montebello, €hiapas;-Est: Biol. “Los: Tuxtlas,’’ V.C.). MONTSERRAT. PANAMA (Barro Colo- rado Island). PUERTO RICO (Humacao, Luquillo Expt. Forest; near Los Marias). SURINAME (Brokopondo Dist., Mazaroni Plateau). VENEZEULA (Saupure, Caura River). Host.—The only host record is Fomes sclerodermeus (Lev.) Sacc. for specimens taken in Panama, Puerto Rico and Costa Rica. Neomida divergicornis Triplehorn (Bie. o17) Neomida_ divergicornis Triplehorn 1994: 420 Figs. 3, 11. The species closely resembles N. clavi- cornis but lacks the deep median concavity on the frons immediately behind the frontal horns. The females of the two species are likewise readily separable; in N. divergi- cornis the frons is coarsely and rugosely punctured (discretely so in N. clavicornis). The antenna of N. divergicornis has the usual eleven antennomeres; whereas N. clavicornis has only ten. The type locality of this species is 10 mi west of Panuco, Veracruz, Mexico. Since the description, I have seen three more (2 3S, 1 2) labeled as follows: 1.8 mi. w. La- gunas de Monte Bello, Chiapas, Mexico (5,000’), VII-27-66, N. Chernoff (OSUC). 327 Neomida distans (Champion), new combination (Figs. 18, 34) Arrhenoplita distans Champion 1886:178, pl. 8, Figs. 8, 8a. Males of this species have the frontal horns widely separated at the base (Fig. 18), with the area between them broad, flat, shiny and virtually impunctate. In well-de- veloped major males (none available to Champion), there is a distinct tooth, facing inward about midway between base and apex of each horn. The male also has a dis- tinct yellowish tubercle on the mentum. In the female, the cephalic horns are re- duced to mere tubercles and the clypeal tu- bercles are much smaller than in the male. A faint Y-shaped raised area is present with the two arms bridging the two frontal tubercles and the base extending posteriorly on the oc- ciput. The frontal area has fine but very sparse punctures and the mentum does not have a large median tubercle. The pronotum of the male is more convex and anteriorly abruptly declivous than that of the female. This is a large species, averaging from 5.7 to 6.5 mm in length and 2.9 to 3.2 mm in width (both sexes). There are four syntypes of this species (Cordova, Mexico, 1836, Sallé Collection, BMNH). For nomenclatorial stability, I have selected and so labeled the male fig- ured by Champion as lectotype. Specimens examined.—60 from the fol- lowing localities: BOLIVIA (El Beni, Beni Sin, Palm Camp, Alto Bosque, NE San Borja). BRAZIL (Manaus, Mann & Baker; Tefé (Ega), 1878; Porto Platon, Amapa, IX- 1957, K. Lenko; Rond6énia 62 km sw Ari- quemes). COSTA RICA (Osa Peninsula, Puntarenas, 21-28-II-1967). MEXICO (type series, Cordova). PANAMA (Barro Colo- rado Island, 6, 10-II-1968, 7-IV-1967, J. EF Lawrence). PERU (Chambireachu, Huala- ga, IV-VITII-1885, M. de Mathau; Tambo- pata Prov., 15 km ne. Pto. Maldonado; nr. Ramon Castilla, 26-II[-1974; Anapati, 80 km se. Satipo, 31 July, 1972). SURINAME 328 (Brokopondo Dist., Mazaroni Plateau, 20- 24-VII-1982, W. E. Steiner). Hosts.—Host data accompanying Pana- ma specimens collected by Lawrence in- clude: Fomes sclerodermeus (Lev.) Sacc., and Ganoderma sp. Neomida dolichocera Triplehorn Neomida dolichocera Triplehorn 1994:417, Figss 1; 9: The large size (7.5 to almost 8 mm in length), the long slender horns, and the flat, smooth head are distinctive. The female is unknown. The type locality for this species 1s Costa Rica, Heredia Province, La Selva Research Station. It is known from only 14 speci- mens, all from Costa Rica. Only one addi- tional specimen was studied since the de- scription: Guanacaste Prov., Est. Pitilla, (JOO m)--9 kines; Stas Cecnia sii 1991" Cc. Marango + P. Rica, (INBC). Neomida telecera Triplehorn, new species (Figs. 19, 35) Holotype male.—Small, elongate-oval, strongly convex, light reddish brown, shin- ing. Head with two short, blunt, cylindrical horns between and in contact with inner margins of eyes; horns porrect and subpar- allel; vertex flat, smooth, almost impuncta- te, not differentiated from frons; clypeus flat, slightly raised, almost impunctate, with two, small, widely spaced tubercles at cor- ners; Clypeal margin continuous with genae, slightly reflexed; eyes large, only slightly emarginate, with dorsal lobe much smaller than ventral lobe; antennae with distal sev- en antennomeres forming a moderately de- fined club, fourth antennomere with only anterior portion expanded; maxillary palpus elongate-oval, rounded apically; mentum convex, impunctate, sparsely setose. Pron- otum strongly convex, widest near middle, all angles obtusely rounded, anterior margin straight, posterior margin feebly bisinuate, lateral margins narrowly reflexed; surface PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON smooth, uniformly finely but not densely punctate. Elytra subequal in width to pron- otum at base, lateral margins straight and parallel, surface smooth and shiny, indis- tinctly punctate-striate, intervals flat and with punctures only slightly smaller than those of striae. Ventral surface concolorous with dorsum, hypomera finely, longitudi- nally wrinkled; prosternum smooth, finely punctate, prosternal process gradually de- flexed behind, its apex not prominent; pro- tibiae straight, not expanded apically, finely punctate and setose; all tarsi clothed be- neath with long, fine, sparse setae; meso- and metasternum rather coarsely punctate laterally, practically smooth medially; me- sosternum shallowly notched anteriorly; metasternum with median incised line deep, extending more than halfway from base to anterior margin; abdominal sterna shiny and microreticulate, finely, shallowly and sparsely punctate; epipleura abruptly abbre- viated opposite middle of last visible ab- dominal sternum. Aedeagus as in Fig. 35. Length: 3.0 mm; width: 1.3 mm. Allotype female.—Similar to male in size, coloration, and punctation; differs in lacking frontal horns and clypeal tubercles; the clypeus is raised as in the male, dis- tinctly set off from frons and deflexed an- teriorly. Length: 3.7 mm; width: 1.5 mm. Variation.—In the six males available for study, there is the usual variation in length of the frontal horns, but the range is not as great as in most species. Types.—Holotype male, allotype female, and nine paratypes (1 3, 8 ¢). Costa Rica, Guanacaste, w. slope Volcan Miravalles [no date], 3,600—3,800 ft. J. T. Doyen, ex In- onotus sp.; paratypes (9 2, 4 3), same lo- cality, collector, and host except VI-29— 1993, 2,800 ft. Holotype, allotype, and paratypes in CASC; paratypes in OSUC. Remarks.—This species is known only from the type series. Despite its small size, it resembles most closely N. distans, mainly because of the widely separated frontal horns (Fig. 19). VOLUME 108, NUMBER 2 KEY TO ADULT MALES OF NEOMIDA (NORTH 5(4’). 6(5’). TK OF COLOMBIA) Epipleura extending to apex of elytra; el- ytra usually confusedly punctured and GluSuliedhy GANS 5 eo Sea oe eee ona ee 2 Epipleura abbreviated caudad of last vis- ible abdominal suture; elytra usually punctate-striate and glabrous ........ W Clypeus with a single median apical tu- bercle Clypeus with two tubercles or horns (or none) On or near anterior margin ..... 4 Eyes large, separated ventrally by about diameter of one eye; pronotal punctures discrete laterally (Fig. 1); body length greater than 3 mm N. picea (Laporte and Brullé) Eyes small, separated ventrally by about 3x diameter of one eye; pronotal punc- tures coalescent laterally (Fig. 2); body length less than 2.5 mm (Brazil, West In- GIGS) Breese OAs Secret |: N. suilla (Champion) Elytra distinctly punctate-striate; inter- vals convex; pronotum with basal angles squared (Fig. 4) (Costa Rica, Honduras, MEXICO) merneee N. occidentalis (Champion) Elytra confusedly punctured, intervals not evident; pronotum with basal angles rounded Frontal horns long, slender, bent sharply forward (Fig. 3), joined by a sharp bridge sometimes bearing two small, acute tubercles; conspicuously setose dorsally (both pronotum and elytra); sur- face luster dull .... N. cioides (Champion) Frontal horns short, blunt, triangular; se- tae usually absent on pronotum; elytral setae short, fine Clypeal tubercles present; no excavation behind frontal horns; deeply and longi- tudinally excavate between frontal horns Bhs ONG ha os Oe N. deltocera Triplehorn Clypeal tubercles absent; transversely and deeply excavate behind frontal horns, but not deeply excavate between them (Guatemala) ets ANA Re N. pentaphylloides (Champion) Frontal horns represented by no more than blunt tubercles (as in females of most other species) (Figs. 5, 6) Frontal horns well-developed Clypeal tubercles present (Fig. 6) § ted Bick) CRE ee N. obsoleta (Champion) Clypeal tubercles absent Ventral surface of head coarsely and densely, almost rugosely punctured; fron- tal tubercles distinct (Fig. 5) (El Salvador, Honduras) ...... N. paurocera Triplehorn 10(7’). 11(10’). 12): 13(12). 14(12’). 15(14). 16(14’). 17(16). oS) N © Ventral surface of head finely and sparsely punctured; frontal tubercles barely evident, sexes indistinguishable externally (Costa Rica, Panama) ... N. acera Triplehorn Prosternal process horizontal, apex acute and prominent; eye not quite in contact with base of frontal horn (Fig. 7); size large (length greater than 7.5 mm).... N. lateralis (Bates) Prosternal process deflexed between pro- coxae, apex not prominent; eye almost always in contact with base of frontal horn; size variable but usually less than 720 tm wlen others eee eee 1] Frontal horns asymmetrical, right always longer, stouter, and expanded apically (Costa Rica, Panama) N. heterocera Triplehorn Frontal horns symmetrical ......... 12 Elytra confusedly punctured (pigment spots may give illusion of striae); frontal horns long, thin, strongly converging from base to apex (Fig. 8) ........ 13 Elytra distinctly punctate-striate; frontal hoOmMsiMotrasfabOVers ae eee ee 14 Frontal horns with dense brush of long, golden setae at apex; dorsal coloration yellowish brown; elytral setae inconspic- UOUS...... FID hee te N. pogonocera Triplehorn Frontal horns with only a few short, dark setae at apex; dorsal coloration dark red- dish brown; elytral setae short but con- spicuous (Costa Rica, Panama) ST AO SES N. inermis (Champion) Usually entire dorsal surface, but at least elytra, green, blue or with bronze-green LeMeCHONS Shin yas anne 15 Entire dorsal surface uniformly yellow- ish to dark reddish brown; luster variable he MA Bh A o Mine ae pee nai HAR 2s, 2 16 Clypeal tubercles prominent and acute (Fig. 9); median portions of abdominal sterna moderately coarsely and densely punctured, each puncture bearing a very short seta (Canada, United States, West Indies) N. bicornis (Fabricius) Clypeal tubercles short and blunt (Fig. 10); median portions of abdominal sterna finely and sparsely punctured, each puncture bearing a long, fine, recumbent seta (Mexico to Costa Rica) ete Le es A SE N. aeneipennis Triplehorn Head deeply concave behind eye and frontal shormvomtubercle wee eae 17 Head not or only shallowly concave be- hind eye and horn or tubercle Pronotum with lateral punctures fine and ee) WwW jo) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sparse, each separated by 2 to 4% their diameters N. lawrencei Triplehorn Pronotum with lateral punctures coarse and dense, each separated by less than their own diameters 18 (17’). cmFrontal horns widely separated at 19(16’). 20(19"). 21(20). 22(20’). base, strongly bowed outwardly, then converging apically, apex of horns abruptly truncate (Fig. 12); pronotum rugosely punctured laterally with punc- tures only slightly smaller medially; el- ytra subrugosely punctured, punctures of intervals not much smaller than those of striae (Mexico) N. punctatissima (Champion) Frontal horns not widely separated, por- rect, subparallel or slightly divergent, bluntly rounded apically (Fig. 11); pron- otum with medial punctures much finer and more widely separated than lateral ones; elytra with strial punctures usually in grooves with intervals convex and minutely punctured (islands and regions bordering Caribbean) N. ferruginea (LeConte) Clypeus without horns or tubercles (Fig. 13), anterior margin narrowly reflexed and continuous with genal margins which are also narrowly reflexed (wide- spread, Mexico to Brazil) N. hoffmanseggi (Laporte and Brullé) Clypeus usually with at least traces of two tubercles or horns on anterior mar- gin, margin not at all reflexed ..... 20 Frons with deep, usually abrupt median concavity behind horns .......... 21 Frons flat, without median concavity be- hind shorms'*. 2 eee, See 22 Pronotum coarsely and densely punc- tured (especially laterally), most punc- tures separated by their diameters or less; elytral striae in grooves, intervals convex; antennae 10-segmented (Mexi- CO) ety. > cpa N. clavicornis (Champion) Pronotum finely and moderately densely punctured (even laterally), most punc- tures separated by 2—4% their diame- ters; elytral striae not in grooves, inter- vals flat; antennae | 1- segmented (wide- spread, Panama to Paraguay) ..... N. castanea (Bates) Clypeus with a pair of long, thin, por- rect horns (not merely tubercles); frontal horns absent (Fig. 15); antenna with three basal and the apical segment light reddish-brown, segments 4 to 10 black (Guatemala to Panama) N. nigricornis (Champion) 24(23’). 26(24'). Clypeus witha pair of short tubercles or none; frontal horns, or at least blunt pro- jections, usually present; antennal seg- ments concolorous Frontal horns connected at base by an abrupt, narrow bridge (Fig. 16) yb iey- Acts: eabgeeoe: N. lecontei (Bates) Frontal horns distinctly separated at base, the area between them flat... . Smaller species, less than 4.0 mm in lenisth Nar ie eee oe ee 25 Larger species, greater than 5.0 mm in length Elytral punctures coarse, separated by more than their diameters; interval punctures minute and sparse, striae well defined; clypeal tubercles acute and prominent (Fig. 17) (Mexico) N. divergicornis Triplehorn Elytral punctures fine, separated by their diameters or less, interval punctures coarse and dense, striae poorly defined; clypeal tubercles obtuse, not prominent (Fig. 19) (Costa Rica) 24 26 N. telecera, new species Frontal horns extremely widely separat- ed at base, bowed slightly outward me- dially, sometimes a distinct tooth or barb about middle on inner side (Fig. 18); pronotum uniformly deflexed an- teriorly N. distans (Champion) Frontal horns not widely separated at base, extremely long and slender, sub- parallel, curved slightly caudad; prono- tum with prominent V-shaped depres- sion anteriorly (Costa Rica) N. dolichocera Triplehorn NOTES ON NEOMIDA SPECIES DESCRIBED FROM SOUTH OF COLOMBIA After resolving the correct names for the species of Neomida from north of Colom- bia, there are still a number of names in the literature which I cannot associate with bi- ological species for various reasons. In the absence of types, most of them will remain nomina dubia. Three of them are easily rec- ognized and I have prepared a brief diag- nosis and a list of localities where speci- mens have been collected. Neomida atricollis (Pic), new combination (Figs. 20, 36) Hoplocephala atricollis Pic 1926:29. VOLUME 108, NUMBER 2 Diagnosis.—Small (2.4—3.6 mm); head and pronotum piceous, elytra reddish; male with clypeus and frons impunctate, shiny, narrowly reflexed apically, with two long, acute, erect tubercles positioned laterally; cephalic horns moderately stout, tapering and erect, deeply and abruptly excavate be- tween and behind cephalic horns. Female with head coarsely and densely punctured, clypeus more finely and sparsely punctured; frons transversely swollen, with distinct tu- bercles (where horns in male are located). Elytra punctate-striate without traces of grooves, punctures of intervals coarse and dense (about 2 size of strial punctures); epi- pleura abbreviated posterior to last visible abdominal suture. Type.—Described from one female from French Guiana (MNHN). Specimens examined.—38 specimens ex- amined from the following localities: BRA- ZIL (Rond6nia, 62 km sw Ariquemes, nr Fazenda Rancho Grande, 12-IX-92, II- 1993, and 8-20-XI-1994; Tapurucuara, Amazonas, Rio Negro, 13-11-1963. C. Lin- deman). BOLIVIA (Guayaramerin, Beni, Estancia Esperanza, 29-30-XI-1966; Santa Cruz, 5-15-XI-01, Thomas and Dozier). FRENCH GUIANA (type). GUYANA (Bartica; Mazaroni-Potaro Dist., Takutu Mts., 31-XII-1982). Neomida luteonotata (Pic), new combinaton (ies 2.1.,.37)) Hoplocephala luteonotata Pic 1926:29. Hoplocephala huedepohli Kulzer 1961:214. New synonymy. Diagnosis.—Moderately large (+ 6 mm); head uniformly reddish, pronotum black, elytra concolorous with head at base, lat- erally and apically; elytral disc with dark blackish blotch (variable in size) with an extension forward to base (or nearly so) and narrowing apically. Male with two small tu- bercles on clypeus and two short cephalic horns. Female without clypeal tubercles, frons coarsely and densely punctured with 331 short prominences laterally (where horns in male are located); epipleura abbreviated be- yond last visible abdominal suture. Types.—Hoplocephala luteonotata was described from one male from French Gui- ana (MNHN). The type series of H. hue- depohli Kulzer (6 3, 4 2) in Museum Frey were unavailable to me, but the description leaves little doubt of the proposed synony- my. The type locality is Iguacu Falls [Pa- rand], Brazil, with an additional female from Nova Teutonia [Santa Catarina] Bra- zil. Specimens examined.—52 specimens from the following localities: ARGENTI- NA (El Quemado; Jujuy). BOLIVIA (Bo- yuebe to Charagua via Cueva Ingri). BRA- ZIL (Santo Antonio da Barra, Bahia; Ma- tusinhos, Minas Gerais; Rio de Janeiro; Nova Teutonia, Santa Catarina). FRENCH GUIANA (type). PARAGUAY (Anasits, Alto Parana, Puerto Bertoni). SURINAME (Brokopondo District). Neomida vitula (Chevrolat), new combination (Figs. 22, 38) Hoplocephala vitula Chevrolat 1878b:214. Diagnosis.—Moderately large (+ 5.0 mm), dark reddish brown to piceous, shiny, the elytral suture reddish. Male with two short, blunt cephalic horns and two very small and inconspicuous marginal clypeal tubercles. Female without clypeal tubercles, frons finely and sparsely punctured with short prominences laterally (where horns in male are located). Elytra finely punctate- striate, striae not in grooves; intervals mi- nutely punctate; epipleura abbreviated be- yond last visible abdominal suture. Types.—Described from a unique type (sex undetermined) from “Brasilia meri- dionalis””> (MNHN). All that remains is the right elytron, both wings, meso- and metas- ternum, both mesothoracic legs and left metathoracic leg. The elytron is reddish brown, punctate striate, with flat, finely and densely punctured intervals (from my notes 832 1980). I am basing the determination of this species on size, locality, abundance in the area (southern Brazil), and the finely punc- tured elytral intervals. Specimens examined.—127 specimens examined from the following localities: ARGENTINA (Iguacu Falls; Prov. Santia- go des Estero, Icano). BRAZIL (States of Bahia, Espirito Santo, Rio de Janeiro, Sao Paulo, Santa Catarina). FRENCH GUIANA (Charvein). SPECIES EXCLUDED FROM NEOMIDA From a study of the types, I have deter- mined that the following two species do not belong to Neomida: Hoplocephala_ ephippiata Chevrolat 1878b:214.—Described from a unique fe- male (studied by me in MNHN 1980) from **Nova Grenada” (Colombia). According to my notes, this is not a Neomida; unfortu- nately, I was not able to place it in any oth- er genus. Hoplocephala_ cavifrons Chevrolat 1878b:209.—Described from a unique specimen, sex undetermined (studied by me in MNHN 1980) from “Brasilia.” This spe- cies belongs in the genus Ulosonia Laporte. UNPLACED SPECIES The following names, listed in chrono- logical order, cannot be associated with any biological species known to me: 1. Diaperis bituberculata Olivier 1791: 274; 1795, 55:8, pl. 1, figs. a, b, described from “‘environs of Paris,” length: 2.1 mm ime): Oplocephala bituberculata (Olivier): La- porte and Brullé 1831:348, reported that this is the only species that is found around Paris, that it lives under bark, and is very rare, length: 2.8 mm long (1 % lines). Hoplocephala bituberculata (Olivier): Chevrolat 1878b:210, pointed out that the Paris locality was in error; it was found in an exotic boletus of unknown origin. Both the original Olivier description and illustrations and the longer redescription by Laporte and Brullé are inadequate to vali- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON date this name. I have seen none of the specimens upon which this name is based. I am not certain whether all of the authors listed above saw the same specimens, nor am I aware that this species is still to be found in France. It will probably always re- main a nomen dubium. 2. Oplocephala_ chalybea Laporte and Brullé 1831:341. Type not seen; described from ‘Philadelphia (Amérique du Nord).” From the size (Length: 4 lines = 8.46 mm), it must be something other than Neomida or else not from North America. Horn (1870:380) did not recognize it and stated that it is “probably not North American.” 3. Oplocephala capra Laporte and Brul- lé 1831:345. Type not seen; described from Jamaica. This is almost certainly a synonym of N. bicornis (Fab.). Marcuzzi (1984) re- ported it as a valid species from Jamaica. Length: 4.23 mm (2 lines). 4. Oplocephala armata Laporte and Brullé 1831:345. Type not seen; described from Cayenne. Eroplus (sic) armatum (La- porte and Brullé): Chevrolat 1878b:210. Marcuzzi (1984) listed this as a valid spe- cies from Cuba and French Guiana. He also listed N. armata var.curticornis (Pic 1925: 29) from French Guiana, but it is not clear if he considered it valid or not. 5. Oplocephala collaris Laporte and Brullé 1831:347. Type not seen; sex not mentioned, but presumably a male. From the size given (2.5 lines = 5.3 mm), it can- not be a synonym of N. bicornis, the only member of the genus that occurs in north- eastern United States. Horn (1870:380) states that it is “unknown to us.”’ The orig- inal description states that this species was from the collection of M. Dupont, which was received from Philadelphia, not that it was collected at Philadelphia. 6. Oplocephala quadrituberculata Cur- tis, 1845:468. Type not seen; described from a pair collected at “‘St. Pauls’’ (= Sao Paulo), Brazil. Only two species known to me are as large as the size given (length: 3 ¥% lines = 7.9 mm): N. lateralis and N. dol- ichocera, neither of which are known from VOLUME 108, NUMBER 2 Brazil. I have no idea what this name rep- resents. 7. Hoplocephala amazonica Bates 1873: 203. Type not seen; described from a unique female from Santarém, Lower Am- azons, Brazil. Length: 4.7 mm (2 % lines). From the stated size, color, head, and elytral structure and type locality, I am reasonably certain that this is a synonym of N. hoff- manseggi (Laporte and Brullé). 8. Hoplocephala lutea Chevrolat 1878a: XCVII. The type is represented only by an empty pin (MNHN) and was published as being from Venezuela. Champion (1886: 177) recorded this species from Esperanza and Jalapa, Mexico, but his identification was based on Chevrolat’s description. I did not see Champion’s Mexican specimens. The size given in the original description (length: 7.5 mm) is larger than any Mexican Neomida known to me. This is very prob- ably a synonym of N. lateralis, based on size, description, and type locality. 9. Hoplocephala convexa Pic 1926:28. Type not seen; described from French Gui- ana. Length: 4.0 mm. Pic places it near N. distans, but I cannot confirm his decision. 10. Hoplocephala testaceipes Pic 1926: 28. Type not seen; described from French Guiana. Length: 4.0 mm. Pic places it near N. convexa. 11. Hoplocephala minuta Pic 1926:28. Type not seen; described from French Gui- ana. Length 3.0 mm. Pic places it near N. cioides which is widespread and quite var- iable in male cephalic armature. This is quite probably a synonym of WN. cioides. 12. Hoplocephala armata curticornis Pic 1926:29. Type not seen; described from a single male from “‘Guyane” (a nine word description, no measurements given). I have no idea what this is. 13. Hoplocephala_ atricollis ‘Kulzer 1961-215. Type not seen; described from one male, 2 females from Nova Teutonia, Santa Catarina, Brazil. From the descrip- tion, this appears to be a synonym of N. luteonotata (Pic) and also a primary hom- onym of Hoplocephala atricollis Pic. 388 ACKNOWLEDGMENTS Only five other collections have been stud- ied since my 1994 paper describing eight new species of Neomida was published. All of the institutions, their curators, and other individ- uals who loaned specimens for that study are acknowledged in that paper. I appreciate the cooperation of the fol- lowing in supplying specimens contributing to the present report: Colecci6n Entomolo- gia, Colegio de Postgraduados (CEAM), Chapingo, Mexico, Thomas H. Atkinson; Hungarian National Collection (HNC), Bu- dapest, Hungary, Otto Merk]; Instituto Na- cional de Biodiversidad, (INBC), Santo Do- mingo de Heredia, Costa Rica, Angel Solis; Muséum National d’Histoire Naturelle (MNHN), Paris, France, Claude Girard; Museu Paraense Emilio Goeldi (MPEG), Belém, Para, Brazil, William L. Overal and Ana Yoshi Harada. Acronyms for other collections men- tioned are as follows: BMNH. The Natural History Museum. London. CASC. California Academy of Sciences. San Francisco. CDFA. California Department of Food and Agriculture. Sacramento. CISG: California Insect Survey, Univer- sity of California. Berkeley. CNCI. Canadian National Collection of Insects, Ottawa, Ontario. EMUS. Entomology Museum, Utah State University, Logan. MCZC. Museum of Comparative Zoolo- gy, Harvard University, Cam- bridge, Massachusetts. OSUC. The Ohio State University Insect Collection, Museum of Biologi- cal Diversity, Columbus, Ohio. USNM. _ Department of Entomology, Na- tional Museum of Natural His- tory, Smithsonian Institution, Washington, DC. I am especially grateful to Kathy Royer for her painstaking efforts to ensure the ac- 334 curacy of my scientific writing despite my atrocious penmanship. Bruce W. Triplehorn contributed to the final stages of editing the manuscript. My colleague, Norman EF Johnson, whose skill and patience are obvious, took the outstanding photographs of these small beetles. His efforts immeasurably enhance the value of this paper and are deeply ap- preciated. Genitalia were drawn by Traci Temple except Figs. 23 and 35, which were pre- pared by Josiah D. Triplehorn. LITERATURE CITED Bates, E 1873. Notes on Heteromera, and descriptions of new genera and species. The Entomologist’s Monthly Magazine 9: 201—204, 233-238. Champion, G. C. 1886. Biologia Centrali- Americana, Insecta, Coleoptera (Tenebrionidae), Vol. 4(1): 137-264. . 1893. Biologia Centrali-Americana, Insecta, Coleoptera. (Supplement to Heteromera) Vol. 4(1): 525:572. . 1896. On the heteromerous Coleoptera of St. Vincent, Grenada, and the Grenadines. Transac- tions of the Entomological Society of London, pp. 1—54, illus. Chevrolat, L. A. A. 1877. Diagnoses de diapérides nouveaux. Petites Nouvelles Entomologiques 2: 170. . 1878a. Diagnoses especes nouvelles de dia- pérides. Annales de la Société Entomologique de Belgique 21: XCVII-C. 1878b. Diagnoses de diapérides nouveaux. Petites Nouvelles Entomologiques 2: 209-210, 214-215. Curtis, J. 1845. Descriptions, etc. of the insects col- lected by Captain P. P. King, R. N., ER. S. & L. S., in the survey of the straits of Magellan. The Transactions of the Linnean Society of London 19: 441-475. Fabricius, J. C. 310 pp. 1776. Genera Insectorum, Chilonii, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fleutiaux, E. and A. Salle. 1889. Liste des Coleopteres de la Guadeloupe et déscriptions d’esleces nou- velles. Annales de la Société Entomologique de France 6(9): 351—484. Gebien, H. 1911. Coleopterorum catalogus, pars 37, Tenebrionidae, IV, pp. 355-585. Horn, G. H. 1870. Revision of the Tenebrionidae of America, north of Mexico. Transactions of the American Philosophical Society, Ser. 2, 14: 253: 404. Kulzer, H. 1961. Neue Tenebrioniden aus Siidamerika (Col.) Entomologische Arbeiten aus dem Museum G. Frey Munich 12: 205-234. Laporte, E L. and G. A. Brullé. 1831. Monographie du genre Diaperis. Annales des Sciences Naturel- les 23: 325-410, pl. 10. LeConte, J. L. 1866. New species of North American Coleoptera, 1 (concl.). Smithsonian Miscellaneous Collections 6(167): 87-168. Maes, J. M. and O. Merkl. 1991. Catologo de los Te- nebrionoidea (Coleoptera de Nicaragua). Revista Nicaragtiense de Entomologia 17: 19-52. Marcuzzi, G. 1984. A catalogue of tenebrionid beetles (Coleoptera: Heteromera) of the West Indies. Folia Entomologica Hungarica Rosartani Kozlemenyak. XLV. 1: 69-108. Motschulsky, Victor von. 1873. Enumération des nou- velles espéces de coléopteres rapportés de ses voyages. Bulletin de la Societe Imperiale des Na- turalistes de Moscou 46(2): 466—482. Olivier, A. G. 1791. Encyclopédie Méthodique. His- toire Naturelle. Insectes, Vol. 6(pt. 1): 1-368. Par- is. . 1795. Entomologie, ou Histoire Naturelle des Insectes, Coleopteres, vol. 3, no. 55 (Diapére). Paris. Pic, M. 1926. Nouveautés diverses. Mélanges Exotico- Entomologiges fasc. 46: 1—32. Triplehorn, C. A. 1965. Revision of Diaperini of America north of Mexico, with notes on extralim- ital species (Coleoptera:Tenebrionidae). Proceed- ings of the United States National Museum. No. 3515: 349-458. . 1994. Studies in Neotropical Neomida; de- scriptions of eight new species (Coleoptera: Te- nebrionidae). Proceedings of the Entomological Society of Washington 96(3): 417—427. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 335-340 STUDIES ON STONEFLIES (PLECOPTERA) OF COLORADO WITH EASTERN FAUNAL AFFINITIES, INCLUDING A NEW STATE RECORD OF THE MIDWESTERN SALMONEFLY, PTERONARCYS PICTETIT HAGEN (PLECOPTERA: PTERONARCYIDAE) ROBERT E. ZUELLIG, BORIS C. KONDRATIEFF, AND ROBERT W. Hoop (REZ) U.S. Geological Survey, Denver Federal Center, MS 415, Denver, CO 80225, U.S.A. (e-mail: rzuellig@usgs.gov); (BCK) Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, CO 80523, U.S.A.; (RWH) U.S. Geological Survey, National Water Quality Laboratory, Denver, CO 80225, U.S.A. Abstract.—Pteronarcys pictetii Hagen nymphs were collected and reared from the South Platte River at Julesburg in eastern Colorado. Including P. pictetii, eight species are now known from Colorado that exhibit eastern North American affinities, Paracapnia angulata Hanson, Taeniopteryx burksi Ricker and Ross, Taeniopteryx parvula Banks, Acroneuria abnormis (Newman), Perlesta decipiens (Walsh), [soperla bilineata (Say), and Isoperla marlynia (Needham and Claassen). A brief discussion of the dispersal of these species into Colorado is presented. Key Words: Currently, 86 species of stoneflies have been recorded from Colorado (Kondratieff and Baumann 2002). Included in this list are seven species, Paracapnia angulata Hanson, Taeniopteryx burksi Ricker and Ross, T. parvula Banks, Acroneuria abnor- mis (Newman), Perlesta decipiens (Walsh), Isoperla bilineata (Say), and I. marlynia (Needham and Claassen), that primarily have an eastern North American distribu- tion. During a recent survey of the South Platte River in eastern Colorado, four Pter- onarcys pictetii Hagen nymphs were col- lected and reared from the South Platte Riv- er at Julesburg (Fig. 1). Including P. pic- tetii, eight species, are now known from Colorado that exhibit eastern North Amer- ican faunal affinities. Specimens were examined from the hold- ings of the C. P. Gillette Museum of Arthro- pod Diversity, Colorado State University, Fort Collins, Colorado (CSUC), U. S. Geo- logical Survey National Water Quality Lab- oratory, Denver, Colorado (NWQL), and II- Plecoptera, Pferonarcys pictetii, Colorado linois Natural History Survey, Champaign, [llinois (INHS). Additional records were provided by the Monte L. Bean Museum, Brigham Young University, Provo, Utah (BYUC), or were taken from the literature. Paracapnia angulata Hanson 1961 Paracapnia angulata is a common spe- cies typically associated with eastern North America (Stark and Baumann 2004), but also can be found in the Black Hills of South Dakota and Wyoming (Huntsman et al. 1999). This species also extends into northern Colorado on both sides of the Continental Divide. The Missouri River Corridor supports the eastern dispersal of this species into the North Platte River Ba- sin where it can be abundant (Kondratieff and Baumann 2002), or into the South Platte River Basin. The presence of this species in the upper Colorado River Basin may be the result of a naturally occurring headwater transfer between the upper South 336 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON North Platte Yampa River River Basin Basin White River Basin Colorado River Basin Dolores River Basin San Juan River Basin Fig. 1. South Platte River Basin Julesburg Republican River Basin peat Smoky Hill River Basin Arkansas River Basin Rio Grande River Basin 210 280 Kilometers Map of the state of Colorado and its counties illustrating the 10 major river basins that flow out of the state and the closed basin of the San Luis Valley (basin boundaries defined by bolded lines). Base map credits are as follows: State of Colorado outline from Environmental Systems Research Institute, Inc. Redlands, California, USA, Series issue: 2000; river basin boundaries from U.S. Geological Survey, Reston, Virginia, 1994: stream and rivers from U. S. Geological Survey in cooperation with the U.S. Environmental Protection Agency, Reston Virginia, 1998. Platte and upper Colorado River basins. Behnke (1992) proposed a similar scenario for the dispersal of cutthroat trout between the upper Colorado, upper South Platte, and upper Arkansas River basins. Another pos- sible scenario is the potential for a naturally occurring headwater transfer from the North Platte River Basin via the Sweetwa- ter River into the upper Colorado via the tributaries of the Green River (R.J. Behnke personal communication, 2004). No trout species are native to the North Platte River Basin (Behnke 1992) indicating that if a headwater transfer took place between the North Platte and Green River, conditions were not conducive for the survival of trout (e.g., unsuitable water temperatures or tur- bidity). Paracapnia angulata is found in a variety of stream types (Stark and Baumann 2004), which could have facilitated the dis- persal of this species into the upper Colo- rado Basin via the North Platte Basin even though conditions were not appropriate for trout. The low lying area between the Sweetwater River and the upper Green Riv- er also may have provided a flight corridor for dispersing adults even if a water con- nection did not occur during a natural head- water transfer. Stark and Baumann (2004) provide records of P. angulata from the North Platte River supporting the headwater transfer theory. Material examined.—Colorado, Jackson Co., Roaring Fork, CR 5 Bridge, 2,461 m, 31-IH-1991, B. C. Kondratieff & R. R. Dur- fee, 3 6, 15 2, 4 nymphs, CSUC; North Platte River, North Gate Canyon, 21-IV- 1993WR. Re Durteey 16, 392 VESUC] Lar VOLUME 108, NUMBER 2 imer Co., Dale Creek, Hwy 287 bridge, 21059 sniw26-lV-1992R Rj Durfee; 1,2, CSUC; 1-IV-1994, H. A. Rhodes, | nymph, CSUC; Cache la Poudre River, Lyons Park Overland Trail Rd. NW of Fort Collins, 1,544 m, 20-II-2001, anonymous, 1 <6, CSUC; Routt Co., Yampa River, upstream from the confluence with Service Creek just below Stage Coach Reservoir, 2,138 m, 31- I-1995, TIT. Schneck, 2 nymphs, CSUC; Grizzly Creek, Hwy 14, 16.1 km W of Wal- den, 2455 m 7-III-1990, B. C. Kondratieff & M. Harris, 1 d, 2 nymphs, CSUC. Stark and Baumann (2004) and Baumann et al. (1977) provide additional P. angulata rec- ords for Colorado. Taeniopteryx burksi Ricker and Ross 1968 Taeniopteryx burksi is a widespread spe- cies ranging over much of eastern and cen- tral North America (Stewart 2000). Previ- ously, Kondratieff and Ward (1987) reported Colorado localities for this species from the North and South forks of the Republican River and Chief Creek. These localities oc- cur in the Great Plains of eastern Colorado. Kondratieff and Baumann (1988) indicated that the presence of 7. burksi in Colorado is a result of the western dispersal of this spe- cies from the east associated with continen- tal glaciation events (Stewart et al. 1974). The Republican River is tributary to the Missouri River via the Kansas River, which probably provided the conduit for westward dispersal into Colorado. During this survey, T. burksi was collected and reared from the South Platte River in North Platte, Nebraska. It is likely 7. burksi will be collected from the northeastern portion of the South Platte River Basin in Colorado (Fig. 1). Material examined.—Colorado, Yuma Co., Chief Creek, CR CC North, 1,111 m, 22-I-1994, B. C. Kondratieff & R. R. Dur- feew2 356,27 2, 18 nymphs, CSUC; North Fork Republican River, River by Rail Road, 1,026 m, 20-II-1994, H. A. Rhodes & B. C. Kondratieff, 6.0.5 &.°8 nymphs;.€SUC. Kondratieff and Ward (1987) and Kondra- 337 tieff and Baumann (1988) provide addition- al T. burksi records for Colorado. Taeniopteryx parvula Banks 1918 Taeniopteryx parvula is a species typical of larger streams in eastern North America (Stewart 2000). Canton et al. (1981) first re- ported unidentified species of Taeniopteryx in Colorado from the North Platte Basin. Kondratieff and Baumann (1988) later con- cluded these populations were T. parvula. The presence of 7. parvula in north central Colorado is probably a result of the western dispersal from the east along the Missouri and Platte River connections. The Colorado records are from the North Platte River Ba- sin supporting this route of dispersal. Kon- dratieff and Baumann (1988) reported that this species might be more common than re- cords indicate because emergence occurs in some populations when streams may be un- der ice. Additional sampling is necessary along the North Platte River corridor to de- termine the distribution of this species in Colorado, Wyoming, and Nebraska. Material examined.—Colorado, Jackson Co., North Platte River, North Gate Can- yon, 2,383 m, 21-IV-1993, R. R. Durfee, 24 Om4n?s CSUC Roaringelork.sCR 5 Bridge, 2,461 m, 31-III-1991, B. C. Kon- dratieff & R. R. Durfee, 3 nymphs, CSUC; Grizzly Creek, Hwy 14 15.8 km W of Wal- den, 2,455 m, 7-IV-1990, B. C. Kondratieff & M. Harris, 2 2, CSUC; same creek, ex- cept 1 6-X1-1 986,45. C Kondratieff, 5 nymphs, CSUC. Kondratieff and Baumann (1986) provide additional 7. parvula rec- ords for Colorado. Acroneuria abnormis (Newman 1938) Acroneuria abnormis has one of the wid- est geographical distributions of any stone- fly in North America (Stark and Baumann 2005). In the Rocky Mountain states this species has been reported from Colorado, New Mexico, Utah, and Wyoming (Stark and Baumann 2005). In Colorado, this spe- cies is found only in the Colorado River and larger tributaries west of the Continen- 338 tal Divide, and reflects a Missouri River dispersal pattern (Baumann et al. 1977, Baumann and Jacobi 1984). The dispersal of A. abnormis to the western slope of Col- orado likely follows the same pattern as P. angulata through a headwater transfer be- tween the upper Sweetwater River Basin and the upper Green River Basin in Wyo- ming. Acroneuria abnormis is found in the North Platte River and upper Green River in Wyoming (unpublished data on file at CSUC) so it is possible that a natural head- water transfer between these two basins may explain the dispersal of stoneflies with eastern affinities across the Continental Di- vide into the upper Colorado River system. Even if a natural headwater transfer did not take place, the low lying topographic sep- aration may have facilitated flight dispersal across the Continental Divide. Material examined.—Colorado, Mesa Co., Colorado River, Corn Lake State Wildlife Area, 1,433 m, 2-IX-1990, R.R. Durfee, 1 nymph, CSUC; at Broadway in Grand Junc- tion, 1,385 m, 23-XI-1990, M. Trammell, | nymph, CSUC; at Palisade, 1,437 m, 6-IV- 1991, B. C. Kondratieff, 1 nymph, CSUC; Hwy 141 at Colorado River State Park, 1,414 m, 20-IX-1998, H. Freeman, | nymph, CSUC; same as above except, 2-VI-1988, S. Jones, 1 2, CSUC; Moffat Co., Green River, Echo Park Dinosaur National Monument, 1,640 m, 19-VIH-1993, B.C. Kondratieff & R.R. Durfee, 3 nymphs, CSUC. Baumann et al. (1977) provides additional records of this species from Colorado including a record from Garfield County. Perlesta decipiens (Walsh 1862) Herbert H. Ross and J. A. Ross collected the earliest record of P. decipiens known from Colorado in 1938 at Longmont (INHS) probably from the Saint Vrain Riv- er. Stark (1989) reported this species to be common in the Republican River Drainage in eastern Colorado. This species appears to be quite common in the eastern portion of the South Platte River Basin and has been recently collected near the Plains/Mountain PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Interface along the Front Range of Colo- rado from Ralston Creek in Arvada, CO, and from lights in Fort Collins, CO. Addi- tionally, a population occurs in the North Fork Cache la Poudre River where it drains the foothills of the northern portion of the east slope of the Rocky Mountains in Col- orado (Zuellig et al. 2002). In this popula- tion, P. decipiens co-occurs with typical mountain taxa (Zuellig et al. 2002, Kondra- tieff and Baumann 2002). This widespread midwestern species probably moved into Colorado from the east via the South Platte River and its downstream tributaries with the Missouri River. This species also is known from New Mexico (Jacobi et al. 2005). Kondratieff and Baumann (2002) reported on a disjunct population in the Conejos River where it also co-occurs with typical mountain fauna. The occurrence of P. decipiens in the Co- nejos River may be explained by its con- nection with the Rio Grande Basin as we have records from the Rio Grande River downstream of Colorado (unpublished data on file at CSUC). It is likely that P. deci- piens will be collected in other river basins of the eastern plains of Colorado such as the Arkansas (Fig. 1). Material examined.—Colorado, Boulder Co., Probably at the Saint Vrain River, Longmont, 1,503 m, 27-VII-1938, H. H. Ross & J. A. Ross, I 2, INHS; Jefferson Co., Ralston Creek above Simms at Ar- vada, 25-VI-2003, 3 nymphs, USGS NWQL, Morgan Co., South Platte River, Cottonwood State Wildlife Area N of Brush, 5-VI-2000, B. C. Kondratieff, 25 3, 10 2, CSUC; Larimer Co., Shields and Horsetooth in parking lot, 1,548 m, 28- VIII-2003, J. Owens, 1 3, CSUC; Irriga- tion Canal, Wellington Game Lands 3.2 km E of Wellington, 1,586 m, 19-X-1986, R. Keith, 3 nymphs, CSUC; North Fork Cache la Poudre River, Phantom Canyon Pre- serve, 1,627 m, 26-VII-1997, B. C. Kon- dratieff, 1 nymph, CSUC; Black light trap, Cranshaw Home, Fort Collins, 1,524 m, 2- VIII-1987, W. Cranshaw, 1 6, 1 2, CSUC; VOLUME 108, NUMBER 2 Conejos Co., Conejos River, CR 28 at state Stream Gage, 2,286 sm, 30-VI-1996, B. C. Kondratieff, 1 ¢,,3 2; CSUC; Weld Co., 17.7 km E SE of Roggen, 1,480 m, 26-VII- [995mPrAe Opler 2°, 12; CSUC; Yuma Cor Cire Creek "ER CC North) 15117 m:; 28-VI-1986, B. C. Kondratieff, 1 ¢, 12 9, 5 nymphs, CSUC. Isoperla bilineata (Say) 1823 The validity of /soperla bilineata in Col- orado is uncertain, but many scientists agree that it is possible /. bilineata once in- habited Colorado (Baumann et al. 1977, Szczytko and Stewart 1978, Ruse and Herrmann 2000, Kondratieff and Baumann 2002). Needham and Claassen (1925) first reported /. bilineata from Colorado but did not give locality information. Needham and Claassen deposited much of their material in the entomological collection at Cornell University, Ithaca, New York. The senior author contacted E. Richard Hoebeke, Cor- nell University, but the specimens of J. bil- ineata reported by Needham and Claassen (1925) were not in the known holdings of the Collection. If this record is valid, it is possible that /. bilineata entered Colorado through the Arkansas River or the South Platte River drainage system. It is likely that /. bilineata occurred in the South Platte and Arkansas basins before the occurrence of urban development and extensive water management. Rhodes and Kondratieff (1996) did not report /. bilineata from the eastern tributaries of the South Platte River in Nebraska. However, adults of this species are known from the Cedar River, North Loup River, Mormon Canal, and the Mis- sour River in north central Nebraska (Kon- dratieff and Zuellig unpublished data on file at CSUC). A recent survey of the Arkansas River Basin in Colorado did not find this species (Ruse and Herrmann 2000). Isoperla marlynia (Needham and Claassen) 1925 Kondratieff and Baumann (2002) report- ed J. marlynia as a new state record for Col- 339 orado from the Republican River Basin. This species also is known from the eastern portion of the South Platte River in Colo- rado. Isoperla marlynia likely colonized eastern Colorado from the east during the post glaciation period via the Missouri Riv- er and its tributaries. This species also is known from the neighboring states of Kan- sas and Nebraska (Stark 2001) where it can be quite common. Material examined.—Colorado, Sedgew- ick Co., South Platte River, Hwy 385 at Ju- lesburg, 1,070 m, 24-I-2004, reared 20-III- 2004, R. E. Zuellig & R. W. Hood, | fe- male, CSUC. Kondratieff and Baumann (2002) provided additional records of J. marlyina from Yuma County, Colorado. Pteronarcys pictetii Hagen 1873 The genus Preronarcys Newman current- ly includes eight species that occur in North America (Nelson and Hanson 1971, Stark and Szczytko 1982). Of these eight species, P. californica Newport and P. princeps Banks are western North American species (Stark 2001); whereas, P. biloba Newman, P. comstocki Smith, P. pictetii Hagen, P. proteus Newman, and P. scotti Ricker are associated with eastern or central midwest- ern North American streams (Nelson 2000). The transcontinental species, P. dorsata (Say) can be a common species in larger rivers of the East but is also recorded from the Northwest Territories, Alaska, and Brit- ish Columbia (Nelson 2000). Our recent survey of the eastern plains portion of the South Platte River included P. pictetii for the first time in Colorado. Pter- onarcys pictetii has been previously reported from the neighboring states of Kansas (Stew- art and Huggins 1977) and Nebraska (Rhodes and Kondratieff 1996). The occurrence of this species in northeastern Colorado reflects a Missouri River dispersal pattern. Material examined.—New State Record, Colorado, Sedgwick Co., South Platte Riv- er, Hwy 385, at Julesburg, 1070 m, Col- lected 24-I-2004, reared 9-III-2004, R. E. Zuellig & "ROW. Hood, 246) 2°2-CSUG) ACKNOWLEDGMENTS Special thanks to Travis Schmidt, De- partment of Fishery and Wildlife Biology, Colorado State University, for assisting in making the map of Colorado used in Fig. 1. Richard W. Baumann provided access to records held in the Monte L. Bean Museum, Brigham Young University, Provo Utah. Edward DeWalt provided access to material in the Illinois Natural History Survey Col- lection. Richard W. Baumann, Kenneth W. Stewart, and Scott Grotheer improved ear- lier versions of this manuscript. LITERATURE CITED Behnke, R. J. 1992. Native trout of western North America. American Fisheries Society Monograph 6. Bethesda, Maryland. 275 pp. Baumann, R. W. and G. Z. Jacobi. 1984. Two new species of stoneflies (Plecoptera) from New Mex- ico. Proceedings of the Entomological Society of Washington 86: 147-154. Baumann, R. W., A. R. Gaufin, and R. E Surdick. 1977. The Stoneflies (Plecoptera) of the Rocky Mountains. Memoirs of the American Entomolog- ical Society Number 31, 208 pp. Canton, S., J, Chadwick, and L. Britton. 1981. A new distributional record for Taeniopteryx (Plecoptera: Taeniopterygidae) from Colorado. Entomological News 92: 155. Huntsman, B. O., R. W. Baumann and B. C. Kondra- tieff. 1999. Stoneflies (Plecoptera) of the Black Hills of South Dakota and Wyoming, USA: dis- tribution and zoogeographic affinities. Great Basin Naturalist 59: 1-17. Jacobi, G. Z., S. J. Cary, and R. W. Baumann. 2005. An updated list of the stoneflies (Plecoptera) of New Mexico, U.S.A. Entomological News 116: 29-34. Kondratieff, B. C. and R. W. Baumann. 1989 (1988). Taeniopteryx of western North America (Plecop- tera, Taeniopterygidae). Pan-Pacific Entomologist 64: 381-390. . 2002. A review of the stoneflies of Colorado with description of a new species of Capnia (Ple- coptera: Capniidae). Transactions of the American Entomological Society 128: 385—401. Kondratieff, B. C. and J. V. Ward. 1987. Taeniopteryx burksi (Plecoptera, Taeniopterygidae) in Colora- do, with notes on aquatic insects of plains streams. Entomological News 98: 13-16. Needham J.G. and Claassen PW. 1925. A monograph of the Plecoptera or stoneflies of America north PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of Mexico. Entomological Society of North Amer- ica. Thomas Say Foundation 2: 397 pp. Nelson, C. H. 2000. Pteronarcyidae (The Salmonflies). In Stark, B. P. and B. J. Armitage, eds. Stoneflies (Plecoptera) of Eastern North America, Volume I, Pteronarcyidae, Peltoperlidae, and Taeniopterygi- dae. Bulletin of the Ohio Biological Survey. New Series 14, 99 pp. Nelson, C. H. and J. K Hanson. 1971. Contribution to the anatomy and phylogeny of the family Pter- onarcyidae. Transactions of the American Ento- mological Society 97: 123-200. Rhodes, H. A. and B. C. Kondratieff. 1996. Annotated list of the stoneflies (Plecoptera) of western Ne- braska, U.S.A. Journal of the Kansas Entomolog- ical Society 69: 191-198. Ruse, L. P. and S. J. Herrmann. 2000. Plecoptera and Trichoptera species distribution related to environ- mental characteristics of the metal-polluted Ar- kansas River, Colorado. Western North American Naturalist 60: 57—65. Stark, B. P. 1989. Perlesta placida (Hagen), an eastern Nearctic species complex (Plecoptera, Perlidae). Entomologica Scandinavica 20: 263-286. Stark, B. P. and R. W. Baumann. 2004. The winter stonefly genus Paracapnia (Plecoptera: Capni- idae). 2004. Monographs of the Western North American Naturalist 2: 96-108. Stark, B. P. and R. W. Baumann. 2005. North America Stonefly List. http://mlbean.byu.edu/plecoptera/ list.asp Version March 30, 2005. Stark, B. P. and S. W. Szczytko. 1982. Egg morphology and phylogeny in Pteronarcyidae (Plecoptera). Annals of the Entomological Society of America Ie SIQ=sy18)- Stewart, K. W. 2000. Taeniopterygidae (The Willow- flies). Jn Stark, B. P and B. J. Armitage, eds. Stoneflies (Plecoptera) of Eastern North America, Volume I, Pteronarcyidae, Peltoperlidae, and Tae- niopterygidae. Bulletin of Ohio Biological Sur- vey. New Series,14: 99 pp. Stewart, K. W. and D. G. Huggins. 1977. Kansas Ple- coptera (Stoneflies). Technical Publications of the State Biological Survey of Kansas 4: 31—40. Stewart, K. W., R. W. Baumann, and B. P. Stark. 1974. The distribution and past dispersal of southwest- ern United States Plecoptera. Transactions of the American Entomological Society 99: 507-546. Szczytko, S. W. and K. W. Stewart. 1978. Isoperla 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. Zuellig, R. E., B. C. Kondratieff, and H. A. Rhodes. 2002. Benthos recovery after an episodic sediment release into a Colorado Rocky Mountain river. Western North American Naturalist 62: 59-72. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 341-346 DESCRIPTION OF THE IMMATURE STAGES OF PYRODERCES BADIA (HODGES) (LEPIDOPTERA: COSMOPTERIGIDAE), WITH A NEW HOST RECORD FROM LOUISIANA DAVID ADAMSKI, JOHN W. BROWN, AND WILLIAM H. WHITE (DA, JWB) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, Smithsonian Institution, PO. Box 37012, MRC 168, Washington, D.C., 20013-7012, U.S.A. (e-mail: dadamski @sei.barc.usda.gov; jbrown @sel.barc.usda.gov); (WHW) Southern Regional Re- search Center, Sugarcane Research Unit, 5883 USDA Road, Houma, LA, 70360, U.S.A. (e-mail: wwhite @srrc.ars.usda.gov) Abstract.—The last larval instar and pupa of Pyroderces badia (Hodges) are described and illustrated based on specimens collected from seed heads of sorghum (Sorghum bi- color (L.) Moench) (Poaceae) in southern Louisiana. The species is a well-known scav- enger on a wide array of plant material. Key Words: The cosmopterigid genus Pyroderces Herrich-Schaffer includes more than 80 species in the tropical, subtropical, and warm temperate parts of the world, with greatest species richness in Australia and the Old World tropics (Hodges 1978). Ob- servations on the early stages suggest that larvae are most likely scavengers on a wide array of plant material. The larvae of Py- roderces trequently are intercepted at U.S. ports-of-entry, associated most often with pineapple (Ananas comosus (L.) Mert.) (Bromeliaceae), and less often with corn (Zea mays L.) (Poaceae), bananas (Musa spp.) (Musaceae), and guava (Psidium gua- Java L.) (Myrtaceae). The U.S. Department of Agriculture, Systematic Entomology Laboratory database includes nearly 150 records of interceptions over the last 15 years, primarily from Central America and the Caribbean (Robert W. Carlson, personal communication). Pyroderces badia (Hodges, 1962) (Fig. 1) is a widespread species of the eastern Cosmopterigidae, Gelechioidea, scavenger, Sorghum seaboard, ranging from Washington, D.C., south to Florida (and possibly the Carib- bean), and west to Louisiana (Hodges 1962, 1978). According to Hodges (1978), the species also is represented by highly dis- junct, probably adventitious (invasive) pop- ulations in southern California and Hawaii. Like most species of Pyroderces, the larvae of P. badia are scavengers, having been reared from seed pods of coffee senna (Cas- sia occidentalis L.) (Fabaceae); peach (Pru- nus persica (L.) Bastch) (Rosaceae) and lo- quat (Eriobotrya japonica (Thunb.) Lindl.) (Rosaceae) mummies; fruits of lime (Citrus aurantiifolia (L.) Swingle) (Rutaceae), grapefruit (Citrus X paradisi Macfad.) (Ru- taceae), and bananas; cabbage (Brassica oleracea L.) (Brassicaceae); coconut blos- soms (Cocos nucifera L.) (Arecaceae); elm leaves (Ulmus spp.) (Ulmaceae); and cones of pine trees (1.e., Pinus elliottii Engelm., P. palustris Mill., P. pinaster Aiton) (Pin- aceae) (Hodges 1962, 1978). Although fre- quently observed, the early stages previ- 342 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ously have not been described or illustrated. While conducting field surveys in Louisi- ana in 2002, we discovered larvae of P. badia feeding in the flower heads of live sorghum, Sorghum bicolor (L.) Moench (Poaceae); we reared adults and preserved examples of the early stages. We take this opportunity to describe and illustrate the last instar larva and pupa of P. badia and compare the larva with that of P. rileyi (Walsingham). MATERIALS AND METHODS Seed heads of sorghum (Sorghum bicolor (L.) Moench) (Poaceae) infested with lar- vae of Pyroderces badia were collected in Avoyelles Parish (29.4525°N, 90.2777°W), Louisiana, on 2 August 2002, and brought into the laboratory for examination. Fifteen larvae were removed, boiled in H,0, and preserved in 70% EtOH. The remaining lar- vae were allowed to complete larval devel- opment and pupate. Pupae were collected periodically (n = 21) and preserved in 70% EtOH. The remaining pupae were allowed to complete development. Newly emerged adults were frozen and later spread and double-mounted. For study using electron microscopy, lar- vae and pupae were cleaned in 10% EtOH with a camel’s-hair brush and dehydrated in increasing concentrations of EtOH to ab- solute EtOH. After dehydration, specimens were critical-point dried using a Tousimis critical point dryer, mounted on stubs, and coated with gold-palladium (40—60%) us- ing a Cressington sputter coater. The ultra- structure of the larvae and pupae was stud- ied with an Amray scanning electron mi- croscope. Gross morphological observations and measurements of the larvae and pupae were made using a dissecting microscope (re- flected light) with a calibrated micrometer. Maps of the larval chaetotaxy were initially drawn using a WILD dissecting microscope with a camera lucida attachment. Terminol- ogy for chaetotaxy follows Stehr (1987). All vouchers (larvae, pupae, and adult spec- imens) are deposited in the National Mu- seum of Natural History (USNM), Smith- sonian Institution, Washington, D.C. Larval specimens of Pyroderces rileyi were ex- amined from the larval alcohol collection at USNM. RESULTS Pyroderces badia (Hodges) (Figs. 1-13) Last Instar larva (Figs. 2—11).—Length 6.0-8.5 mm, mean = 7.4 mm (n = 15). Body pale brownish gray with a trace of pink (live specimens pink); integument ap- parently smooth under light microscopy, granular under SEM at magnifications above 200; epicranium, prothoracic shield, and L-group pinaculum slightly darker than anal plate; pinacula slightly darker than body; thoracic legs concolorous with body; spiracles on Tl and A8 about equal in size, twice diameter of spiracles on A1-A7. Head (Figs. 2, 3): Hypognathous, adfrontal sclerites narrow, meeting dorsally between lateral hemispheres of epicranium; coronal suture absent; AF-group setae on apical half of adfrontal sclerites, AF2 near apex, about equidistant to AFI as AFI is to Fl; AF1 and AF2 slightly longer than F1; P1 dorsolaterad to AF1, about 3—4 times as long as P2; Cl beneath and slightly anterad to Fl, C2 above base of mandible, slightly longer than Cl; mandible (Fig. 10) with 5 dentiform processes, and bearing 2 sube- qual setae; maxilla and antenna with sen- sillar types and arrangement as_ figured (Figs. 4—5); 6 stemmata in a semicircular array, with stemma III and IV proximate; setae associated with stemmata in genal area illustrated in Fig. 3. Thorax (Fig. 8): Prothoracic shield with XD1 and D2 along anterior margin, SD1 along anterolateral margin; XD2 closer to SD1 than to XD1; XD1 about equal in length to XD2; SD1 about twice as long as XD1 and XD2; SD2 equally distant from XD2 and SD1, form- ing a triangular array within anterolateral angle; D1 posteriorad to XD1, both setae VOLUME 108, NUMBER 2 / Fig. 1. Adult of Pyroderces badia. parallel to median longitudinal axis, D1 and D2 in straight line perpendicular to axis; D2 and SD1 about equal in length; DI and SD2 about equal in length and about % as long as XD1 and XD2. L-group pinaculum sub- rectangular and anteriad to spiracle; L2 be- neath L1, both anteriad to L3; L2 about twice as long as LI] and at least 3 times as long as L3. SV-group bisetose, SV2 about ¥% as long as SVI. Distance between coxae of Tl about % distance of that between cox- ae on T2-T3; V-group setae on T1-T2 2-3 times farther apart than on prothorax. T1- T2 (Fig. 8) with DI and D2 on same pin- aculum; D2 about 3 times as long as D1; SD1 and SD2 on same pinaculum, slightly anteriad to D-group pinaculum, SD1 about 2 times as long as SD2 and about equal in length to D2; LI and L2 on same pinacu- lum, L1 about equal in length to D2 and SD1, and about % times as long as L2 and about 3 times as long as L3; SV-group un- isetose, SV1 posterior to L3. Abdomen (Figs. 9, 11): Al-A2 (Fig. 9) with D2 about 2.5 times as long as D1; SD1 above spira- cle, about equal in length to D2; LI and L2 343 on same pinaculum, slightly anteriad of spi- racle; L2 about % as long as L1; L3 in ver- tical line with D2, about *%4 as long as SD1 and D2; SV-group trisetose, SV2 about 3 times as long as SV1 and SV3; V-group setae slightly farther apart than V-group se- tae on A3-A6. A3-A6 (Fig. 11) with SD1 dorsoanteriad to spiracle, in approximate straight line with D1 and pinaculum bearing L1-L2; SV-group trisetose, SV1 about 2 times as long as of SV3 and at least 3 times as long as SV2; proleg with uniserial cro- chets, long mesally, gradually shortened along lateral margin. A7 with S V-group bis- etose, SV1 about 2 times as long as SV2. A8 with D2-pinaculum large, but not as large as D2-pinaculum on A9; SDI anter- oventrad to spiracle; SV-group unisetose, SV1 slightly shorter than L3. A9 with D2 pinaculum the largest of abdomen; D2 in line with SD1, with D1 slightly anteriad; D2 about equal in length to SD1, both setae about 5 times as long as D1; L1-L2 on same pinaculum, L1 about 3 times as long as L2; L2 slightly longer than D1 and L3; D1 about equal in length to L3. AlO with anal 344 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-7. plate with D2, SD1, and SD2 along outer margin, SD1 closer to D2 than to SD2; D1 slightly anteriad to SD2; SD2 about equal in length to SD1, both about twice as long as D2; D2 slightly longer than D1; prolegs (Fig. 6) bearing crochets in curved line, with large crochets medially, gradually de- creasing in size mesially and laterally. Pupa (Figs. 12, 13).—Length 3.5—4.5 mm, mean = 4.0 mm (n = 21). Body smooth; spiracles protuberant; terminal six Scanning electron micrographs of the larva of Pyroderces badia. 2, Head, frontal view (scale = 100 j). 3, Head, ventrolateral view (scale = 100 2). 4, Maxillary palpus (scale = 10 wp). 5, Antenna (scale = 10 w). 6, Abdominal proleg (scale = 100 1). 7, Anal plate (scale = 100 p). abdominal segments articulate as a unit; ventral surface of posteroapical part of ab- domen with 5 pairs of setae with hooked apices, dorsal surface of posteroapical part of abdomen with 4 pairs of setae with hooked apices, tergum-9 with 2 setal pairs with hooked apices; tergum-8 with 2 setal pairs with hooked apices; and tergum-7 with | setal pair with hooked apices on pos- terior half; other abdominal terga with setae without hooked apices. VOLUME 108, NUMBER 2 Figs. 8-11. 3. 10, Manible. 11, Abdominal segments 6-10. DISCUSSION In his catalog of the Cosmopterigidae, Si- nev (2002) recently transferred both P. bad- ia and P. rileyi to Anatracynthis. This ac- tion, however, was not explained and has received little support. Hence, we have tak- en a conservative approach and continue to treat these two North American species as members of Pyroderces until more compel- ling evidence is presented to the contrary. Stehr (1987) figured the head, thorax, one of the proleg-bearing abdominal seg- ments, and the crochet arrangement of Py- roderces rileyi (original illustrations from Peterson 1948), a sympatric congener of P. badia. Although generally adequate, a few inaccuracies in the drawing are noted here, based on the examination of specimens in the USNM collection. The pinacula are not 345 11 Chaetotaxy of larva of Pyroderces badia. 8, Thoracic segments (1—3). 9, Abdominal segments 2— shown; the setae are much too short; the placement of L3 and SV1 on TI is too an- teriad; and the crochets should be uniserial, long mesally, gradually shortened along the lateral margin. Although several larvae of Pyroderces rileyi were examined, we could find no diagnostic characters to reliably dif- ferentiate them from the larvae of P. badia. ACKNOWLEDGMENTS We thank James EF DiLoreto, Office of Imaging and Photographic Services, Smith- sonian Institution, Washington, D.C., for the digital processing and preparation of the plates. Robert W. Carlson, USDA System- atic Entomology Laboratory (SEL), provid- ed statistics on interceptions of Pyroderces from the SEL database. We thank the fol- lowing for reviewing the manuscript: Lauri 346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON = iz Figs. 12-13. Kaila, Finnish Museum of Natural History, University of Helsinki, Finland, and Mi- chael Gates and Thomas J. Henry, SEL, USDA, National Museum of Natural His- tory, Washington, DC. LITERATURE CITED Hodges, R. W. 1962. A revision of the Cosmopterigi- dae of America north of Mexico, with a definition of the Momphidae and Walshiidae (Lepidoptera: Gelechioidea). Entomologica Americana 42: 1— 166. . 1978. Fascicle 6.1 Gelechioidea, Cosmopter- 13 Pupa of Pyroderces badia. 12, Ventral view. 13, Dorsal view. Scale = 1.0 mm. igidae. In The Moths of America North of Mexi- co. E. W. Classey Limited and The Wedge Ento- mological Research Foundation, London. 166 pp. Peterson, A. 1948. Larvae of Insects. An Introduction to Nearctic species. Part 1. Lepidoptera and Plant Infesting Hymenoptera. Published by the author, Ann Arbor, Michigan, 315 pp. Sinev, S. Y. 2002. World catalogue of cosmopterigid moths (Lepidoptera: Cosmopterigidae). Russian Academy of Sciences, Proceedings of the Zoolog- ical Institute, St. Petersburg 293: 1-183. Stehr, E 1987. Cosmopterigidae (Gelechioidea), pp. 391-392. In Stehr, E, ed. Immature Insects, Vol- ume 1. Kendall/Hunt Publishing Company, Du- buque, Iowa, 754 pp. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 347-365 A REVISION OF THE GENUS MACROSIPHONIELLA DEL GUERCIO (HEMIPTERA: APHIDIDAE) FROM THE KOREAN PENINSULA, PART II: SUBGENUS MACROSIPHONIELLA (SENSU STRICTO) JAROSLAV HOLMAN, SEUNGHWAN LEE, AND JAN HAVELKA (JHolm JHay) Institute of Entomology, Academy of Science, BraniSovské 31, Ceské Budéjovice, Czech Republic (e-mails: holman@entu.cas.cz; jhav@entu.cas.cz); (SL) Di- vision of Entomology, School of Agricultural Biotechnology, Seoul National University, San 51-1 Shillim-dong, Gwanak-gu, Seoul, 151-742, Korea (e-mail: seung@snu.ac.kr). Corresponding author: Seunghwan Lee Abstract.—Thirteen species of the subgenus Macrosiphoniella del Guercio sensu stricto (Hemiptera: Sternorrhyncha: Aphididae) are recorded from the Korean Peninsula, includ- ing Macrosiphoniella (M.) capillaricola, n. sp. (apt. & al.) and Macrosiphoniella (M.) pseudotanacetaria, n. sp. (apt. & al.). The former is from South and North Korea, found on Artemisia capillaris Thunb. and A. japonica Thunb. The latter is from North Korea, found on an unidentified Asteraceae: Anthemideae. Macrosiphoniella (M.) kikungshana ssp. sylvaticae Szelegiewicz is synonymized with its respective nominate taxon (new synonymy). A key to apterous viviparous females of the Korean species in the subgenera Macrosiphoniella and Asterobium (in part: M. (A.) yomenae (Shinji)) is given, and a list of host plants for all Macrosiphoniella species of the Korean Peninsula is also presented. In addition, descriptions and measurements of the new species, and microscopic photos of all Korean species treated are given. Key Words: Hemiptera, Aphididae, Macrosiphoniella, Korea, list of host plants, Korean Peninsula As the second part of “The Genus Ma- crosiphoniella del Guercio (Hemiptera: Aphididae) from the Korean Peninsula,” this paper includes 13 species of the nom- inate subgenus Macrosiphoniella del Guer- cio. All species are listed in alphabetical or- der, disregarding phylogenetic relation- ships. Five subgenera of the genus Macro- siphoniella (Asterobium, Chosoniella, Papillomyzus, Phalangomyzus, and Sinosi- Phoniella) were treated in the first part of this study (Holman et al. 2006). MATERIALS AND METHODS Most specimens and all holotypes used in this study are deposited in the College of Agriculture and Life Sciences, Seoul Na- tional University (CALS SNU, Korea) and the National Institute of Agricultural Sci- ences and Technology (NIAST, Suwon, Ko- rea), and some paratypes and other speci- mens are deposited in the Institute of En- tomology, Czech Academy of Sciences (IE CAS, Ceské Budéjovice, Czech Republic). A list summarizing the host plants of all the Macrosiphoniella spp. of the Korean Pen- insula is given in Table 2. Abbreviations used are as follows: al.— alate viviparous female, alata; Ant.—anten- nae; Ant.I, II, VIb—antennal segment I, III, base of VI, respectively; apt.—apterous viviparous female, aptera; BDAnt.III—bas- 348 al diameter of antennal segment III; BL— length of body; C—cauda; HFEM—hind femur; HTIB—hind tibia; HT2—second segment of hind tarsus; HW—width of head across the compound eyes; PT—pro- cessus. terminalis; SIPH—siphunculi; URS—ultimate rostral sement (segment IV + V). Provincial names of North and South Korea for the collection data: PN—Pyoun- gannando; RG—Ryanggangdo; HB— Hwanghaebukdo; HN—Hwanghaenamdo; GG—Gyeonggido; GW—Gwangwondo; CB—Chungcheongbukdo; CN—Chungch- eongnamdo; GN—Gyeongsangnamdo; JB—Jeollabukdo; JN—Jeollanamdo. Names of plants with an asterisk (*) in- dicate new host records for the respective species of aphid. KEY TO SPECIES OF THE SUBGENERA MACROSIPHONIELLA AND ASTEROBIUM (IN PART) OF THE KOREAN PENINSULA Apterous viviparous females 1. Siphunculi pale (at least at base), cauda pale or dusky (e.g., Fig. 1B). Abdomen without SGlErotlzatlonn(Gse=p hom Ay) Seen 2 — Siphunculi and cauda entirely dark brown to black (Fig. 2B). Abdomen usually with some sclerotized, dark areas (e.g., Fig. 3A)...... 3 Head uniformly dusky to brownish, concolo- rous with Ant.I and Ant.II. Frons concave. URS shorter than HT2. Longest hairs on Ant.II. and on anterior abdominal terga 1.0— 1.3 times as long as BDAnt.III. On Artemisia annua L and A. iwayomogi Kitam. (Fig. 1) M. (M.) abrotani chosoni Szelegiewicz — Head and Ant.I dark brown to black, Ant.II pale to smoky. Frons not concave, nearly straight. URS longer than HT2. Longest hairs on Ant.III, up to twice as long as BDAnt.II] and, and longest hairs on anterior abdominal terga 2.5-3.5 times as long as BDAnt.III. On Artemisia princeps Pamp., A. feddei H. Ley. & Vaniot, A. mongolica Fisch. Ex Bess.., and A. gmelini Webb. Ex Stechm (Fig. 13) ae: ee M. (M.) taesongsanensis Szelegiewicz 3. Tibiae uniformly dark, sometimes with mid- dle part paler but never concolorous with body (Fig. 2C) — Tibiae with middle part pale, concolorous with body (Fig. 10C) 4. Primary rhinaria naked or with very short cil- ia and distinctly protruding membrane .... 5 — Primary rhinaria ciliate, usually with cilia N PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bent over the flat or slightly convex mem- brane Dark green in life. Body more than 3 mm long. Siphunculi up to 0.9 times as long as cauda. Secondary rhinaria more than 30, ir- regularly arranged on basal half of Ant.III. URS short, at most 0.9 times as long as HT2. On Tanacetum boreale Fisch. Ex DC. (Fig. 11) M. (M.) pseudotanacetaria, n. sp. Shiny black in life. Body less than 3 mm long. Siphunculi more than 0.9 times as long as cauda. URS long, more than 0.93 times as long as HT2. Secondary rhinaria 1—15 in one row on basal 4—% of Ant.II. — 6 Trochanters and basal %—% of femora pale. URS maximum 1.1 times as long as HT2, 0.75—1.0 times as long as Ant.I, 0.7—1.0 times as long as Ant.VIb. On Artemisia capillaries Thunb. (Fig. 3)... Trochanters dark, femora pale, at most on bas- al 1/4. URS long, narrow, 1.25—1.40 times as long as HT2, about 1.1—1.6 times as long as Ant.I, and 1.0—1.4 times (rarely less than 1.1 times) as long as Ant.VIb. On various Arte- misia spp., including A. capillaries Thunb. (Fig. 5) M. (M.) formosartemisiae Takahashi Hind tibia with dorso-posterior row of short, thorn like hairs, different from other tibial hairs. On Artemisia iwayomogi Kitam (Fig. 7) ape mcaraecr ioe as bet M. (M.) jaroslavi Szelegiewicz Hind tibia without a row of short, thorn like hairs. Dorso-posterior hairs not differing sub- stantially from other tibial hairs Abdominal dorsum with well-developed, dark antesiphuncular sclerites and with large scle- roites at base of dorsal hairs (Fig. 2A) .... 9 Abdominal dorsum membranous with no dis- tinct sclerotization, rarely with faint antesi- phuncular sclerites, and with small scleroites on terga 7 and 8 (Fig. 6A)............ 10 Body widely oval, 1.5—2.6 mm long. Dorsal hairs about 2.5 times as long as BDAnt.III. Accessory hairs on URS much longer than the apical hairs. On various Artemisia and Tanacetum boreale Fisch. Ex DC. (Fig. 2) M. (M.) atra latysiphon Holman and Szelegiewicz Body slender, 2.5—4.0 mm, but rarely less than 2.8 mm long. Dorsal hairs less than 2 times as long as BDAnt.III. Some accessory hairs on URS short, not exceeding apical hairs in length. On Aster, Erigeron and some Artemisia spp. M. (Asterobium) yomenae (Shinji) (see Part. I, Holman et al. 2005) M. (M.) capillaricola, n. sp. . Body yellowish green or pinkish in life, with- out wax powder. Coxae and trochanters pale, femora pale on basal 4%—% and gradually dark- er toward apex. Ant.I (outer side) 0.12—0.15 VOLUME 108, NUMBER 2 mm long, %—%4 of URS. On various Artemisia. (Fig. 6) M. (M.) hokkaidensis Miyazaki — Body green in life, covered with fine wax powder. Legs including coxae black except extreme base of femora. Ant.I (outer side) 0.16—0.19 mm, ¥% of, or equal to URS. On various Artemisa spp., Chrysanthemum spp.. Dendranthema morifolium (Ramat) Tzvelev, Tanacetum boreale Fisch. Ex DC. (Fig. 14) M. (M.) yomogifoliae (Shinji) 11. Primary rhinaria ciliate. Green in life, abdom- inal dorsum without scleroites at base of dor- sal hairs. On various Artemisia spp. (Fig. 10) Bert s +, 5 M. (M.) pseudoartemisiae Shinji — Primary rhinaria naked. Shiny brown to black in life, usually with scleroites at base of some dorsal hairs 12. Anterior abdominal terga each with maximum 15 hairs. Antennal hairs maximum 1.5 times ASMOngease DD Ant Ile ee ee 13 — Anterior abdominal terga each with 24 or more hairs. Antennal hairs 2.5 or more times as long as BDAnt.III. On Artemisia spp. (Fig. Sars os M. (M.) kikungshana Takahashi 13. URS longer than HT2, with 6 accessory hairs. Abdominal dorsum without or with only small scleroites at base of some hairs. Cauda with distinct constriction, bearing 17-21 hairs. Secondary rhinaria 10—25, irregularly distributed on basal % to entire length of Ant... On Tanacetum, Dendranthema, Chry- santhemum and some Artemisia spp. (Fig. 12) M. (M.) sanborni (Gillette) — URS shorter than HT2, with 4 accessory hairs. Abdominal dorsum with rather large scleroites at base of nearly all hairs. Cauda elongate tri- angular, usually without distinct constriction, bearing 13—15 hairs. Secondary rhinaria |—2 on basal third of Ant.III. On Artemisia japonica Thunb. (Fig. 9)... M. (M.) oronensis Szelegiewicz Subgenus Macrosiphoniella del Guercio, sensu stricto Macrosiphoniella del Guercio 1911: 331. Type species: Siphonophora atra Ferrari 1872 by original designation. Pyrethromyzus Boérner 1950: 15. Type spe- cies: Macrosiphum sanborni Gillette 1908 by original designation. Mediosiphum Wojciechowski 1993: 108. Type species: Mediosiphum caucasicum Wojciechowski 1993 by monotypy. 349 Macrosiphoniella (M.) abrotani chosoni Szelegiewicz 1980 (Fig, 1) Macrosiphoniella (M.) abrotani chosoni Szelegiewicz 1980: 427 Specimens examined.—NORTH KO- REA: Taesong-san (Botan. Garden), Pyong- yang-si, PN, 30.v.1988, on Artemisia annua (88Ha2699, 88Ha2700, 88Ha2701, 88Ha2702: apt. & al.); Taesong-san, Pyong- yang-si, PN, 6.vi.1988, on Artemisia annua (88Ha2856, 88Ha2865: apt. & al.). Distribution.—Korea (North), Russia (Far East). Host plants.—Artemisia (annua L., iway- omogi Kitam., montana Fisch. Ex Bess.), Tanacetum boreale Fisch. Ex DC. (Pash- chenko 1998). Macrosiphoniella (M.) atra latysiphon Holman and Szelgiewicz 1978 (ies 2) Macrosiphoniella (M.) atra latysiphon Hol- man and Szelgiewicz 1978: 181 Samples examined.—NORTH KOREA: Chonryong-san, Pyongsong-si, PN, 7.Vi1.1985, on Artemisia gmelini (85Ha405: apt. & al.); Hyesan, RG, 8.vii.1985, on Ar- temisia japonica (85Hal523: apt. & al.); Puryong, Chongjin, HB, 17.vi.1987, on A. iwayomogi (87Hal 837, 87Hal 838, 87Hal839: apt.); Hyeryong, 18.vi.1987, on A. iwayomogi (87Hal870, 87Hal872: apt. & al.); Chongryong-san, Pyongsong-si, PN, 29.vi-6.vil.1987, on A. japonica & A. iway- omogi (87Ha2056, 87Ha2150: apt. & al.); Taesong-san, Pyongyang-si, PN, 2-— 4.v1.1988. A. iwayomogi (88Ha2758, 88Ha2763, 88Ha2765, 88Ha2767, 88Ha2768, 88Ha2769, 88Ha2770, 88Ha2771, 88Ha2887: apt. & al.); Hyesan, RG, 26.vi.1988, on -2 1k, 242219) Geibio- nomics); Ward 1992: 210 (catalog). Anopheles sp. nr. salbaii of Ribeiro et al. 1985: 689-692 (A biology). Anopheles stephensi of Gad 1967: 172—174 (L*, L bionomics); Gad and Kamel 1967: 249-252 (L bionomics, A, L morpholo- gy); Gad and Salit 1972: 581 (A biolo- gy); El-Said and Kenawy 1983: 69, 73 (collection record); Kenawy 1988: 74 (bi- onomics, distribution); Kenawy 1990: 270, 271 (collection record). Anopheles (Cellia) n. sp. of Glick 1992: 129, 130, 140, 150 (distribution, A key). Diagnosis.—Anopheles ainshamsi 1s a member of the Neocellia Series based on the absence of upper proepisternal setae in adults and the presence of one branched long propleural, one branched long meso- pleural, and two branched long metapleural setae (9-P, 9-M, and 9,10-T, respectively) in larvae. Adults of An. ainshamsi are distin- guished from Oriental members of the se- ries, except An. stephensi, in having both spotted legs and hindtarsomere 5 dark- scaled, and from Afrotropical members of the series, except An. dancalicus, An. herv- yi, and An. salbaii, by the combination of spotted legs, dark hindtarsomere 5, and scaling on abdominal segments [-—VIII. The absence of scales on the scutal fossa distinguishes An. ainshamsi from An. Ste- phensi and the three similar Afrotropical species. Anopheles ainshamsi resembles An. dancalicus and differs from the other three species in lacking a pale fringe spot at the apex of the anal vein of the wings. Larvae are distinguished from other mem- 368 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sternal A Lituah— Fig. 1. Pupa and male genitalia of Anopheles (Cellia) ainshansi. A, Pupa, left side of cephalothorax, dorsal to right. B, Pupa, dorsal (left) and ventral (right) aspects of metathorax and abdomen. C, Male genitalia, aspects as indicated. Ae, aedeagus; c, club on dorsal lobe of claspette; Cl, claspette; CT, cephalothorax; Gc, gonocoxite; Gs, gonostylus; InS, internal seta; LAe, leaflets of aedeagus; Pa, paddle; PBS, parabasal setae; Pr, proctiger; I- IX = abdominal segments I-IX; 1-14 = setal numbers for specified areas, e.g., seta 3-I. Scales in mm. VOLUME 108, NUMBER 2 ber of the series by one or more of the fol- lowing characters: seta 2-C not brush-like; 2,3-C aciculate or frayed; 1,2-P well sepa- rated, inserted on small tubercles; 2-P with fewer than 15 branches; 11-P without short barblike branches; 1-III—VII lanceolate branches or blades with weakly developed shoulders and a short filament. Pupae of species belonging to the series are not well known and are difficult to distinguish. Female.—Overtly brown and pale yel- low. Head: Truncate erect scales of vertex pale (white) anteriorly and becoming pro- gressively darker (yellowish to brown) pos- teriorly and posterolaterally; eyes widely spaced, erect scales grade into elongate semierect fusiform scales on interocular space, these scales interspersed with long golden setae, lateral margins of interocular space lined with white decumbent scales that become longer and give rise to long sinuous setae above antennal pedicels. Clypeus bare. Antenna length 0.9—1.2 mm (mean 1.0 mm); pedicel with yellowish to brown integument and usually few incon- spicuous pale scales on dorsal surface; fla- gellomeres 1—3 with elongate pale scales, particularly dense on mesal surfaces. Pro- boscis 1.4—1.7 mm (mean 1.6 mm), slightly longer than forefemur (about 1.1); pre- mentum entirely dark-scaled, scales ap- pressed throughout except for few slightly erect scales at base; labella slightly paler than prementum. Maxillary palpus length 1.4—-1.6 mm (mean 1.5 mm), usually with 3 pale (white) bands—apical (apex of pal- pomere 4 and all of 5), preapical (apex of 3 and base of 4) and proximal (apex of 2)— apical pale band about length of preapical dark band (middle of palpomere 4), palpus with 4-banded appearance when middle of palpomere 5 occasionally dark-scaled; pal- pomere 2 with semi-erect scales giving a slightly bushy appearance to proximal por- tion of palpus; ventral surface of palpus without scales. Thorax: Integument dark brown; scutum with broad median pale pru- inose area confluent with scutellum of sim- ilar appearance; anterior promontory and 369 antedorsocentral areas with white semierect scales that grade into yellowish to golden decumbent fusiform scales on acrostichal and dorsocentral areas, scales on these areas converge at middle of scutum and extend posteriorly between posterior dorsocentral and lateral prescutellar setae to scutellum, with small posterior medial area of pre- scutellar area void of scales, scales at lateral margins of this bare area become white be- fore margin of scutellum; golden to golden- brown setae on acrostichal, dorsocentral, fossal, antealar, supraalar and prescutal ar- eas; narrow line of decumbent pale scales on mesal side of supraalar setae extends to near parascutellar seta, this line of scales separated from scales on posterior dorso- central and prescutellar areas by rather broad pale pruinose area. Scutellum with row of white to golden fusiform scales ad- jacent to posterior row of long golden- brown setae. Mesopostnotum and postpro- notum bare. Antepronotum without scales, with long golden-brown setae. Pleura with golden-brown setae: O—4(1) prespiracular area, 1—3(2) prealar, 2,3(2) upper and 1— 3(2) lower mesokatepisternal and 3-—6(4) upper mesepimeral; upper proepisternal se- tae absent. Wing (see Fig. 62 in Glick 1992): Length 2.7—3.3 mm (mean 2.9 mm); dark scaling brown, stark on costa, subcosta and R-R,, subdued on posterior veins, pale scaling pale yellow, not white; costa with humeral dark spot, dark basal to humeral crossvein; costa, subcosta and R with pre- sector and sector dark spots, presector equally long on 3 veins, sector about half as long on R, sometimes with pale inter- ruption; costa and R, with equally long preapical and apical dark spots; remigium and base of R pale to presector dark spot; dark spots on other veins often faint (some sometimes absent, fully present as follows: Rs dark except at base, R, with dark spot opposite apical dark spot on R,, spur of R,,; dark, postbasal and preapical dark spots on R,,;, M,, M>, M,;,, and CuP, distal areas of M and mcu dark-scaled, 1A with 2 dark spots in distal half (preapical and 370 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Range (mode) of numbers of branches for pupal setae of Anopheles (Cellia) ainshamsi. a eee an See Abdominal Segments | rate No. (Gir I I Il IV Vv VI VII Vil IX Pa i oO — oo ] 1 ] l 1 — = 1 24(3) ~80 5-10(8) 4-8(6) 3-10(4) 14(@) 1,2) 1.20) — 2G) aa De, 12=4((2) 2-810) 4-96) 3-714), 3-6) 3=5'G) 2 3513) ee (3) = 2-5 (4) 3 256) 2c) iA@) I @) 2=7G) etch) 2c Ite) = = = A 9-516) 2264 1276) 2276) s-64@) 543) £2030)" — — Sr 491) y wl ol) e263) 8 (62-814) 3 =6) 214) 2-9 —_— — 6002-5 Gin 1E4G) 3-56) 3286) 3-714) 256) 246) Ses = = It 2G 2intd=614)., 2-616). 3-614). 3-66), 2-416) 32 1-3) = 8s 2 e@)) = a 2-5 (4) 146) 14@2) 1408) 1306) — = = OR l= 3) (2) 2 Ci 1 ] ] 1 1-3 (1) 7-11(8) — —= 10 1-4(2) — D= 513) ls CL) eel 21) ee =Si(2) a) = = 11 1-5 (4) — — 2D) Ne ZB) he 2) 14(2) — = = 14 (2 — —_ | — a = alveolus only. just beyond midlength), apex of vein with- out scales; faint pale fringe spots at apices of M,, M,, M;,, and CuP (total of 4 pale fringe spots). Halter: Pedicel and scabellum pale, capitellum dark-scaled. Legs: Fore- coxa with few inconspicuous dark scales among setae anteriorly at base, midcoxa usually with few inconspicuous pale scales among setae laterally at base, hindcoxa without scales; femora, tibiae and first tar- someres with speckles and blotches of pale yellow scaling; foretarsus with pale bands across joints, mid- and hindtarsi with nar- row pale bands or dorsal pale spots at api- ces of tarsomeres 1—4, sometimes faint or absent on midtarsomeres 2—4, hindtarso- mere 5 usually narrowly pale at base. Ab- domen: Integument dark, with golden se- tae; tergum I and sterna I-VI without scales, terga II-VIII largely covered (except laterally) with pale yellow to golden fusi- form and narrow spatulate (primarily) scales, cerci with similar scales, sternum VIII, and sometimes posterior area of ster- num VII, with scattered pale scales (see Glick 1992: Fig. 64). Male.—Similar to female except for ob- vious sexual differences; other differences include the following. Head: Proboscis slightly longer, 1.6—1.8 mm (mean 1.7 mm), about 1.6 length of forefemur. Maxillary palpus largely pale-scaled, dark scaling on palpomere 1, proximal 0.5 or less of pal- pomere 2 and narrowly across joints be- tween palpomeres 2—3, 3—4 and 4—5. Wing: Length 2.5—-3.0 mm (mean 2.7 mm); gen- erally paler and scaling reduced, dark spots of posterior veins very faint or absent, fringe spots unapparent. Genitalia (Fig. 1C): Gonocoxite with pale yellow scales on lateral surface; with 5 parabasal setae, most sternocaudal seta long and slender, similar to unspecialized setae of gonocoxite; gon- ostylus strongly and evenly curved in distal half, with row of minute setae along ster- nomesal margin and | or 2 longer setae on tergal side near apex; claspette with long apical seta about 1.5 length of club and 3 or 4 shorter subapical setae, club formed of 4 fused setae; aedeagus with 2 or 3 pairs of smooth, slender, attenuated leaflets; proctig- er membranous, with lightly sclerotized long narrow lateral paraprocts. Pupa (Fig. 1A, B).—Character and po- sitions of setae as figured; numbers of branches in Table 1. Cephalothorax: Light- ly tanned, legs darker, scutum and meta- notum with darker blotches. Seta 7-CT VOLUME 108, NUMBER 2 about 1.8 length of 6-CT, usually double, sometimes triple; 8-CT normally single, rarely double; 10-CT usual double or triple (1—4 branches) with branches arising near base; 11-CT usually split distally into 2—5 short branches, sometimes single. Trumpet: Angusticorn, moderately tanned, borne on tubercle, tracheoid absent, pinna without slit; length 0.37—0.46 mm (mean 0.42 mm); meatus fairly long, 0.15—0.22 mm (mean 0.18 mm); pinna slightly longer, 0.19—0.25 mm (mean 0.23 mm). Abdomen: Length 2.22—3.04 mm (mean 2.54 mm); lightly tanned, anterior margins of sterna darker, progressively lighter after sternum IV. Seta Q-II-VIH minute, simple, inserted anterior and usually slightly mesad of seta 2; seta 1- H-IV with multiple thin flexible branches, 1-V usually double (1—4 branches) and lon- ger than following tergum, 1-VI,VII usually single (infrequently double) and longer than following tergum; seta 6-II generally triple (3—5 branches) and nearly twice length of seta 7, 6-III—VII multiple branched, number of branches generally progressively de- crease from 6-III to 6-VII; seta 7-III,1V of- ten inserted within striations of fold line, 7- V—-VII always inserted on fold line, 7-VII inserted at posterior margin of segment, seta 7-III-V short, branched, 7-VI,VII usu- ally single, long, about length of following sternum; setae 8,10,11-II absent, alveolus of 8-II usually present; seta 9-I relatively short, about 0.35 length of 6-I, usually sin- gle, occasionally double; 9-II-IV_ small, peg-like; 9-V—VII long, curved, simple and sharply pointed, length not substantially in- creasing from segment V to segment VII; 9-VIII plumose with 7—11(8) branches aris- ing from a normally thickened non-flat- tened central stem. Genital lobe: Length about 0.25 mm in female; about 0.45 mm in male, with nipple at apex. Paddle: Lightly pigmented (hyaline), buttress and midrib slightly darker, midrib distinct to near seta 2-Pa; length 0.64—0.77 mm (mean 0.71 mm), width 0.43—0.56 mm (mean 0.50 mm), index 1.35—1.48 (mean 1.42); outer part with spicules ending before seta |-Pa, 37] marginal serrations (refractile border) begin 0.26—0.40 from base and end 0.48—0.66 from base where they grade into short hy- aline filaments; refractile index 0.25—0.43 (mean 0.35). Seta 1-Pa long, sinuous, with hooked tip, about one-third length of pad- dle; 2-Pa well developed, relatively long, with 2—5(4) branches. Larva, fourth-instar (Fig. 2).-Character and positions of setae as figured; numbers of branches in Table 2. Head: Length 0.60— 0.72 mm (mean 0.67 mm), width 0.64—0.77 mm (mean 0.71 mm); moderately tanned, darker patches behind setae 5—7-C and pos- terior to eyes, collar and dorsomentum darkly tanned. Seta 2-C single, aciculate or frayed in distal half; 3-C generally single but often with aciculae or dendritic pro- cesses; 4-C single, rarely double, relatively long, extending well beyond base of 2-C; 6,7-C relatively short, about half length of 5-C, 11-C with comparatively few branches (8-18, commonly 15). Antenna: Moderate- ly tanned; mesal and ventral surfaces with relatively sparse needle-like spicules; length 0.20—0.28 mm (mean 0.25 mm). Seta 1-A small, about as long as diameter of antenna, single, inserted about one-third distance be- tween base and apex of antenna. Thorax: Integument hyaline, smooth. Seta 1-P on small setal support plate, with 5—8(7) rather widely spaced branches; 2-P on margin of plate bearing seta 3-C, single; 11-P signi- ficantly larger than 11-M,T, with 2—4(3) branches; support plate of pleural setal group 9—12-P with small lateral spine, 9- P.M,T and 10-T always branched, 10-M,T and 12-PM,T often single but sometimes with 2 or 3 branches, 12-T more often tri- ple; 14-P with relatively few branches (2— 5, usually 3); 4-M usually with 2 or 3 branches arising from short stem, occasion- ally with 4 branches, rarely single or with 5 branches; 6-M rather long, usually with 3 or 4 branches arising from short stem, range 3-8 branches; 7-M farther ventrad of 6-M than usual (not evident in Fig. 2), with 3— 6(4) branches arising from short stem; 3-T very often single, sometimes with 2 or 3 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘pajunos jou = ou ‘sassa0oid ontsipuap Jo avpnoioe Buoy YM a[suls A[[vssuaH ‘sassaooid ayl][-Yyouvig Japuays 10 avpnoiow /—-Z YIM dISUIS x i — == == —- == == os — — = == = ou Cl == I I I I I I a = as (9) 8-+ (p) S-€ ou fp] (9) 8-€ (€) S-€ (ty) S-€ (¢) S-€ (¢) 9-+ (tv) 9-€ (€) p‘€ (9) 6-S (€) S-@ (C)is=2 EI (Z) €-1 Cs—c @e-1 (Oye=I (Z) €-I (€) S-Z ici CDiget (Migr (HiIS=e rl (7) v-7 (Z) S-Z (Z) €-1 (2) c= (1) €-1 (€) pI I I (Direc (GW) S18) Sur (€) v-I (Z) €-1 (Z) 7 'I ie-1 (€) --I CDical (8) pI-€ (1) ZT (1) €-1 Oi’ Te OL (p) S-€ (bp) S-€ @ Se OOS SG@ieL (S) 6-+ (8) €I-9 (9) OI-Z (6) ZI-S (p)9-% 6 @is-t (ict @Mic-I (Z) € ‘7 (Z) p-7 — (61)SZ-IIl (SI)07-8 (61) P7-EI (e)rI 8 (e) r‘€ (€) S-€ (p) 9-€ (yp) L-€ (OZ) ZE-ZL, (0%) 7e-FI1_~—s (81) STS @o-ce CEDILIETI C1) SIS0L WL (€) S-€ (Qis=s (Sc) i-p= (EDISI-= MED Isl (6Dieceri (€) p-7 (€) 8-€ li (ODA ey) CP ise mG) Svan OIE b, (S) 9-€ (vy) S-€ (S) 9-b (9) 8-9 (vy) L-€ LE-tl Io UCECCS-8 K6SISE. US ie (AS ote: AGO aM I (€) vl @it-z (S)IS—Z (b) 9-€ (bp) 9-€ (€) €-1 (tI C©D6i-1l (Mert (OG=San AS) eC Cea I Qa (€) p-Z I I I GD isel Dial lL) Qe Wee (SiN iat (G)eEe (Syst I I Wiz (S) S-I @yiee (€) p-7 I Cicat (8) OI-L be oS Gis Ie @ie—1e i= (8) 6-S (SQ) OI-C —=(S)01-r (9) 8-¢ (bp) 9-€ (€) p-@ (Glee ECO 91 (D8=6 Te == I I I I I I I = a = I i 0) Pe a ee ee ee ee ee ee ee pe SS TS SS eS a bo ee ee ee L W d o oN peoyH 19S s}usWIZ9g FeUTWOpgY xvloUu ee ‘ISWUDYSUID (DIIJAD) Sajaydouy JO IeIOS [BAIR] IeISUI-YLANOJ JOF sayouvsq JO ssoquinu JO (9pout) asuey “7 FIGP.L VOLUME 108, NUMBER 2 378 NN Up Y/ +—— 0-5 ——J Fig. 2. Fourth-instar larva of Anopheles (Cellia) ainshamsi. A, Head, dorsal (left) and ventral (right) aspects of left side. B, Thorax and abdominal segments I-VI, dorsal (left) and ventral (right) aspects of left side. C, Abdominal segments VII—X, left side. A, antenna; C, cranium, P, prothorax; M, mesothorax; MANP, median accessory tergal plate; S, spiracular lobe; T, metathorax; TP, tergal plate; I-VUI,X = abdominal segments I— VIII and X: 1-15 = setal numbers for specified areas, e.g., seta 5-C. Scales in mm. 374 branches; 11-T minute, sometimes absent; 12-T usually triple, sometimes double, rare- ly single. Abdomen: Integument hyaline, smooth; tergal plates on segments I-VI, roughly triangular, small, width 0.1 or less diameter of segments, median accessory tergal plates on segments II-VI. Seta 1-I weakly developed, with normal branches, usually triple, occasionally double, rarely with 4 branches, 1-II—-VII palmate, with rel- atively few leaflets, leaflets darkly pig- mented distally, generally lanceolate but some with weakly developed shoulders and short filaments; 3-I-HI,V,VI fairly long, sin- gle (3-V occasionally double), 3-IV usually triple (2-4), 3-VII frequently double or tri- ple but most often single; 9-I,II inserted more or less mesal to seta 7; 2-II, III, VII branched, 2-IV—VI single (2-VI rarely dou- ble); 6-[V—VI well developed, with long branches arising well beyond the base of central rachis. Pecten plate moderately tanned, darker on anterior margin, with 12— 17(15) spines, spines with denticles princi- pally on dorsal margins, one to few minute spicules on ventral margins, longer spines usually at each end with few interspersed among shorter spines. Seta 1-S large but with only 4—6(5) branches; 2-S_ usually with 4 or 5 branches (2—7). Saddle moder- ately tanned, short, dorsal length 0.24—0.30 mm (mean 0.27 mm), lateral margins irreg- ular in outline. Seta 1-X inserted on margin of saddle, long, about twice length of sad- dle, single or double, more often single; 2- X with 12—21(15) branches, most branches on dorsal side of rachis, relatively straight, with fine tapering tips; 3-X with relatively few (5-9, mode 6) mostly long, thick, slightly curved, apically hooked branches; 4-X (ventral brush) with 9 offset pairs of setae. Anal papillae very small, short, ba- cillus-shaped, length about 0.09 mm. DNA sequence.—Specimens available for this study included the type series that comprises material collected in 1983 (see below) and a series of pinned adults taken from a laboratory colony maintained at Harvard University in 1982. Ten of these PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON specimens, including two paratypes, were used for DNA extraction. Unfortunately, PCR amplification of the extracted DNA was unsuccessful using primers for the nu- clear ITS2 region and the mitochondrial COI gene. Specimens preserved explicitly for DNA studies are needed for the molec- ular characterization of An. ainshamsi. Etymology.—The species is named in recognition of Ain Shams University, Cai- ro, Egypt, and its support of mosquito bi- ological research and vector control. Systematics.—Anopheles ainshamsi, originally identified as An. stephensi, was first discovered near Ras Gharib on the Gulf of Suez coast in 1966 (Gad 1967). Larvae and a few adults reared from larvae were sent to Dr Peter Mattingly in London, who noted ‘“‘marked differences’’ between these specimens and specimens of An. stephensi in the BMNH (Gad and Kamel 1967). De- spite the apparent differences, the Egyptian specimens were added to the museum’s col- lection (now the NHM collection) of An. stephensi. The specimens include seven lar- vae mounted on a single microscope slide, four pinned females, and a pinned male with dissected genitalia on a microscope slide. The microscope slides are labeled ““An. (Cellia) stephensi/EG YPT/Shokeir/ near Ras Ghareb/x1:1966/A.M. Gad/From brackish swamp/near seashore, Red/Sea coast’’, and the pinned adults each bear a label inscribed with ““EGYPT/Shokeir/near Ras Ghareb/xi:1966/Brackish swamp/near seashore”’. Thus, it seems that Mattingly did not seriously question Gad’s identifica- tion of the species despite the differences he observed in the adult and larval stages. After quoting the differences noted by Mat- tingly, Gad and Kamel (1967) suggested that the “‘marked differences in the Egyp- tian material might indicate that the mos- quito has existed for a long time in the area.” Gad (1967) may have used Mattingly and Knight’s (1956) keys to the mosquitoes of Arabia (the Arabian Peninsula south of the Sinai, Jordan and Iraq, and adjacent islands) VOLUME 108, NUMBER 2 to identify the Anopheles mosquito from the Gulf of Suez coast. Adult females and lar- vae of An. ainshamsi both key to An. ste- phensi in these keys. Although the two spe- cies are similar, they are easily distin- guished by the characters listed in Table 3. Some of these characters are illustrated and used to distinguish adult females of the two species in Glick’s (1992) pictorial key to the anopheline mosquitoes of southwestern Asia and Egypt. Females of An. ainshamsi lead to An. dancalicus in the pictorial key to the anoph- eline mosquitoes of Ethiopia constructed by Verrone (1962a), and are also identified as this species in the computer key to the Anopheles of the Afrotropical Region de- veloped by Hervy et al. (1998). Females key to couplets that distinguish An. salbati and An. dancalicus, and are identified as An. salbaii, in the keys to the Afrotropical anophelines by Gillies and de Meillon (1968) and Gillies and Coetzee (1987). It should be borne in mind, however, that An. hervyi is not included in the last two keys because it was unknown when these keys were developed, and An. salbaii is not in- cluded in Verrone’s key because it is not known to occur in Ethiopia. Hervy et al. (1998) included An. hervyi in their com- puter key even though it was not formally described until the following year. Larvae of An. ainshamsi key to An. dancalicus in the Verrone’s (1962b) pic- torial key to the anopheline larvae of Ethi- opia. They also key to this species in the keys of Gillies and de Meillon (1968) and Gillies and Coetzee (1987), but they are not identifiable as either An. dancalicus or An. salbaii in the computer key of Hervy et al. (1998). As in the case of Verrone’s key for adult females, his key for larvae does not include An. salbaii. Unfortunate- ly, the immature stages of An. hervyi are unknown. Pupae fail to be identified as either An. dancalicus or An. salbaii in the keys of Gil- lies and de Meillon (1968) and Gillies and Bis Coetzee (1987), which are the only avail- able keys for the identification of this life stage of Afrotropical Anopheles. Identifi- cation terminates at couplet 25 because seta 1-V,VI is long in An. ainshamsi and must be short to key to An. dancalicus and An. salbaii. Anopheles ainshamsi obviously belongs to the Neocellia Series, a group of species that breed in open temporary pools of water and are characterized by the presence of broad scutal scales and the absence of upper proepisternal setae in the adults. Larvae have one long mesopleural and two long metapleural setae branched. The series, as currently defined, includes 16 species divid- ed between three species groups and 14 species, including An. dancalicus, An. herv- yl, An. salbaii and An. stephensi, that are unassigned to species groups (Harbach 2004). As An. ainshamsi does not exhibit features that diagnose any of the currently recognized species groups, and shares sa- lient anatomical features with four unas- signed species, it must be regarded as an- other unassigned species of the Neocellia Series. Based on overall morphological similarity, as indicated in Table 3 and re- flected in the use of the identification keys mentioned above, An. adinshamsi appears to be more closely related to An. dancalicus than to the other three species. Areas of the scutum and wings of the adults bear the same ornamentation and markings, pupal setae 1-V—VII and 7-VI,VII are similarly developed, and the leaflets of larval setae 1-IV—VII are lanceolate (usually) or have weakly developed shoulders and a short fil- ament. Bionomics.—Larvae of An. occur in shallow clear saltwater pools, usu- ally shaded by halophilic shrubs, Avicennia marina (Forsk.) (Avicenniacae: Verbena- ceae), and various grasses. The water some- times contains dense mats of grass and fil- amentous green algae. Larvae are also abundant in depressions and drilled holes without vegetation. Larvae of Ochlerotatus ainshamsi PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 376 JUdSa1g JUdSIIg sods yep JOUTSIP © OVC UA JUDSAIg eJSOO UO UISLIO 0} [RISIP SUIS -9q “Y UIOA UO 19.10Y4S pouoney ‘oyduuts oys Joye ‘sired p ajeymjeds Ajpediourg JUdSII1g }UdSd1g sjods ep ¢ YUM ATES) juasgy BISOD UO UISLIO 0} [RISIP suIsaq ‘yy UIDA UO Jaz10Ys AT]eNnsp juasqy ajyeymeds Ajjedioung juasqy JUOSgGYW juasqe 3ods [Pe -seq ‘sjods yep Z WA juasqy RISOD UO UISLIO 0} [RISIP SUIS -aq *Y UIOA UO IazI0YS aspo JaYysIeNsS SUO|R PayeLlas ‘d¥1] -apriq ‘peoig ‘sired g—¢ unlojisny Aypedioutg Ayuo sj}ods ged jeordvosiop Aye -nsn ‘juasqe 10 JUasaIg JUDSdI1g sjods yep snonoidsuoos ¢ YA juasoid Ajjens~y RISO UO UISIIO 0} [BISIP SUIS -3q “Y UIBA UO 1aI0YS > -ejas ‘aduiis “suoy Jaye ‘sired ¢ 10 Z ajyepmeds Ajpedioursg JUDSOId juasqy juasqe sAvmye ods [er -seq ‘sjods yep ponp -qns ZT NOYWM JO YA juosqe A]yens~ Y UIOA pur e}JsOoqgns ‘ejsoo uo Suoy Atjenby snsvapar jo sjayeaj— p soyeos [euTWOpqy snsieja1o} JO sjurof ssoioe spurq aed UIDA [Rue JO xode ye jods asus ayeg SUIM JO UIA [Ruy Sy uraa uo jods yieq jods yrep 10}DasaIg MOL JUOU MOI ]UU { -tuoid s10ul Jopre0g uy] MOI MOLIeU UT -TWoOId a10U Japie0g UT MOI MOLIRU UT sayeos iepeeidns JUaSaI1g Judas JUdSa1dg }UISIIg juasqy SQ[ROS [BSSO} [BINDS i. snojouinu ‘y.1eq May ATOATLIAI ‘ale snojouinu ‘yIeq Maj A[AANPIAI ‘ayeq = SayTBOS [e.QUDDOSIOpaAUYy OU AA aIYM Aj[e1suayH MO][OA A][e1QUaDH ayM AjperduayH MO]JOA A]Te1AUdH SuUI[BOS [eINIS (penods) payros € { -oyed pure -y1ep Ayjenby poyeos-y1eq poyeos-yrep A[urey poyeos-ayed Ayjurepy ~—s pure Z sarauodjed— p sjods ajed YM ¢ o10lU JUISI1g JUISd1g juosqy }UISIIg jUdSo1g -odyjed jo alppru— 4 sy[npVy 1AALOY “UY 1Dq]DS “Uy snIDIUDp “Uy isuaydajs “uy ISUUDYSUID “Uy soinjeoj 2501S jeormojeuy v yey) saiworpur (Z) yaeu UONsenb y ‘saLiag eIJIOIN ay) JO satoads payefar inoy pue iswyYysuID “Uy UZdM}9q SadUaJEJJIP [edioutid Jo uosteduroD ‘uMOUyUN aie Masay “uy Jo ednd pure ‘ease ‘oyeuu oY, “‘UMOUYUN st sin voy ‘€ 31981 S/T VOLUME 108, NUMBER 2 oC nm kx "a G é é }UOUI -R[y pure siaprnoys JOUNSIP YIM soyourig [SUIS afduuts oyduris poyouriq 10 o]Surs ‘UINS.19} JO YSUI] jjey uey) ssay uoys uINnS.1a} URY) 108 -uO] pue o[surs AyTensA oy duis Ajyensn ‘jnojs ‘suo7y xode junyq uM ayIpsag JUDUIe|Y 110Ys pur SJopynoys yeom YIM IO d]v[OINuR] SoyouPIg [SUIS oyduis ayepnoroe 10 ayduig gI3 -uls Ayjensn ‘wuins.19} JO Ysus] jnoge ‘Suo7y uins.1a} uy) JOSUO] JOU ‘payouRIg avon JL] Jo -1q A[jensn ‘rap -ug]s AJOANRIOI ‘SuOT] xode aynor YIM oylpourds ssa] 10 a10jy }UOU -R[Y pure siapynoys JOUNSIP WIM soyourig a[Suls oyduuig pokey IO ayeyNo1INR UdJO poyouriqg Ayyensn “UINS.10] JO WSUS] jyey uey) ssoy 0ys uins.19} ueY) Jasuo] ‘apsuis Ayjensy~ oyduits ‘ynoYs ATAAVIA1 ‘SuoT7T xode junyq WWM oyI[seq JUSUIe]Y WOYS pur slop[noys yeam YIM JO aye[OoouR] soyouPsg a[qnop 10 aysuls podAvly 10 ae[noIoy pokey 10 ayepnoly opsurs Aypensn ‘uims.10} JO YSug] ynoqe ‘suoT uns -19] UB) JaSUO] ‘aySuIS ‘SuO'T] xode junyq 1oyyer YIM ayl[sod ssa] 10 dO ojduuts ‘1apueys HA-AI-[ 9819S xX-[ BIOS O-€ BIBS O-T BES HA TA-L ®19§ ITA-A-T 89S HA-A-6 8319S AL6 89S SBAIV™T] ordng rs IMAMOY “UW 1Dq]DS “Uy SNIDIUDP “Uy isuaydals “uy ISUIDYSUID “UY ‘ponunuog "€ a1qeL 378 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ics 3) lia) ainshamsi near Ras Shukeir on the Gulf of Suez coast. Typical breeding site of Anopheles (Cel- detritus (Haliday) also occur in these hab- itats. The type series consists of larvae and specimens reared from larvae and pupae collected from salt encrusted pools and drilled holes ranging from 0.2—2.0 m in di- ameter. A typical saline pool inhabited by larvae is shown in Fig. 3. The pH and salinity of breeding sites were recorded using methods recommended by the World Health Organization (WHO 1975). The pH of pools near Ras Gharib ranged from 6.9—7.3; those that were not encrusted with salt contained 33-37.5 g Cl/ liter whereas those encrusted with salt contained 54—72 g Cl/ liter. Pools at the site north of Ras Shukeir were more saline and more acidic: 72—77.5 g Cl/ liter, pH 5.8—6.3. However, An. ainshamsi develops well in water of various salinities. Gad et al. (1987) reared them successfully both in seawater (21—22 g Cl / liter) and water with only 9 g Cl / liter. Anopheles ainshamsi occurs in areas that are uninhabited (Riberio et al. 1985, Kenawy 1988), but females will attack humans who visit their realm (Gad and Salit 1972). Lizards, deer, and passerine birds have been seen near breeding places (Gad and Kemal 1967, Ribeiro et al. 1985), and camel skeletons were observed in the salt springs area, but whether fe- males feed on these or other animals is unknown. As noted previously, An. ain- shamsi is an autogenous mosquito capable of developing and laying eggs in the ab- sence of hosts (Ribeiro et al. 1985). Distribution.—Anopheles ainshamsi is known only from coastal areas of Ras Shu- keir District, El-Bahr El-Ahmar (Red Sea) Governorate, Egypt. Larvae have been col- lected from sites not far from the Gulf of Suez coast. One site, where this species was originally discovered and identified as An. stephensi by Gad (1967), is located near Ras Gharib. Another site is located about 19 km north of Ras Shukeir town, and the specimens that comprise the type series in- clude fourth-instar larvae and adults reared from larvae and pupae collected about 5 km west of Ras Shukeir. Type series.—Two hundred and eighteen specimens (2 adults used for DNA extrac- tion; failed PCR): (32 25533 6,46 ent talia, 40 Le, 65 Pe, 44 L. Holotype, @° (EG146-34), with LePe on microscope slide, EGYPT: El-Bahr El-Ahmar (Red Sea) Governorate, 5 km W_ of Ras Shukeir, mixed small ground pools and holes, some drying up and encrusted with salt, 30 April 1983 (Harrison, Gad, Gamal). Paratypes (same locality and collectors as holotype), 25 2LePe (EG146-2, -4, -5, -6, -9, -11, -12, -18, -23, -25 through -40; EG146-23 used for DNA extraction, failed PCR); 14 SLePe (EG146-1, -3, -7, -8, -10, -13 through -17, -19 through -22, -24; EG146- 1, -3, -7, -8 with dissected genitalia on mi- croscope slides; EG146-14 used for DNA extraction, failed PCR); 7 2 Pe (EG146-100 VOLUME 108, NUMBER 2 through -103, -106, -109, -121); 18 dPe (EG146-104, -105, -107, -108, -111 through) -1207=122> -123, -124): "44 1 (EG146). The holotype (EG146-34) and the following paratypes are deposited in the NMNH: EG146-1, -2, -3, -6 to -9, -11 to -16, -18 to -22, -24 to -40, and 40L. The remaining paratypes are deposited in Ain Shams University (EG146-4, -17, -103, -104, and 2L) and the NHM (EG146-5, -10, -14, -23, -109, -122, and 2L). ACKNOWLEDGMENTS We are grateful to Ali Hafez and Saad El Masry for assistance in the field in Egypt; to Taina Litwak (formerly of WRBU) for preparing pencil drawings of the larva and pupa; to Theresa Howard (NHM) for mod- ifying and inking the pencil drawings and illustrating of male genitalia; to Joanna Elphic (formerly of NHM) for recording larval and pupal chaetotaxy; and to Yvonne Linton and Fredy Ruiz (NHM) for con- ducting DNA extractions and PCR in an at- tempt to obtain DNA sequences from avail- able specimens. LITERATURE CITED Brunhes, J., G. le Goff, and B. Geoffroy. 1999. Afro- tropical anophelines mosquitoes. III. Description of three new species: Anopheles carnevalei sp. nov., An. hervyi sp. nov., and An. dualaensis sp. nov., and resurrection of An. rageaui Mattingly and Adam. Journal of the American Mosquito Control Association 15: 552-558. El-Said, S. and M. Kenawy. 1983. Geographical dis- tribution of mosquitoes in Egypt. Journal of the Egyptian Public Health Association 58: 46—76. Gad, A. M. 1967. Anopheles stephensi Liston in Egypt, UAR. Mosquito News 27: 171-174. Gad, A. M. and O. M. Kamel. 1967. Further notes on Anopheles stephensi in Egypt, U.A.R. Journal of the Egyptian Public Health Association 42: 249— DSP: Gad, A. M. and A. Salit. 1972. The mosquitoes of the Red Sea area, Egypt. Journal of Medical Ento- mology 9: 581—582. Gad, A. M., S. El Said, A. N. Hassan, and A. Shoukry. 1987. The distribution and ecology of the mos- quitoes in the Red Sea Governorate, Egypt. Jour- nal of the Egyptian Society of Parasitology 17: 207-221. Gad, A. M., W. A. Maier, and G. Piekarski. 1979a. S79 Pathology of Anopheles stephensi after infec- tion with Plasmodium berghei berghei. 1. Mor- tality rate. Zeitschrift fiir Parasitenkunde 50: 249-261. . 1979b. Pathology of Anopheles stephensi after infection with Plasmodium berghei berghei. I. Changes in amino acid contents. Zeitschrift fiir Parasitenkunde 60: 263-275. Gillies, M. T. and M. Coetzee. 1987. A supplement to the Anophelinae of Africa south of the Sahara (Afrotropical Region). Publications of the South African Institute for Medical Research 55: 1— 143. Gillies, M. T. and B. de Meillon. 1968. The Anophel- inae of Africa south of the Sahara (Ethiopian zoo- geographical region). 2nd ed.. Publications of the South African Institute for Medical Research 54: 1-343. Glick, J. I. 1992. Illustrated key to the female Anoph- eles of southwestern Asia and Egypt (Diptera: Cu- licidae). Mosquito Systematics 24: 125-153. Harbach, R. E. 2004. The classification of genus Anopheles (Diptera: Culicidae): a working hy- pothesis of phylogenetic relationships. Bulletin of Entomological Research 94: 537-553. Harbach, R. E. and K. L. Knight. 1980. Taxonomists’ Glossary of Mosquito Anatomy. Plexus Publish- ing, Inc., Marlton, New Jersey. xi + 415 pp. 1982 [1981]. Corrections and additions to Taxonomists’ Glossary of Mosquito Anatomy. Mosquito Systematics 13: 201-217. Hervy, J.-P, G. le Goff, B. Geoffroy, L. Hervé, J.-P. Manga, L. Manga, and J. Brunhes. 1998. Les anopheles de la région afro-tropicale. CD-ROM. ORSTOM, Montpellier, France. Kenawy, M. A. 1988. Anopheline mosquitoes (Dip- tera: Culicidae) as malaria carriers in A.R. Egypt “history and present status”. The Journal of the Egyptian Public Health Association 63: 67-85. . 1990. Fauna of anopheline mosquitoes (Dip- tera: Culicidae) in A.R. Egypt “historical back- ground and present situation’. The Journal of the Egyptian Public Health Association 65: 263— 281. Mattingly, P. F and K. L. Knight. 1956. The mosqui- toes of Arabia I. Bulletin of the British Museum (Natural History) Entomology 4: 91-141. Ribeiro, J. M. C., P. A. Rossignol, and A. Spielman. 1985. Salivary gland apyrase determines probing time in anopheline mosquitoes. Journal of Insect Physiology 31: 689-692. Verrone, G. A. 1962a. Outline for the determination of malarial mosquitoes in Ethiopia. Part I-adult fe- male anophelines. Mosquito News 22: 37—49. . 1962b. Outline for the determination of ma- laria mosquitoes in Ethiopia. Part II— anopheline larvae. Mosquito News 22: 394—401. Ward, R. A. 1992. Third supplement to “A Catalog of 380 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the Mosquitoes of the World’? (Diptera: Culici- | Wilkerson, R. C. and E. L. Peyton. 1990. Standardized dae). Mosquito Systematics 24: 177-230. nomenclature for the costal wing spots of the ge- WHO. 1975. Manual on Practical Entomology in Ma- nus Anopheles and other spotted-wing mosquitoes laria. Part II. Geneva, WHO offset publication, no. (Diptera: Culicidae). Journal of Medical Entomol- 13. 191 pp. ogy 27: 207-224. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 381-388 DESCRIPTION OF THE LARVA OF GOMPHUS SANDRIUS TENNESSEN (ODONATA: GOMPHIDAE) KENNETH J. TENNESSEN 125 N. Oxford St, Wautoma, WI 54982, U.S.A. (e-mail: ktennessen @centurytel.net) Abstract.—The final stadium larva of Gomphus sandrius Tennessen is described based on reared specimens from Tennessee. The larva is distinct from G. exilis Selys and G. lividus Selys (the only species in the subgenus Gomphus sympatric with G. sandrius) by the greater width to length ratio of abdominal segment 9 venter (mean 1.82, range 1.69— 1.96 in G. sandrius vs. mean 1.43, range 1.23—1.57 in G. lividus and mean 1.40, range 1.26—1.52 in G. exilis). It differs further from G. lividus in the narrower prementum (2.40— 2.90 mm vs. 3.00—3.75 mm and shorter antennal segment 3 (1.15—1.35 mm vs. 1.50—1.90 mm). The larva of G. sandrius is most similar to the allopatric G. graslinellus Walsh, but antennal segment 3 is shorter (G. sandrius: mean 1.25 mm, range 1.15—1.35 mm: G. graslinellus: mean 1.45 mm, range 1.35—1.55 mm). Key Words: Gomphus sandrius Tennessen is a rare dragonfly restricted in geographic range to south-central Tennessee (Dunkle 2000, Donnelly 2004). There are seven known lo- calities in five contiguous counties in the Central Basin (Fig. 1). Bick (2003), who listed Alabama (Colbert Co.) in error, rated the species “critically imperiled” (Natural Heritage category Gl); only 3 of the 27 odonate species considered to be “‘at-risk”’ in the United States were rated as G1. For possible future conservation efforts, ability to distinguish the larva of G. sandrius from its congeners is critical. Gomphus sandrius belongs to the sub- genus Gomphus as defined by Needham et al. (2000), who provided a key for all lar- vae of the subgenus except for the previ- ously unknown larva of G. sandrius. Lar- vae of Gomphus are relatively difficult to identify because of similarity in form, in- traspecific variability, and lack of a detailed comparative study of all species in the sub- genus. Previous errors in association also Odonata, Gomphidae, Gomphus, larva, Tennessee account for some of the difficulty. For ex- ample, the larva of Arigomphus lentulus (Needham) was mistaken for Gomphus mil- itaris Hagen by Bird (1934) and this error existed until Landwer and Sites (2003) cor- rected it. I associated larvae and adults of G. sandrius and herein provide the follow- ing description, illustrations, and diagnosis based on exuviae and other preserved lar- vae from several locations. MATERIALS AND METHODS I collected Gomphus larvae with an aquatic dip net in sand and gravel substrates of small, shallow tributaries in central Ten- nessee. Live individuals were transported to Florence, Alabama, and reared in an aerated aquarium. Specimens were preserved in 80% ethanol. Morphological terminology follows that of Needham et al. (2000), except the apical tooth on the labial palp is called the end tooth, not end hook. Abdominal segments are abbreviated with an *‘S” preceding the Ww ie) i) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. “apical angle” (CQ) of abdomen in Gomphus larvae. 3, Measuring length of posterolateral spine of abdominal S9, ventral view. number of the segment (e.g., S10 = abdom- inal segment 10). Measurements were taken with an ocular micrometer on a Wild ste- reomicroscope, making sure that each struc- ture was perpendicular to my line of sight; a camera lucida was used to draw the fig- 1, Map of central Tennessee showing known distribution of Gomphus sandrius. 2, Measuring ures. Length of antennal segment 3 was measured dorsomedially from base to apex. Length of hind tarsal claws was measured as a Straight line (even though the claws are curved) from the dorsobasal notch to the apex. Cleaning with a fine brush was nec- VOLUME 108, NUMBER 2 essary to count the serrations on the lateral margins of S6-S9 and to measure antennal segment 3. Length and width of S9 and S10 were measured ventrally. To quantify the degree of taper at the apex of the abdomen, I measured the angle formed by two lines drawn (using a camera lucida) from the an- terolateral margins of S7 to the apex of the epiproct (Fig. 2); the resulting angle was designated as the “‘apical angle.”’ The pos- terolateral spine of S9 was measured ven- trally as shown in Fig. 3. The epiproct was measured dorsomedially; cercus length was measured along the dorsomedial margin. For the five characters in which G. sandrius and G. graslinellus were found to differ, I made no statistical comparisons, but cal- culated 95% confidence limits for the means using Microsoft Excel® (Table 1). Final Stadium Larva of Gomphus sandrius . (Figs. 4—9) Description.—Based on 28 specimens (listed below). Body elongate and dorso- ventrally flattened, general color pale brown with few dark markings, dorsum speckled with tiny raised dots and numer- ous, long, hairlike setae, abdomen lanceo- late (Fig. 4). Total length 24.5—30.0 mm. Head: Width 5.17—5.90 mm; rounded anteriorly, posterolateral corners slightly produced. Antennal segment 3 length 1.17— 1.37 mm, 3.2—3.4 times longer than wide. Prementum 2.42—2.87 mm wide at distal margin, 2.71—3.20 mm long, lateral margins indented in basal third from which margins converge slightly to distal margin (Fig. 5), rarely parallel; ligula convex (Figs. 5 and 6a); palpal lobe incurved, end tooth usually slightly longer than adjacent tooth (Fig. 6a), sometimes twice as long (Fig. 6b). Thorax: With dorsolateral, diagonal, dark brown stripes (Fig. 4). Apex of hind wing pad extending to apex of S4 or to an- terior half of S5. Fore and middle tibiae with well-developed apical burrowing hook; hind femur length 4.9—5.7 mm, distal end extending nearly to posterior margin of 383 S4 (Fig. 4); hind tarsal claw 0.83—0.93 mm long. Abdomen: Widest at S5. Lateral spines on S6-S9 increasing in length posteriorly (Fig. 4), on S9 ranging from 0.53—0.77 mm long and extending to about midlength of S10. S2-S9 each with a middorsal posterior prominence, which on S3-S9 bears a dis- tinct dorsal hook that overlies intersegmen- tal membrane (Fig. 7). Posterior margins of terga 6—9 with stout spines, small and pale on S6, darker and more developed on S7- S9 (Fig. 8). S9 with middorsal, full-length, rounded ridge (Figs. 4, 8, 9), S8 with sim- ilar ridge on posterior half only. Lateral margins of S6-S9 appearing serrated due to small stout spines, numbering 0—4 on S6, 3—13 on S7, 9-18 on S8, and 16—28 on S9; distance between most serrations on S8 and S9 less than basal width of serrations. Width/length ratio of S9 ranging from 1.69 to 1.96; width/length ratio of S10 about 1.22—-1.45. Apical angle (Fig. 2) 46—54° (mean = 50°). Epiproct 1.15—1.35 mm long, usually longer than cerci (1.03—1.21 mm) and paraprocts, occasionally shorter (Fig. 8); tips of cerci sharply acuminate, tips of paraprocts much more blunt than cerci. Specimens examined (n = 28).—TEN- NESSEE: Bedford Co.: Weakly Creek, Halls Mill Road, V-11—1983, KJT, 1 asso- ciated female exuvia, 3 unassociated exu- viae; X-10-1083, 2 larvae, KJT; Marshall G€o= Wilson Creek, 3.2 km SE of Chapel Hill, V-11—1983, KJT, 5 exuviae; Maury Co.: Flat Creek, Hwy. 431, IV-26-1984, KJT, 1 associated male exuvia, 7 larvae; XI- 4-1983, KJT, 4 larvae; Rutherford Co., Middle Fork Stones River, near Elam Rd., I-15-2004, KJT, 4 larvae; Wilson Co.: Round Lick Creek, nr. I-40, III-4-1998, J. S. Tindell, 1 larva. All specimens are de- posited in the Florida State Collection of Arthropods. For comparative purposes, I examined 24 larvae/exuviae of each of the following: G. exilis” Selys: (Als (GA: NGPA, SG; TN: WV, WI), G. graslinellus Walsh (AR, MO), 384 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Gomphus sandrius larva, dorsal aspect. G. lividus Selys (AL, GA, MI, SC, TN, WI), and G. minutus Rambur (FL, GA). Diagnosis.—In the larval key by Need- ham et al. (2000: 310—312), G. sandrius keys to G. graslinellus Walsh, a widespread species that is mainly Midwestern in distri- bution. These species are allopatric (see Donnelly 2004): the nearest record of G. graslinellus in Arkansas (Mississippi Co.) is roughly 270 km W of the westernmost G. sandrius record (Maury Co., TN), whereas the nearest G. graslinellus record VOLUME 108, NUMBER 2 385 end tooth = 6 b adjacent tooth —” 10 - lividus Figs. 5-10. 5-9, Gomphus sandrius. 10, G. lividus. 5, Prementum. 6a, Ligula and palpal lobe. 6b, Variation in palpal lobe. 7, Dorsal prominences of abdominal S2—9, lateral view. 8, Abdominal S9, S10 and anal append- ages, dorsal view. 9, Abdominal S9 in cross-section. 10, Palpal lobe. in Kentucky (Edmonson Co.) is about 120 (Fig. 11). The larva is similar to G. san- km N of the northernmost G. sandrius re- drius in possessing a convex ligula, a var- cord (Wilson Co., TN). Based on its pres- iable but usually reduced end tooth on the ently known geographic range, it is possible — palpal lobes, S9 much wider than long, and that G. graslinellus occurs in western TN S9 with a full-length middorsal ridge. I 386 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 11. Map of north-central U.S. showing distribution, by county, of Gomphus graslinellus (gray) and G. sandrius (black); note —G. graslinellus ranges further northwest of the area shown on the map. compared larvae of these species based on 22 morphological characters and found sig- nificant differences in 5 characters (Table 1). The most reliable difference was length of antennal segment 3, as no overlap in range was found. Another useful character is the ratio of S9 width to length, as the confidence limits did not overlap with the VOLUME 108, NUMBER 2 387 Table 1. Summary of five morphological characters for larvae of Gomphus sandrius and Gomphus grasli- nellus; measurements for antennal segment 3 length and hind tarsal claw length in mm (C.L. = confidence limits around the mean). Character Species n Mean Range 95% C.L. AntSeg3L G. sandrius 24 1.27 1.17—1.37 1.25—1.29 G. graslinellus 24 1.45 1.37-1.53 1.43-1.47 S9W/L G. sandrius 24 1.82 1.69-1.96 1.79-1.85 G. graslinellus 24 1.65 1.46—-1.77 1.62—1.68 CercL/EpiL G. sandrius 24 0.93 0.87—1.02 0.92—0.94 G. graslinellus 24 0.86 0.80—0.93 0.85—0.87 PrementL/W G. sandrius 24 1.13 1.09-1.19 1.12-1.14 G. graslinellus 24 1.20 1.13—1.28 1.19-1.21 HindTarsClawL G. sandrius 24 0.89 0.83—0.93 0.87-0.91 G. graslinellus 23 0.99 0.89-1.05 0.97-1.01 range of either species. The confidence lim- its of 1) cercus length to epiproct length, 2) prementum length x width, and 3) hind tar- sal claw length did not overlap; however, some Overlap between the confidence limits and the ranges in these characters was found and therefore these characters are less reliable for differentiating G. sandrius and G. graslinellus larvae. Three species of Gomphus (Gomphus) are recorded in Tennessee other than G. sandrius, namely G. exilis Selys, G. lividus Selys, and G. quadricolor Walsh. In G. quadricolor, S9 and S10 are about as wide as long (L/W ratio of S9 = 0.95-1.10, L/ W ratio of S10 = 0.85—1.08), whereas in G. exilis, G. lividus and G. sandrius, S9 is much wider than long (range 1.23—1.96) and S10 is usually significantly wider than long (range 0.91—1.45). Furthermore, G. guadricolor has small middorsal hooks only on S8 and S9, best seen in dorsal view; even though S2-—7 are slightly prominent posteromedially, no dor- sal hooks are present on these segments. Gomphus sandrius, G. exilis, and G. lividus are more likely to be confused with one an- other in Tennessee. Gomphus lividus is dis- tinct from G. sandrius in the straight ligula (convex in sandrius), greater width of pre- mentum (3.03—3.77 mm vs. 2.42—2.87 mm in sandrius), greater length of prementum (3.20—3.85 mm vs. 2.71—3.20 mm in san- drius), longer antennal segment 3 (1.49— 1.89 mm vs. 1.17—1.37 mm in sandrius), longer hind tarsal claw (1.01—1.21 mm vs. 0.83—0.93 mm in sandrius), and longer S9 (2.79-3.36 mm vs. 2.17—2.62 mm in san- drius). Gomphus exilis has a narrower S9 (2.87—3.69 mm vs. 4.02—4.67 mm in san- drius), narrower S10 (1.15—1.44 mm vs. 1.56—1.78 mm in sandrius), and shorter cer- ci (0.87—1.01 mm vs. 1.03—1.27 mm in san- drius). Total length averaged less for G. ex- ilis (23.3 mm) than for G. sandrius (27.5 mm), but both species are quite variable in size (exilis 21.0—26.0 mm vs. sandrius 24.5—30.0 mm). Other helpful differences are: G. lividus has more strongly curved palpal lobes (Fig. 10), G. exilis has fewer granulations on S9 (2-3 dozen vs. more than 6 dozen in /ividus and sandrius), in G. lividus the diameter of the granules on S9 is at least twice that in exi/is and sandrius (0.05—0.06 mm vs. 0.010—0.024 mm), G. sandrius has dorsal prominences on S2-S9 whereas exilis and lividus lack prominences on S2 and S3. Gomphus sandrius is also morphologi- cally similar to the allopatric G. minutus Rambur that occurs in much of FL and southern parts of GA, AL and SC (Don- nelly 2004). The stout spines (serrations) on the lateral margins of S6—9 in G. minutus are very small and hidden by setae (difficult to detect even at 50X magnification), whereas these spines are larger and more easily detected (visible at 10X) in G. san- 388 drius (Fig. 8). The ratio of S9 width to length was much lower in G. minutus (1.20—1.39 vs. 1.69-1.96 in sandrius), as was the width/length ratio for S10 (0.83— 1.00 vs. 1.19—1.45 in sandrius). The tip of the abdomen in G. sandrius and G. graslinellus is more broadly tapered than in other members of the subgenus Gomphus (in G. sandrius mean apical angle = 50°, range 46—-54°, in G. graslinellus mean = 49°, range 44—53°). Other broadly tapered species of the subgenus Gomphus had lesser values (G. lividus mean = 46°, range 42—50°, and G. exilis mean = 43°, range 40—51°). In this characteristic, G. sandrius is closest to species I measured in the subgenera Gomphurus (range 52—61°) and Hylogomphus (range 58—66°). Remarks.—I found Gomphus lividus in two of the seven streams occupied by G. sandrius (Fall Creek in Bedford County, Middle Fork Stones River in Rutherford County). The microhabitats differed slight- ly: G. lividus larvae usually occupied slow- er edges where more silt/mud had accu- mulated, whereas G. sandrius larvae were usually in mixed gravel with less silt. I did not find G. exilis or G. quadricolor in any of the G. sandrius localities. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ACKNOWLEDGMENTS I thank Carl Cook (Center, KY), Dave Etnier (University of Tennessee), Brett Landwer (University of Missouri), and Bill Mauffray (Florida State Collection of Ar- thropods, Gainesville) for advice and spec- imen loans. LITERATURE CITED Bick, G. H. 2003. At-risk Odonata of conterminous United States. Bulletin of American Odonatology 7(3): 41-56. Bird, R. D. 1934. The emergence and nymph of Gom- phus militaris (Odonata, Gomphidae). Entomolog- ical News 45: 44—46. Donnelly, T. W. 2004. Distribution of North American Odonata. Part 1: Aeshnidae, Petaluridae, Gomphi- dae, Cordulegastridae. Bulletin of American Odonatology 7(4): 61—90. Dunkle, S. W. 2000. Dragonflies Through Binoculars: A Field Guide to Dragonflies of North America. Oxford University Press, Oxford, 266 pp. Landwer, B. H. P. and R. W. Sites. 2003. Redescription of the larva of Gomphus militaris Hagen (Odon- ata: Gomphidae), with distributional and life his- tory notes. Proceedings Entomological Society of Washington 105(2): 304-311. Needham, J. G., M. J. Westfall, Jr., and M. L. May. 2000. Dragonflies of North America. Scientific Publishers, Gainesville, 939 pp. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 389-395 A KEY TO SPECIES OF THE GENUS ONITIS FABRICIUS (COLEOPTERA: SCARABAEIDAE: SCARABAEINAE) FROM CHINA, WITH THE DESCRIPTION OF A NEW SPECIES AND A NEW RECORD FOR CHINA MING BAI, XINGKE YANG, AND YOUWEI ZHANG Institute of Zoology, Chinese Academy of Sciences, 36 box, 25 Bei SiHuanXiLu, Hai- dian, Beijing, 100080, China, (e-mail: baim@ioz.ac.cn); (MB) Graduate School, Chinese Academy of Sciences, Yuquanlu, Shijingshan, Beijing, 100039, China. Corresponding author: Xingke Yang (e-mail: yangxk @ioz.ac.cn) Abstract.—The genus Onitis Fabricius in China includes 10 species, among them Onitis brevidens, new species, and Onitis feae Felsche, a new record for China. The distinctive characters of the new species are discussed. A key to species of Onitis from China, habitus photographs, and illustrations of the protibiae, profemora, mesocoxae, and genitalia are provided. Key Words: China, Onitis, Scarabaeinae, Scarabaeidae, Coleoptera, new species The genus Onitis (Coleoptera: Scara- baeoidea: Scarabaeidae: Scarabaeinae: On- itini) was established by Fabricius in 1798. Scarabaeus inuus Fabricius, 1781 from the Congo was selected by Fabricius as the type species. In 1871, Harold regarded S. inuus as a synonym of Onitis sphinx (Fabricius 1775). Lansberge revised Onitis in 1875, and 61 world species were known at that time. Janssens published a revision of On- itini in 1937, and 113 species of Onitis were described in the monograph. There are 154 worid species of Onitis with ten species currently known in China, including one new species described here. There are 124 species in the Afrotropical Region, 5 spe- cies in the Palaearctic Region, and 25 spe- cies in the Oriental Region. The male legs usually show modifica- tions. Species of Onitis are tunnellers, and their nesting behavior is complex (Philips et al. 2004). They can be collected from various animal dung, such as cow, elephant, gaur, buffalo, and pig (Hanboonsong and Masumoto 2000). In this paper, ten species of Onitis from China are treated, among them Onitis brev- idens, new species, and Onitis feae Felsche, a new record for China. A key to species from China, habitus photographs, and illus- trations of protibiae, profemora, mesocox- ae, and genitalia are given. Onitis Fabricius, 1798 Onitis Fabricius 1798: 2. Type species: Scarabaeus inuus Fabricius, 1781. Desig. by Fabricius 1798. Description.—Male: Oblong in shape, with legs stout and short (except proleg). Head: Shape narrow. Ocular lobes united by a carinate suture with clypeus. Clypeal margin rounded or slightly excised at mid- dle. Antenna 9-segmented, all segments short except basal; club compact, with first segment cup-shaped, smooth, chitinous, en- closing succeeding segment. Mandible oval, with long terminal fringe. Membra- nous lobe of maxilla broad, palpus short. Labium and labial palpus clothed with long 390 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON if | Oe | YY B BBY AB) I) Figs. 1-4. Onitis species. 1, O. brevidens. 2, O. excavatus. 3, O. falcatus. 4, Onitis feae. A = genitalia in ventral view; B = genitalia in dorsal view; C = genitalia in lateral view; D = protibia and profemur; E = mesocoxa. VOLUME 108, NUMBER 2 bristles, palpus flat, basal joint weakly di- lated, 2nd long, 3rd minute. Pronotum: Surface without process or excavation. Base with small prominence at middle, usu- ally without complete margin, but with pit or impression on each side near middle. Scutellum: Visible but minute. Elytron: Sur- face with simple, narrow epipleura, delim- iting carina strongly marked and straight. Abdomen: Completely covered with a ca- rina around sides of all segments and py- gidum. Legs: Proleg elongate; procoxa prominent; protibia slender and_ strongly curved toward apex, armed with 4 external teeth and with | or more teeth on lower face, without articulated terminal spur, but with apex produced into fingerlike process; protarsus absent. Mesocoxa long, parallel, widely separated; mesotibia with angular projection at inner edge. Pro-, meso- and metafemora, or. some of them, toothed at edge, or mesotrochanter spinose. Meso- and metatarsi with progressively shortened seg- ments, basal joint more than twice as long as 2nd. Female: Head sometimes with frontal tu- bercle or short horn. Protibia always broad, with stronger teeth, and with articulated ter- minal spur. Diagnosis.—Onitis is recognized easily by the absence of protarsi, mesotibiae short and dilated, the presence of 2 basal impres- sions on the pronotum, visible scutellum, and strong lateral carina on the elytra. KEY TO ADULT MALES OF ONITIS FROM CHINA 1. Pygidium smooth, glabrous ............. 2 Pygidium more or less setose 2. Frontoclypeal carina simple; pronotum green, elytra dark yellow (Figs. SA—-C, 14) ......... SCE on eA Onitis humerosus (Pallas 1771) — Frontoclypeal carina interrupted or with tuber- cle at middle; pronotum and elytra black ... 3 3. Pronotum feebly punctate, punctures shallow ANGIVACUC) oh sksleis svete SY Se tees Ain Sh ctinedeteie ce 4 — Pronotum distinctly punctate, punctures deep amnG! GIA 54 concacesoaugceaaegede guns 5 4. Metasternum transversely excavated at middle (Figs. 2A—C, 11) O. excavatus Arrow, 1931 — Metasternum not excavated at middle (Figs. 3A=Co 12) ne Oe O. falcatus (Wulfen 1786) 5. Metasternum longitudinally grooved in front OREN OME gE REED. tack Gh b,)ot by-b) O01) ONO neecneaE 6 — Metasternum flat, not grooved ........... 8 6. Ventral surface entirely metallic (Figs. 6A—C, Se a a as Sent O. philemon Fabricius, 1801 Ventral surface not metallic ............. J 7. Basal margin of pronotum straight .......... 5 Me 3 ANS hc O. intermedius Frivaldszky, 1892 — Basal margin of pronotum round (Figs. 7A—C, iC) MR Re ecmea ts cat. chee O. spinipes (Drury 1770) 8. Frontoclypeal carina broadly interrupted; pro- tibia with terminal external tooth projecting in fronti(Figs7 SACs weyeanveweocn eae O. subopacus Arrow, 1931 — Frontoclypeal carina narrowly interrupted; pro- tibia with terminal external tooth tapering in front (Figs. 9A-C, 18) O. virens Lansberge, 1875 9. Profemur and protibia with strong tooth; me- socoxa toothed at its posterior end (Figs. 4A— E13) O. feae Felsche, 1907 — Profemur and protibia without or with feeble tooth; mesocoxa not toothed at its posterior end (Figs. 1A—E, 10) O. brevidens, n. sp. Onitis brevidens Bai, Yang, and Zhang, new species (Figs. 1A—E, 10) Holotype male.—Length 14.3 mm, wid- eth 8.1 mm. Shape broadly oval, compact and convex. Color black, subopaque; head, lower surface, and legs dark red; pygidium and lower surface clothed with brown setae; elytra bearing a fringe of similar setae on hind margin. Head: Clypeus elliptical, front margin imperceptibly notched at middle, surface finely and closely rugulose, with short, transverse, slightly curved or angular cari- na. Frontoclypeal region with straight, sharply elevated carina. Frons slightly ru- gopunctate, not shining. Vertex with short- er, transverse, curved carina immediately before carinate posterior margin of head. Pronotum: Strongly, densely punctate, without smooth median line, but with fee- bly indicated median groove. Base almost completely margined; basal foveae deep with intervening space finely rugose, opaque, distinctly lobed in middle. Elytron: Surface convex and entirely opaque except PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 392 | \ I] \ \ | \ \ oa i ‘a y Sy meal bs C5 dais See A : - a c ~*~ ) solu G Mp Onitis species. 5, O. humerosus. 6, O. philemon. 7, O. spinipes. 8, O. subopacus. 9. O. virens. A Figs. 5-9. = genitalia in ventral view; B = genitalia in dorsal view; C = genitalia in lateral view o>) \O ioe) VOLUME 108, NUMBER 2 16 18 Figs. 10-18. Habitus photographs of Onitis species. 10, O. brevidens. 11, O. excavatus. 12, O. falcatus. 13, Onitis feae. 14, O. humerosus. 15, O. philemon. 16, O. spinipes. 17, O. subopacus. 18, O. virens. 394 at humerus. Shape parabolic; elytra 0.4 times wider than long medially. Striae faint; intervals slightly carinate along middle, outer ones slightly, unevenly punctate. Py- gidium: Surface feebly rugulose, thinly clothed with erect, long, reddish setae. Shape parabolic; 1.9 times wider than long medially. Metasternum: Surface with con- spicuous granules, granules closer and finer at front and sides, with long, erect setae; middle broadly grooved. Abdomen: Sur- face setae at sides, smooth at middle. Legs: Profemur with feeble tooth near middle of front edge. Protibia elongate, with tip strongly curved and produced, with finely serrate carina beneath, without tooth near base (Fig. 1D). Mesocoxa not toothed at posterior apex, mesotibia not excised at base (Fig. 1E). Male genitalia as in Figs. 1A-C. Female.—Length 16.8 mm; width across humeri 8.9 mm. Similar to male except pro- femur without tooth near middle of front edge. Variation in paratype series.—Some of the four paratypes have a much darker el- ytra and pronotum. Measurements: Length 13.4-16.1 mm, width across humeri 7.9— 8.3 mm. Type material.—Holotype: P. R. CHINA. 6, Yunnan: Menghai, 1,250 m, 24 February 1957, coll. Fu-Ji PU. Paratypes: P. R. CHI- NA. 1 ¢, 1 2, Yunnan: Kaida, 1956, coll. unknown; | 6, Yunnan: Jingdong, 1,200 m, 2 March 1957, coll. Monchadsky; 1 2, same data as holotype. Holotype and para- types deposited in the Institute of Zoology, Chinese Academy of Sciences, Beijing, P. R. China (IZAS). Distribution.—China (Yunnan). Diagnosis.—The new species is similar to Onitis feae but differs primarily by its smaller size, darker color, profemur and protibia with a feeble tooth, protibial apex with a short, fingerlike process, and meso- coxa not toothed at its posterior end. Onitis feae is larger, shining, the profemur and protibia each with a sharp, long tooth, pro- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tibial apex with a long, fingerlike process, and the mesocoxa is toothed at its posterior end. Etymology.—The name is from the Latin brevis, referring to the profemur with a fee- ble tooth near the middle of its front edge. Onitis feae Felsche, 1907 (Figs. 4A—-E, 13) Onitis feae Feischea 1907: 293. Material examined.—P. R. CHINA. Yun- nan: Menghai, 1 6, 1,300 m, 28 February 1957, coll. Da-Hua LIU; Jingdong, 1 @, 1,300 m, 6 March 1957, coll. Panfilov. VI- ETNAM. Hanoi, 4 6, 6 2, 20 m, 23 De- cember 2000, coll. Wen-Zhu LI. All speci- mens are deposited in the Institute of Zo- ology, Chinese Academy of Sciences, Bei- jing, P. R. China (IZAS). Distribution.—China (Yunnan), Vietnam, Thailand, Burma, India. Notes.—This is a new record for China. The specimens were collected from cow dung. ACKNOWLEDGMENTS We gratefully acknowledge the assistance of Dr. Ales Bezdek (Institute of Entomol- ogy, Academy of Sciences of the Czech Re- public), Dr. Giuseppe M. Carpaneto (Univ- ersita degli Studi Roma Tre, Italy), Dr. Bert Kohlmann (Universidad Earth, Costa Rica); and Dr. Patrick Prevost (France) for obtain- ing literature. Dr. Denis Keith (Muséum d’Histoire Naturelle et de Préhistoire, France), Dr. Andrew B. T. Smith (Canadian Museum of Nature, Canada), Dr. Dirk Ah- rens (Deutsches Entomologisches Institut, Germany), Dr. Si-Qin Ge (IZAS), Dr. Li-Jie Zhang (IZAS), Mr. Wen-Zhu Li (IZAS), and Mr. Shu-Yong Wang (IZAS) provided useful advice. The project was supported by a Chinese National Science Foundation Grant (No. 30200025). LITERATURE CITED Fabricius, J. C. 1798. Supplementum Entomologiae Systematicae. Hafniae, 572 pp. VOLUME 108, NUMBER 2 Felsche, C. 1907. Coprophage Scarabaeiden. Deutsche Entomologische Zeitschrift 1907: 293 Hanboonsong, Y. and K. Masumoto. 2000. Dung bee- tles (Coleoptera, Scarabaeidae) of Thailand, part 2, Genus Onitis. Elytra 28(1): 101-114. Harold, E. von. 1871. Diagnosen neuer Coprophagen. Coleopterologische Hefte 7: 114. Janssens, A. 1937. Revision des Onitides. Mémoires Musée Royal d’Histoire Naturelle de Belgique 11(2): 1-200, pls.1—2. Lansberge, G. 1875. Monographie des Onitides. An- nales Société Entomologique Belgique 18: 1—148. Philips, T. K., E. Pretorius, and C. H. Scholtz. 2004. A phylogenetic analysis of dung beetles (Scara- beinae: Scarabaeidae): unrolling an evolutionary history. Invertebrate Systematics 18: 53-88. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 396-404 DELPHACODES CAMPESTRIS (VAN DUZEE) AND D. LUTULENTA (VAN DUZEE) (HEMIPTERA: FULGOROMORPHA: DELPHACIDAE): ASSOCIATION WITH COMMON HAIRGRASS, DESCHAMPSIA FLEXUOSA (POACEAE), AND NOTES ON HABITATS, SEASONALITY, AND TAXONOMY A. G. WHEELER, JR. AND CHARLES R. BARTLETT (AGW) Department of Entomology, Soils, and Plant Sciences, Clemson University, Clemson, SC 29634-0315, U.S.A. (e-mail: awhlr@clemson.edu); (CRB) Department of Entomology and Wildlife Ecology, 250 Townsend Hall, University of Delaware, Newark, DE 19716-2160, U.S.A. (e-mail: 02542 @udel.edu) Abstract.—Common hairgrass, Deschampsia flexuosa (L.) Trin. (Poaceae: Pooideae), is recorded as a new host for the widespread Nearctic planthoppers Delphacodes campestris (Van Duzee) and D. lutulenta (Van Duzee). We found D. lutulenta in Maryland, New York, Pennsylvania, Virginia, and West Virginia at 10 sites, ranging from disturbed areas such as roadside slopes to more natural communities such as a mid-Appalachian shale barren and high-elevation rock outcrops. In North Carolina, South Carolina, and Tennes- see, the delphacid associated with common hairgrass was D. campestris, which we col- lected at six sites in the southern Appalachians and upper Piedmont; it was found in grassy balds, on a high-elevation granitic dome and high-elevation granitic outcrop, and on a low-elevation granitic dome. In addition to characterizing their habitats, we give notes on the seasonality and taxonomy of both planthoppers. Key Words: Insecta, common hairgrass, grass feeding, southern Appalachians, grassy balds, granitic outcrops, shale barrens Delphacodes campestris (Van Duzee) METHODS and D. lutulenta (Van Duzee) are common, widely distributed Nearctic planthoppers generally found in meadows and pastures (e.g., Spooner 1920; Osborn and Drake 1922; Osborn 1938, 1939). Unlike most North American Delphacidae, including the majority of other Delphacodes species (Bal- lou et al. 1987, Calvert et al. 1987), their Common hairgrass, D. flexuosa, was sampled (by AGW) for planthoppers in Maryland, New York, Pennsylvania, Vir- ginia, and West Virginia from late July to early August 2002, and in North Carolina, South Carolina, and Tennessee from 2002 to 2005. Plants were tapped over a white host plants (grasses; Poaceae) are reason- ably well known, and both species have been studied in the field and laboratory (e.g., Giri and Freytag 1983a, b; Wilson et al. 1993). Here we give new state records; record common hairgrass, Deschampsia flexuosa (L.) Trin. (Poaceae: Pooideae), as a new host; and provide taxonomic and bi- ological notes. enamel tray with an ax handle, and a plastic vial was used to collect dislodged nymphs and adults from the tray. Notes on season- ality are based mainly on collections and observations of D. campestris at irregular intervals at Glassy Mountain, Pickens County, South Carolina, from March 2002 to March 2005. Nymphal instars, when sorted in the field, are indicated by roman VOLUME 108, NUMBER 2 numerals in the Seasonality and Material examined sections; otherwise, nymphs are referred to generally as early, mid-, or late instars. Brachypterous and macropterous adults are denoted by ‘“‘b”’ and ‘‘m,”’ re- spectively; asterisks indicate new state rec- ords. Records of D. campestris from Roan Mountain in Carter County, Tennessee, are listed under Mitchell County, North Caro- lina. Determinations of the Delphacodes species were made by CRB, and voucher specimens were deposited in the University of Delaware Insect Reference Collection, Newark, Delaware, and the Clemson Uni- versity Arthropod Collection, Clemson, South Carolina. We compiled distribution records from the literature, citing the first reference that lists a particular state. Older records, espe- cially those prior to R. H. Beamer’s work, may be suspect. For each species, we also give a critical synonymy and taxonomic notes. The list of synonymy includes the original combination and synonyms since publication of Metcalf’s (1943) catalog of Delphacidae, which includes an exhaustive synonymy up to 1940. Morphological ter- minology follows Asche (1985). Delphacodes Fieber As currently defined, the genus Delpha- codes Fieber, 1866 (Delphacinae: Delpha- cini; type species Delphax (Delphacodes) mulsanti Fieber, 1866), consists of 10 Old World species (Wagner 1963, Asche and Remane 1983, Asche 1985), with all New World Delphacodes in incertae sedis. Ap- proximately 95 species of Delphacodes are found in North America, but comprehen- sive keys to species are lacking. Delpha- codes species can be definitively identified only by features of the male genitalia. Del- phacodes campestris and D. lutulenta (Figs. 1-2) are similar in size and shape of the head; size, shape, and dentition of the hind tibial spur; and in lacking processes on seg- ment 10, and the posteriorly projecting di- aphragm armature. They are most easily recognized using the works by DuBose 397 (1960) or Wilson and McPherson (1980a). They often can be distinguished in the field by color (D. campestris is paler), but ex- amination of male genitalia is needed to confirm their identity. Delphacodes cam- pestris has broader parameres in caudal view and teeth on the aedeagus (lacking in D. lutulenta). Delphacodes campestris and D. lutulenta are similar in having no pro- cesses on segment 10 and the armature of the genital diaphragm posteriorly projecting (Figs. 5-10). Both species fall within the same clade among the more advanced Del- phacini in preliminary phylogenetic analy- ses using combined molecular and morpho- logical data (Cryan and Bartlett, unpub- lished). Delphacodes campestris (Van Duzee) (Figs, 13; 5-7) Liburnia campestris Van Duzee 1897: 254. Liburnia osborni Van Duzee 1897: 250, synonymy by Oman 1947: 210. Liburnia unda Metcalf 1923: 207, synony- my by Metcalf 1949: 56. Van Duzee (1897) described D. campes- tris from Michigan, Mississippi, New Hampshire, New York, and Ontario. Libur- nia osborni (Van Duzee 1897) and L. unda (Metcalf 1923), considered synonyms of D. campestris, were described from Iowa, Michigan, New Jersey, and New York, and from North Carolina, respectively. Delpha- codes campestris also is known from Ari- zona, California, Colorado, Connecticut, District of Columbia, Kansas, Louisiana, Massachusetts, Nevada, Pennsylvania (Met- calf 1943): Illinois, Minnesota, Missouri, Nebraska, Ohio (DuBose 1960); South Da- kota (Stoner and Gustin 1980); Kentucky (Giri and Freytag 1983a); and North Da- kota, Oklahoma, Tennessee, and Virginia (Wilson 1992). Canadian records are avail- able for Alberta, Yukon (Wilson 1992); British Columbia, Manitoba, New Bruns- wick, Newfoundland, Nova Scotia, Ontario, Prince Edward Island, Quebec, and Sas- 398 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Adult male habitus and distribution. 1, 3, Delphacodes campestris. 2, 4, Delphacodes lutulenta. On maps, shaded states are published records; hatched states are new records. Delphacodes lutulenta also is known from Puerto Rico. katchewan (Maw et al. 2000). In addition to these previously published records and those reported herein, CRB has observed specimens in collections from Delaware, Idaho, Maryland, Montana, Washington, and Wyoming (Fig. 3). Data on the bionomics of D. campestris include the collection of nymphs and adults in Nebraska on eight irrigated, cool-season grasses of the subfamily Pooideae. Densi- ties varied seasonally on different hosts and were highest on intermediate wheatgrass (Thinopyrum intermedium (Host) Barks- worth and D.R. Dewey). The delphacid produced two generations in the study plots; nymphs of a partial third generation Overwintered (Whitmore et al. 1981). In Kentucky, nymphs and adults were collect- ed from plots of mixed grasses (chloridoid, panicoid, and pooid species) from June to November. The incidence of macroptery was 31.3% in 1980 and 42.5% in 1981 (Giri et al. 1985). When tested on seven grasses in the laboratory, D. campestris colonized four species: Johnson grass (Sorghum hal- epense (L.) Pers.), oats (Avena sativa L.), tall fescue (Lolium arundinaceum (Schreb.) S. J. Darbyshire), and wheat (Triticum aes- tivum L.) (Giri and Freytag 1983a). Wilson et al. (1993) collected adults from a tall- grass prairie in Missouri. In the laboratory, using a colony initiated from eggs depos- ited by females collected in South Dakota, D. campestris reproduced successfully (ovi- position plus nymphal development) on 39 native and introduced C, and C, grasses of several subfamilies. Cereals such as barley (Hordeum vulgare L.), oats, rye (Secale cereale L.), and wheat allowed a rapid buildup of populations for maintaining stock colonies. In the laboratory at 25°C, total development time (egg to adult) av- VOLUME 108, NUMBER 2 eraged 31 days for males and 35 days for females. Longevity of mated females ranged from 18: to 55 days; fecundity ranged from 213 to 919 eggs (Stoner and Gustin 1980). Habitats.—In the southern Appalachians of North Carolina (and in Tennessee on Roan Mountain), we found D. campestris on common hairgrass in grassy balds on Black Balsam Knob (ca. 1,735 m) and Roan Mountain (ca. 1,685 m); in a high-elevation granitic outcrop (ca. 1,125 m) in Alleghany County; and in a red oak (Quercus rubra L.) forest on Whiteside Mountain (ca. 1,490 m), a high-elevation granitic dome. We found the planthopper in South Carolina in the Mountains (Blue Ridge) ecoregion near the overlook (ca. 975 m) at Caesars Head State Park, a high-elevation granitic out- crop, and in the Piedmont ecoregion on Glassy Mountain (ca. 515 m), a monadnock that is a low-elevation granitic dome. Seasonality—On Glassy Mountain in South Carolina, D. campestris overwintered mainly as late instars. Nymphs were more numerous than newly developed adults on 10 March 2002, but by 16 March only adults (brachypters) were found. In 2003, a brachypterous male was observed with late instars on 9 March. In 2005, the first 20 individuals observed during sampling on 25 March included 18 brachypterous adults (4 3, 14 2) and two fifth instars. Nymphs (in- stars I-V) of a new generation were present with smaller numbers of brachypters on 12 May 2002; late instars and brachypterous and macropterous adults were observed on 29 May. The planthopper was more difficult to find in mid- to late June 2002, with late instars and adults present on 15 June and only adults on 22 June. Nymphs (early to late instars) of a second generation were found on 8 July when only one adult was observed; a late-July collection consisted of small numbers (<10) of early and late in- stars and a brachypterous female. Sampling on 17—18 August did not yield nymphs or adults. On 21 August, fewer than five early instars and a brachypterous female were ob- 399 served on Whiteside Mountain in western North Carolina. Taxonomic notes.—Delphacodes cam- pestris (Fig. 1) is light tan, sometimes with an orangish abdomen, with dark brown spots on the front coxae and lateral com- partments of the mesonotum. Delphacodes campestris is most similar morphologically to D. atralabris Beamer, 1948a, which 1s smaller and lacks the dark markings on the mesonotum, and D. lutea Beamer, 1946 and D. penelutea Beamer, 1948b, which lack dark markings. All four species differ in the shape of the parameres and aedeagus but have similarly shaped diaphragm armatures and lack processes on segment 10 (Beamer 1946, 1947, 1948a, b; Wilson & McPherson 1980a). Material examined.—NORTH CARO- LINA: Alleghany Co., ca. 10 km E of Lau- rel Springs, 16 June 2002, 3beG, 3b 2; 4 Aug. 2002, 5b 6, 5b 2; Haywood Co., Black Balsam Knob, 18 July 2002, 14 b d, sm, 6b 2; fim 2a 298Sepie 20027 3b 6; Jackson Co., Whiteside Mountain, ca. 6 km NE of Highlands, 21 Aug. 2002, 1b 9; Mitchell Co., Roan Mountain, Carvers Gap, 27 Oct: 2002, lam; 6, Tbr SSilateimstars-23il Aug. 2003, adults, nymphs; 7 Nov. 2004, 3b 6, 1m dé, II-V. *SOUTH CAROLINA: Greenville Co., overlook, Caesars Head State Park, 6 Nov. 2004, IV—V; Pickens Co., Glassy Mountain, 4.2 km NE of Pick- ens, 10 Mar. 2002, brachypters, III—V; 16 Mar. 2002, brachypters; 12 May 2002, bra- chypters, macropters, I-V; 29 May 2002, brachypters, macropters, late instars; 15 June: 2002, 3bid ; 2b, 2. Noam, 9 a Vz 22 June: 2002. 2bad.,, 2timid 2 bee aaa S July 2002, 1b 3, early-late instars; 27 July 2002, 1b 2, 2 early, 3 late instars; 9 Mar. 2003, 1b 3, 9 mid-late instars; 13 Apr. 2003, 1b 3; 6 Nov. 2004, III-V; 25 Mar. 2005, 4b ¢.,. 14b. 2, 2V. TENNESSEE: See NORTH CAROLINA: Mitchell Co. Delphacodes lutulenta (Van Duzee) (Figs. 2, 4, 8-10) Liburnia lutulenta Van Duzee 1897: 252. 400 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 10 Figs. 5-10. Male genitalia. 5-7, Delphacodes campestris. 8-10, D. lutulenta. 5, 8 = pygofer, caudal view. 6, 9 = pygofer, lateral view. 7, 10 = aedeagal complex, lateral view. Scale bar = 0.2 mm. Described from New York (Van Duzee 1897), D. lutulenta has been recorded from 15 additional states: California, Colorado, Connecticut, Kansas, Massachusetts, Min- nesota, New Hampshire, Ohio (Metcalf 1943); North Carolina, South Dakota (DuBose 1960); Illinois (Wilson and Mc- Pherson 1980b); Kentucky (Sperka and Freytag 1975); Maryland (Bean and Denno 1980; the study site was not listed but was in Maryland [R. A. Bean, personal com- munication]); Missouri (Wilson et al. 1993); and Tennessee (Bartlett and Bow- man 2003). Canadian records include Al- berta, British Columbia, Nova Scotia, On- tario, Quebec, and Saskatchewan (Maw et al. 2000). It also is known from Puerto Rico (Osborn 1929, 1935). In addition to these records, and those reported herein, CRB has observed specimens in collections from Ar- izona, Delaware, Indiana, Maine, and Ver- mont (Fig. 4). In Kentucky, D. lutulenta was trivoltine and the most abundant delphacid in mixed- grass pastures (Giri and Freytag 1983a, Giri et al. 1985). Under controlled conditions, the planthopper developed on 13 grasses in the subfamilies Panicoideae and Pooideae; wheat was the preferred host. At 26°C, de- velopment times for males (33.4 days) and females (34.2 days) were not statistically different. Mean fecundity was 430.5 eggs, VOLUME 108, NUMBER 2 and longevity was 27 days for males and 34 days for females (Giri and Freytag 1983b). Other host grasses are Ischaemum muticum L. (as Andropogon repens (Roxb.) Steud.) and Puccinellia nuttalliana (Schul- tes) A. Hitchce. (Wilson et al. 1994). Habitats.—We collected D. lutulenta in New York in a pitch pine-oak-heath rocky summit of the northern Shawangunk Moun- tains (ca. 365 m), in oak-hickory and mixed deciduous forests in Pennsylvania, a shale barren in Maryland, and at the edge of woodlands and along shaded roadside banks in Maryland and West Virginia. The Virginia collections were from high-eleva- tion (ca. 1230 m) rock outcrops. Seasonality.—Collections were nearly limited to late July 2002. Late instars were found with brachypters at all sites in Mary- land, Pennsylvania, Virginia, and West Vir- ginia. No macropters were observed during 21-23 July. An early-August collection in Virginia included an early instar with bra- chypterous adults. Taxonomic notes.—Delphacodes lutulen- ta (Fig. 2) is brownish tan, usually with a darker abdomen. It is most similar morpho- logically to D. lutulentoides Beamer, 1948a and D. perusta Beamer, 1947. Delphacodes lutulentoides differs mainly in having teeth on the aedeagus and more spatulate para- meres (Beamer 1948a). Delphacodes perus- ta has short processes on segment 10, each with a mediobasal tooth, and parameres that are broad and laterally directed basally, and that narrow and are medially directed be- yond midlength (Beamer 1947). Delpha- codes perusta was found in this study on Danthonia sericea Nuttall (Poaceae) in Pickens Co., South Carolina. Material examined.—MARYLAND: Washington Co., Rt. 144, 5.5 km W of Han- cock, 22 July 2002, 4b 6, 7b 2, 2V; Wood- mont Rd. 2.5 km S of Exline, 22 July 2002, 2b 6, 3b 2, IV— V; Boy Scout shale bar- ren, NE of Little Orleans, 22 July 2002, 12b 3, 10b 2, III-V. NEW YORK: Ulster Co., Mohonk Preserve, Bonticou Crag, 4 Sept. 20025b 6) 2ba2) *PENNSYEVANIA: 401 Franklin Co., Warren Township SW of Syl- van, lower slope Keefer Mountain, N of T301 (Red Rock Rd.), 23 July 2002, 2b 6, 5b 2, IV—V; Franklin Co., Warren Town- ship, SW of Sylvan, N of Rt. 456 E of jet. T301 (Red Rock Rd.), 23 July 2002, 8b 6, 5b 2, IV—V. *VIRGINIA: Bath Co., Rt. 703, Bald Knob, 3.5 km N of Alleghany Co. line, 3 Aug. 2002, 16b 3, 3b &, 1 early instar; Madison Co., Hawksbill Mountain, 2ivduly 2002;°9b-6, 9b. 23 IV—-V.. +WEST VIRGINIA: Morgan Co., Nebo Rd. E of Rt. 9, ca. 9 km S of Great Cacapon, 22 July 2002, libeé')2be 2s, latesimstars- RE Oca 3 km N of Largent, 22 July 2002, 2b 6, 1V. DISCUSSION Delphacodes campestris and D. lutulenta are multivoltine grass feeders (Whitmore et al. 1981, Giri et al. 1985). Under controlled conditions in the greenhouse or laboratory, they can reproduce on many of the same hosts, mainly pooid grasses (Stoner and Gustine 1980; Whitmore et al. 1981; Giri and Freytag 1983a, b). The planthoppers have a similar distribution—essentially transcontinental in Canada (Maw et al. 2000), south to the southeastern states, and west to California (Metcalf 1943; DuBose 1960)—and sometimes co-occur (e.g., Spooner 1920, Osborn and Drake 1922, Giri et al. 1985, Wilson et al. 1993). They also both serve as hosts of the dryinid par- asitoid Dicondylus americanus (Perkins) (Giri et al. 1985, Giri and Freytag 1988). Both planthopper species might also be grass generalists (sensu Whitcomb et al. 1988), although information on their host- plant range is based mainly on the screen- ing of cultivated grasses in the greenhouse or laboratory, or on studies of grasses in single- or mixed-species field plots. Little is known about the wild grasses that serve as hosts of these delphacids. In the eastern United States, we found nymphs and adults of D. campestris and D. lutulenta on common hairgrass. Several Pa- learctic delphacids are known from this Holarctic grass (e.g., Holzinger et al. 2003, 402 Nickel 2003), but Deschampsia flexuosa apparently is a new host for Nearctic plant- hoppers. Delphacodes campestris and D. lutulenta, though found on common hair- grass from New York to South Carolina, were never syntopic. Delphacodes lutulenta was associated with common hairgrass at more northern sites (New York to northern Virginia), whereas D. campestris was found on the grass at southern sites (North Caro- lina, South Carolina, and southern Tennes- see). The north-south disparity in host use was not based solely on differences in ele- vation; D. lutulenta in Virginia was found at higher elevations than was D. campestris at certain sites in North Carolina, South Carolina, and Tennessee. The ecological factors influencing the planthoppers’ use of common hairgrass in different geographic areas are unknown and might be an artifact of our limited number of sample sites. Van Duzee (1897) noted that D. campestris is numerous in dry pastures, whereas D. lu- tulenta is found on grass in damp situations. Phytophagous insects typically feed on a particular host plant in only part of the plant’s range (e.g., Strong et al. 1984), as in Delphacidae associated with the cord- grass Spartina patens (Ait.) Muhl. in east- ern North America (Denno et al. 1981). Certain European delphacids (assuming cryptic or sibling species are not involved) show geographic variation in host use, some species having reduced diet breadth in northern portions of their range (Nickel 2003). An interpretation of the observed patterns of host use in D. campestris and D. lutulenta must await experimental field and laboratory studies, and perhaps information on the genetic basis of their host-plant ad- aptation. Either or both might represent a complex of sibling species. ACKNOWLEDGMENTS We thank Stephen Bennett (South Caro- lina Department of Natural Resources) for issuing a collecting permit for Glassy Mountain Heritage Preserve; Patrick Mc- Millan (Clemson University) for identifying PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Deschampsia flexuosa and providing distri- butional information; Paul Huth and John Thompson (Mohonk Preserve) and Larry Klotz (Shippensburg University) for taking A.G.W. to D. flexuosa colonies in New York and Pennsylvania, respectively; Wil- liam Brown (University of Delaware) for constructing the species range maps; An- thony Gonzon (University of Delaware) for assisting with specimen processing, identi- fication, and digital photography; and Peter Adler (Clemson University) for suggesting improvements to an earlier draft of the manuscript. LITERATURE CITED Asche, M. 1985. Zur Phylogenie der Delphacidae Leach, 1815 (Homoptera: Cicadina: Fulgoromor- pha). Marburger Entomologische Publikationen 2(1): 1-910. Asche, M. and R. Remane. 1983. Zur Problematik von 1866) Kenntnis einiger benachbarter Taxa (Homoptera Delphacodes mulsanti (Fieber und zur Auchenorrhyncha Fulgoromorpha Delphacidae) (Vorlaufige Mitteilung). Marburger Entomologis- che Publikationen 1(8): 25—5S6. Ballou, J. K., J. H. Tsai, and S. W. Wilson. 1987. Del- phacid planthoppers Sogatella kolophon and Del- phacodes idonea (Homoptera: Delphacidae): de- scriptions of immature stages and notes on biol- ogy. Annals of the Entomological Society of America 80: 312-319. Bartlett, C. R. and J. L. Bowman. 2003. Preliminary inventory of the planthoppers (Hemiptera: Ful- goroidea) of the Great Smoky Mountains National Park, North Carolina and Tennessee, U.S.A. En- tomological News 114: 246-254. Beamer, R. H. 1946. Some new species of Delphaco- des (Homoptera, Fulgoridae, Delphacinae). Jour- nal of the Kansas Entomological Society 19: 139— 144. Beamer, R. H. 1947. 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Field studies of delphacid planthopper populations (Homoptera: Delphacidae), with notes on their dryinid parasites (Hymenoptera: Dryinidae). Jour- nal of the Kansas Entomological Society 58: 69— 74. Holzinger, W. E., I. Kammerlander, and H. Nickel. 2003. The Auchenorrhyncha of Central Europe, Vol. 1: Fulgoromorpha, Cicadomorpha excl. Ci- cadellidae. Brill, Leiden, The Netherlands, 673 pp. Maw, H. E. L., R. G. Foottit, K. G. A. Hamilton, and G. G. E. Scudder. 2000. Checklist of the Hemip- tera of Canada and Alaska. NRC Research Press, Ottawa, 220 pp. Metcalf, Z. P. 1923. A key to the Fulgoridae of eastern North America with descriptions of new species. Journal of the Elisha Mitchell Scientific Society 38: 139-230, plates 38-70. . 1943. General Catalogue of the Hemiptera. Fascicle [TV Fulgoroidea, Part 3 Araeopidae (Del- phacidae). Smith College, Northampton, Massa- chusetts, 552 pp. . 1949. The redescription of twenty-one species of Areopidae described in 1923. 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An ecological study of the Hemiptera of the Cranberry Lake Region, New York. New York State College of Forestry at Syracuse University Technical Publication 16: 5-86. Sperka, C. and P. H. Freytag. 1975. Auchenorrhynchus [sic] hosts of mermithid nematodes in Kentucky. Transactions of the Kentucky Academy of Science 36: 57-62. Spooner, C. S. 1920. Some notes on the occurrence of Delphacinae (Hemip. Homop.). Entomological News 31: 44—46. Stoner, W. N. and R. D. Gustin. 1980. Biology of Del- phacodes campestris (Homoptera: Delphacidae). Annals of the Entomological Society of America 73: 372-374. Strong, D. R., J. H. Lawton, and Sir R. [T-R.E.] South- wood. 1984. Insects on Plants: Community Pat- terns and Mechanisms. Harvard University Press, Cambridge, Massachusetts, 313 pp. Van Duzee, E. P. 1897. A preliminary review of the North American Delphacidae. Bulletin of the Buf- falo Society of Natural Sciences 5: 225-261. Wagner, W. 1963 [1962]. Dynamische Taxionomie, an- gewandt auf die Delphaciden Mitteleuropas. Mit- teilungen aus dem Hamburgischen Zoologischen Museum und Institut 60: 111-189. Whitcomb, R. F, A. L. Hicks, D. E. Lynn, H. D. Blocker, and J. P. Kramer. 1988. Host specificity: a major mechanism enhancing insect diversity in grasslands, Paper 11.06. /n Davis, A. and G. Stan- ford, eds. The Prairie: Roots of our Culture; Foun- dation of our Economy. Proceedings of the Tenth North American Woman’s University, Denton, Texas, June 22—26, 1986. Native Prairie Association of Texas, Dallas. Whitmore, R. W., K. P. Pruess, and J. T. Nichols. 1981. Leafhopper and planthopper populations on eight Prairie Conference of Texas 404 irrigated grasses grown for livestock forage. En- vironmental Entomology 10: 114-118. Wilson, S. W. 1992. The Delphacidae of Yukon Ter- ritory, Canada (Homoptera: Fulgoroidea). Insecta Mundi 6: 79-100. Wilson, S. W. and J. E. McPherson. 1980a. Keys to the planthoppers, or Fulgoroidea, of Illinois (Ho- moptera). Transactions of the Illinois State Acad- emy of Science 73(2): 1-61. . 1980b. The distribution of the Fulgoroidea of the eastern United States (Homoptera). Transac- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tions of the Illinois State Academy of Science 73(4):7—20. Wilson, S. W., C. Mitter, R. E Denno, and M. R. Wil- son. 1994. Evolutionary patterns of host plant use by delphacid planthoppers and their relatives, pp. 7-113. In Denno, R. E and T. J. Perfect, eds. Planthoppers: Their Ecology and Management. Chapman & Hall, New York. Wilson, S. W., J. L. Smith, and P. D. Calvert. 1993. Planthoppers of a Missouri tallgrass prairie (Ho- moptera: Fulgoroidea). Journal of the Kansas En- tomological Society 66: 75-80. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 405-410 REDESCRIPTION OF NEALIOLUS CURCULIONIS (FITCH) (HYMENOPTERA: BRACONIDAE), WITH A NEW HOST RECORD AND DISTRIBUTION DATA VICTOR LOPEZ-MARTIiNEZ, JOSE ANTONIO SANCHEZ-GARCIA, ANTONIO HUERTA-PANIOGUA, GLORIA CALYECAC-CORTERO, NESTOR BAUTISTA-MARTINEZ, AND JOSE ISAAC FIGUEROA-DE LA ROSA (VL-M) Universidad Aut6noma del Estado de Morelos, Facultad de Ciencias Agro- pecuarias, Av. Universidad 1001, Col. Chamilpa, Cuernavaca, Morelos 62210, México (e-mail: vilomar@uaem.mx); (JAS-G) CIIDIR-IPN-Unidad Oaxaca, Area de Control Biolé6gico, Santa Cruz Xoxocotlan, Oaxaca, C.P. 71230, México: (AH-P, GC-C, NB-M, JIF-D) Colegio de Postgraduados, IFIT-Entomologia, Km 35.5 carr. México-Texcoco, Montecillo, Edo. de México, C. P. 56230, México Abstract.—A detailed description of the adult of Nealiolus curculionis (Fitch), with illustrations of diagnostic characters including the male genitalia, is provided. Trichobaris championi Barber (Coleoptera: Curculionidae), a pest of Physalis ixocarpa Brot., is re- ported as a new host record for N. curculionis. New distribution data in Mexico and the first records from Guatemala and Honduras are presented. Key Words: Nealiolus curculionis (Fitch) is a member of the tribe Brachistini of the subfamily Helconinae. It has the forewing vein r-m absent, the first three metasonal tergites forming a carapace with metasomal terga I and II not fused, and the lateral tergite of the second tergum fused with the third. The genus Nealiolus can be identified as an member of the subfamily Helconinae using the keys of March et al. (19987), Shaw (1995), and Sharkey (1997). In México, N. curculionis has been re- ported as a parasite of Anthonomus grandis Boheman (Coleoptera: Curculionidea), a pest of Gossypium hirsutum L. in Guerrero (Cross and Chesnut 1971). In the United Staes, N. curculionis is rocognized as the principal natural enemy of Cylindrocoptu- rus adspersus (LeCOnte) (Coleoptera: Cur- culionidae) (Charlet 1983b), and it is asso- ciated at least with ten curculionid species Helconinae, Brachistini, Neotropical, husk tomato, stalk borer in Canada, México, and the United States (Charlet 1983a, 1994; Charlet et al. 1992; Marsh 1979). Specimens collected in the Mexican State of Puebla revealed a wasp species found emerging from Trichobaris championi Bar- ber pupae on stems and branches from husk tomato, Physalis ixocarpa Brot. Study of these specimens revealed that they are Nealiolus curculionis. This represents a new host record for N. curculionis. This species is redescribed as a necessary part of a forthcoming paper of the Mexican Braconidae fauna (L6pez, in preparation). Furthermore, the male genitalia of N. cur- culionis is described for the first time, and we describe the potential of this structure as morphological character in future system- atic studies of the genus. We also present new data about its distribution in three Cen- tral American countries. 406 MATERIALS AND METHODS Specimens were reared from husk tomato containing 7. championi pupae as part of a bioecological pests program through the Universidad Autonoma del Estado de Mo- relos, Colegio de Postgraduados and CON- ACYT, in central Mexico. Other specimens were borrowed from the following collec- tions: Canadian National Collection, Otta- wa, John. T. Huber (CNC) and Museum of Zoology, Lund University, Sweden, Roy Danielsson (MZUL). The Mexican material is housed in the Coleccion Entomoldgica del Instituto de Fitosanidad (CEAM) and the Facultad de Ciencias Agropecuarias, Universidad Aut6noma del Estado de Mo- relos (VLM). Measurements were performed using an image analyzer Image Pro plus version 3.1 (Media Cybernetics 1997) adapted to a vid- eo camera (Hitachi KP-D51) and a micro- scope (Olympus BX- 50). Terminology fol- lows Sharkey and Wharton (1997) and Martin (1956). We examined male genitalia of 15 specimens from different localities. Preparation technique and terminology of genitalia follows Sdanchez-Garcfa et al. (2003). RESULTS Nealiolus curculionis (Fitch) (Figs. 1-9) Female.—Body length: 3.87 mm. Color: Body black, except following: legs brown, hind tibia dark brown basally; antenna dark brown; ovipositor and clypeus brown; me- tasoma dark brown; mandible brown; api- cally dark brown. Head: Quadrate in front view (Fig. 2); setae sparse on face, clypeus and mandi- bles; vertex temple and gena evenly setose; face surface punctate, vertex, temple and gena polished, fine rugae below antennal fosae; frons polished with shallow and scarce sculptured depressions, but with a strong short carina; face 2.66 times higher than wide; clypeus transverse, 2.31 times PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON wider than high, ventral margin strongly rounded, and with an obtuse angulation near base of each mandible; mandible base width 0.12 mm; eye 1.61 times higher than wide; ocelli partially surrounded by an im- pressed groove; antenna shorter than body, length 3.43 mm, with 27 flagellomeres, 1r- regular placodes present in flagellomeres 1— X, first flagellomere shorted than second (Fig. 3): Mesosoma: \.31 times longer than high; lateral face of pronotum strongly sculptured on medial surface; notaulus deeply im- pressed, strongly foveolate and converging in a foveolate area (Fig. 1); scutellar sulcus deeply impressed, polished and with a dis- tinct midridge; posterior margin of scutel- lum with long foveola; sternaulus deeply impressed (Fig. 5), epicnemial carina with finely rugulose striae; metapleuron densely and uniformly rugose with short whitish se- tae; propodeum rugose, with a distinct pen- tagonal areola, inner surface punctate; lat- eral protuberances almost absent, propodeal spiracle raised at surface level; propodeal carina short. Forewing 3 times longer than high (Fig. 4); stigma broad 3.94 times lon- ger than high; 1Cub 3.8 times longer than 1Cua; 1M distinctly bowed; (RS+M)a sin- uate; im-cu 1.8 times longer than (Rs+M)a sinuate; Im-cu 1.8 times longer than (rs+M)b, 2cu-a present; Rla 4.4 times lon- ger than RIb, the latter not extending to wing tip. Hind wing 4.15 times higher than wide; M+CU 3.13 times longer than IM. Hind femur short, broad, 4.47 times longer than maximum width; hind tibia 6.2 times longer than basal width; tarsonmeres H—IV 1.58 times longer than first tarsomere; tarsal claws with distinct basal tooth (Fig. 7). Metasoma: Carapace oval, short in dor- sal view (Fig. 1), length 1.56 mm; first transverse groove well developed and mov- able, second groove as a fine transverse line; first terguite with centro-lateral carinae strongly developed, reaching first groove, first tergite with a strong lateral lamella, ex- tending from point of articulation with tho- rax to first transverse groove (Fig. |); third 407 VOLUME 108, NUMBER 2 res. 4, Fron is. 1, Dorsal view. 2, lion tal view of head. 3, Basal flagellome a. 6, End view of carapace. 7, Posterior 408 tergite with a very narrow, delicate, lateral lamella (Fig. 6); first two tergites longitu- dinally striate, second tergite with a pol- ished and unsculptured medial area; striae on both first and second tergites not reach- ing transversal grooves (Figs. 1, 6, 8). Ovi- positor length 2.78 mm. Male genitalia: Male specimens with a simple pattern: Aedeagus bifid with two apical lobes, a medial deep canal, and api- cal pits; gonoforceps wider, its apical part rounded and with some lobes, with many straight apical setae and some at its median part; digitus with three or four teeth; cuspis pointed, lamina volsellaris with six setae (Fig. 9). Variation.—Female length 3.79 mm (3.87-—3.96), male 3.78 mm (3.58—3.99). Striae on lateral pronotum variable in num- ber and size, many specimens with more longitudinal striae on basal margin. One male specimen with apex of carapace with a short distance concavity medio-ventrally. Hosts.—Trichobaris championi associat- ed with husk tomato (P. ixocarpa) in Mex- ico 1s a new host record for N. curculionis. Other host associations are Anthonomus grandis Boheman (in cotton squares) (Cur- culionidae), Coccotorus scutellaris (Le- Conte) (Curculionidea), Conotrachelus nenuphar (Herbst) (Curculionidae), C. near tibialis Schoof (Curculionidae), C. posti- catus? Boheman (curculionidae), Crapon- ius inaequalis (Say) (Curculionidae), Cylin- drocopturus adspersus (LeConte) (Curcu- lionidae), Grapholita molesta (Busck) (Tor- tricidae), Magdalis armicollis Say (in elm) (Curculionidae), and Smicronyx fulvus LeConte (in sunflower) (Curculionidae) (Carlet 1983a, b, 1994; Charlet et al. 1992; Marsh 1979; Martin 1956; Pinkham and Oseto 1987). New distribution data.—Previously cited from Canada, United States, and México. We record it from Guatemala and Hondu- ras. This species is also quite common in Costa Rica (S. Shaw, unpublished data). GUATEMALA: | @, Zacapa, below San Lorenzo, XI/1986, 750 msnm, M. Sharkey PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (CNC). HONDURAS: 2 @2, Atlantida, Lan- cetilla.. Tela, 31/VIN/1995s leg.» RaGave: Malaise trap in lowland ran forest, 15°43’N 87°27'W (MZLU); 1) SayYoro, Palowde Comba, 27/1[X/1995, leg. R. Cave, Malaise trap in mid-elevation secondary forest, 15°11'N, 87°39’W (MZLU); 1 6, same data but 29/IX/1995; 3 3d and 2 2, same data but 5/VIII/1995; 1 2, same data but Yus- caran, 15°43’N. 87°27'W. MEXICO: 2 9 and 2 6, Puebla, Tecamachalco, km 22 carr. Carretera México-Veracruz, 2-VIII-2000, Néstor Bautista, Tomate de cascara Physalis ixocarpa Brot., hosp. Trichobaris cham- pioni (CEAM); 1 @, Puebla, Quecholac, Tuzuapan, 16-VI-2001, A, Huerta, tomate de cascara, hosp. Trichobaris championi (CEAM); 2 2 and 1 6, same data but, 9- VI-2001. A. Huerta, Tomate de cascara Physalis ixocarpa Brot., hosp. Trichobaris championi (VLM); 2 2, same data but, 16- VI-2001 (CEAM); | 2, Tuzuapan, 24-VIII- 2001, Antonio Huerta emerged from pupa of Trichobaris championi (VLM). Diagnosis.—This species is quite similar to Neoliolus rufus (Riley), but N. rufus has a rufo-testaceous color. Nealiolus curcu- lionis 1s a dark brown species with the clyp- eus transverse, third tergite finely striate, oval shaped and a slightly curved carapace. DISCUSSION Trichobaris championi is a pest of husk tomato and is distributed in at least ten Mexican states (Barber 1935, Bautista and Morales 2000). Larvae of T. championi feed mainly on internal tissue of the stem and principal branches as a borer, while the mature larvae weave a cocoon with host fi- bers. This species hibernates as a pupa or adult in the husk tomato stems, and reaches 92% infestation levels in Tenango del Valle, Estado de México (Bautisa and Morales 2000) and 15.3% to 87.3% in the Puebla high plateau (Huerta et al. 2003). Our col- lections were made in commercial plots of husk tomato, where 10 insecticide applica- tions are the regular program for T. cham- VOLUME 108, NUMBER 2 409 Figs. 8-9. Nealiolus curculionis. 8, Lateral view of metasoma. 9, Male genitalia. pioni control. From our collections only two braconids species were sampled, WN. curculionis and an undetermined species of Bracon. Nealiolus curculionis represented 80% of the parasitoids reared from this host. The number of collected adults was low due to insecticide applications, but fu- ture collections in untreated fields may give a better idea of the impact of parasitism. Nealiolus curculionis is an endoparasi- toid of some curculionid species. The fe- male oviposits into early first instar larvae and exhibits a great synchronization with its host (Charlet 1994). The use of male genitalia for separation of species in Braconidae has been discussed by Sanchez-Garcia et al. (2003). Male gen- italia can provide useful characters for the higher classification for Braconidae, using setal patterns, dimensions of various struc- tures, form of the gonoforceps, aedeagus, number and position of setae, and number of teeth on the digitus. It is a tool useful for separating species of Braconidae in several 410 genera including Aphaereta Foerster. Bla- cus Nees, Digonogastra Viereck, Ephedrus Haliday, Epsilogaster Whitfield and Mason, Leiophron Nees, Triaspis Haliday, and many genera of Microgastrinae. In the ma- terial used in this study, only one pattern of male genitalia was discovered for N. cur- culionis from the different localities ana- lyzed. If this character is present in other species on the genus, it could provide a tool useful for separating species of Nealiolus. ACKNOWLEDGMENTS Thanks to all collection managers listed in the Materials and Methods for loan of specimens, and to Robert W. Brooks, (Snow Entomological Collections, University of Kansas), Larry D. Charlet (Northern Crop Science Library, USDA), and Colin Favret (Illinois Natural History Survey) for loan of determined material of Nealiolis curculion- is. Thanks to Charles O’Brien (Florida A&M University) for the corroboration of curculionid pest material. J. Romero N. re- viewed the first draft of this paper. Thanks to two anonymous reviewers for helpful comments. This paper was funded by SNI- CONACYT (31351) and PIFI 2004-18-12 grants to the first author. LITERATURE CITED Barber, H. S. 1935. The tobacco and solanum weevils of the genus Trichobaris. United States Depart- ment of Agriculture, Miscellaneous Publication 226, 28 pp. Bautisa, M. N. and O. G. Morales. 2000. Melanagro- myza tomaterae Steyskal (Diptera: Agromyzidae) plaga del tomate (Physalis ixocarpa Brot.) en México. Folia Entomolégica Mexicana 110; 129— 130. Charlet, L. D. 1983a. Inset stem fauna of native sun- flower species in western North Dakota. Environ- mental Entomology 12: 1286-1288. . 1983b. Parasitoids of a stem weevil, Cylin- drocopturus adspersus(Coleoptera: Curculioni- dae) in sunflower: Incidence and parasitization in the northern Great Plains. Environmental Ento- mology 12: 888-890. . 1994. Seasonal abundance and impact of the sunflower stem weevil parasitoid, Nealiolus cur- culionis (Hymenoptera: Braconidae), in the north- ern Great Plains. Biological Control 4:26-31. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Charlet. L. D., G. L. Brewer, and V. H. Beregovoy. 1992. Insect fauna of the heads and stems of na- tive sunflowers (Asterales: Asteraceae) in eastern North Dakota. Environmental Entomology 21: 493-500. Cross, W. H. and T. L. Chesnut. 1971. Arthropod par- asites of the boll weevil Anthonomus grandis 1. An annotated list. Annuals of the Entomological Society of America 64(2); 516-527. Huerta P, R. A., N. Bautista M., and J. Romero N. 2003. Present status of Trichobaris championi Barber (Coleoptera: Curculionidae) in Mexico. Southwestern Entomologist 29(2): 153— 154. Marsh, P. M. 1979. Family Braconidae, pp. 144-313. In Krombein, K. V., P. D. Hurd, D. R. Smith, and B. D. Burks, eds. Catalog of Hymenoptera in America North of Mexico, Vol. 1. Symphyta and Apocrita (Parasitica). Smithsonian Institution Press, Washington, D.C. pp. 1— 1198. Marsh, P. M., S. R. Shaw, and R. A. Wharton. 1987. An identification manual for the North American genera of the Family Braconidae (Hymenoptera). Memoirs of the Entomological Society of Wash- ington No. 13, 98 pp. Martin, J. C. 1956. A taxonomic revision of the trias- pidine braconid wasps of Nearctic America (Hy- menoptera). Canadian Department of Agriculture Publication 965:1—157. Media Cybernetics. 1997. Image-pro plus reference guide, version 3.1 for Windows, Media Cybernet- ics, Silver Spring, Maryland. Pinkham, J. D. and C. Y. Oseto. 1987. Natural enemies and their maturity rates on Smicronyx fulvus LeConte (Coleoptera: Curculionidae) larvae. En- vironmental Entomology 16: 1302-1304. Sanchez-Garcia, J. A., R. A. Wharton, J. Romero-Na- poles, A. G6nzalez- Hernandez, V. L6pez-Marti- nez, A. Equihus-Martinez, H. Gonzalez-Hernan- dez, and J. L. Carrillo-Sanchez. 2003. Description of a new species of Blacus Nees (Hymenoptera: Braconidae) from Mexico, utilizing characters of the male external genitalia. Pan-Pacific Entomol- ogist 79(2): 135-144. Sharkey, M. J. 1997. Subfamily Key, pp. 39-64; Hel- coninae, pp. 261—274. In Wharton, R. A., P. M. Marsh, and M. J. Sharkey, eds. Manual of the New World Genera of the Family Braconidae (Hymenoptera). International Society of Hymen- opterists. Special Publication No. 1, 439 pp. Sharkey, M. J. and R. A. Wharton. 1997. Morphology and Terminology, pp. 19-37. In Wharton, R. A., P. M. Marsh, and M. J. Sharkey, eds. Manual of the New World Genera of the Family Braconidae (Hymenoptera). International Society of Hymen- opterists. Special Publication No. 1, 439 pp. Shaw, S. R. 1995. Braconidae. Chapter 12.2, pp. 431— 463. In Hanson, P. E. and I. Gauld, eds. The Hy- menoptera of Costa Rica, Oxford University Press, Oxford, 893 pp. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 411-417 FEROSETA PRISCUS (NEUROPTERA: MANTISPIDAE), A NEW GENUS AND SPECIES OF MANTIDFLIES IN DOMINICAN AMBER GEORGE POINAR, JR. Department of Zoology, Oregon State University, Corvallis, OR 97331, U.S.A. (e-mail: poinarg @science.oregonstate.edu) Abstract.—Feroseta priscus (Neuroptera: Mantispidae), a new genus and new species of mantidflies, is described from Dominican amber. Diagnostic characters of the new genus are: |, The cubitus of the hindwing is parallel to the first anal vein and does not touch it, 2, the absence of a pterostigma in all wings; 3, the shape and pedicellated nature of the flagellomeres; 4, the extended tip of the terminal flagellomere; 5, the structure of the thorax; and 6, the elongated male ectoprocts. Key Words: mantidfly, Tertiary Mantidflies of the family Mantispidae (Neuroptera) are a cosmopolitan group of lacewings with a poor fossil record. The family is divided into four or five subfam- ilies depending on whether the South Af- rican Rhachiberothidae is included with the more traditional groups (Redborg 1998). The Neotropical mantispids fall into four subfamilies, the Calomantispinae, Sym- phrasinae, Drepanicinae, and the Mantispi- nae. These can be separated by the structure of the forelegs, with members of the former three subfamilies having two claws and an aroleum on the foretarsi, and the Mantis- pinae with only a single claw and no aro- leum on the fore tarsi (Penny 1982a, b; Hoffman 2002). A recently discovered male mantidfly in Dominican amber could not be placed in any of the extant genera and is described below in a new genus in the subfamily Mantispinae. MATERIALS AND METHODS The amber piece was repolished to obtain a better view of the diagnostic characters. Feroseta priscus, new genus, new species, Mantispidae, Dominican amber, The final amber piece weighs 4.9 g, is 32 mm in greatest length, 21 mm in greatest width, and 13 mm in greatest depth. The specimen was obtained from the La Toca mine in the Cordillera Septentrional of the Dominican Republic. Dating of Dominican amber is still controversial with the latest purposed age of 20-15 mya based on fo- raminifera (Iturralde-Vinent and MacPhee 1996) and the earliest as 45—30 mya based on coccoliths (Cépek in Schlee 1999). Do- minican amber is secondarily deposited in sedimentary rocks, which makes a definite age determination difficult (Poinar and Mastalerz 2000). A range of ages for Do- minican amber may be likely since the am- ber is associated with turbiditic sandstones of the Upper Eocene to Lower Miocene Mamey Group (Draper et al. 1994). Do- minican amber was produced by the legu- minous tree, Hymenaea protera Poinar (1991). A reconstruction of the Dominican amber forest based on amber fossils indi- cated that the environment was similar to that of a present day tropical moist forest (Poinar and Poinar 1999). Terminology is 412 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON based on the works of Penny (1982a, b) and Lambkin (1986a, b). Observations, drawings, and photographs were made with a Nikon SMZ-10 R stereo- scopic microscope and Nikon Optiphot compound microscope with magnifications up to 600X. Family Mantispidae Leach, 1815 Subfamily Mantispinae Leach, 1815 The presence of only a single claw and absence of an aroleum in the foreleg place the fossil in the subfamily Mantispinae (Penny 1982a, b; Lambkin 1982a, b). The fossil is well preserved and complete except for the absence of the foretarsus of the right leg. Feroseta Poinar, new genus Type species.—Feroseta priscus Poinar. Description.—Body medium size; with a single claw without aroleum on foretarsus; antennal flagellomeres pedicellated; termi- nal flagellomere with extended tip; prono- tum with few scattered setae in prozonal re- gion, mesoscutal furrow inconspicuous; cu- bitus of hind wing runs parallel to anal vein without touching it; pterostigma not scler- otized in all wings, replaced by bristles; ec- toprocts elongated and narrow, with well- developed median lobes; mediuncus with extended terminus. Diagnosis.—The combination of the above characters separates Feroseta from all other genera in this subfamily, including those recently described from Costa Rica (Hoffman 2002). Etymology.—‘‘Feroseta”’ is from the Latin “‘fero”’ for to bear or carry, and ‘seta’ for bristle in regards the bristles in the area of the pterostigma. The gender is feminine. Feroseta priscus Poinar, new species (Figs, 1-10) Description—wWith characters listed un- der generic description. Total length 14.2 mm (as positioned in the amber) but 15.9 mm when body divisions measured sepa- rately. Head: Length, 1.6 mm, width, 2.1 mm; occiput only slightly raised above level of compound eyes; distance between eyes, 2.1 mm; upper portion of head (vertex and frons) dark, lower portion (clypeus and mouthparts) light; antennal length, 2.8 mm; antenna with 30 segments; distance be- tween antennal bases, 440 wm; flagello- meres ranging from as wide as long to twice as wide as long; segments appearing glabrous, but with few, fine, short setae; fla- gellomeres pedicellated (with pedicel or ex- tension at base); terminal 6 flagellomeres darker than remainder, terminal flagello- mere with extended tip; head region be- tween eyes brown, lower portion of head light, without inverted V-shaped marking or vertical or horizontal stripes above or be- neath antennae. Thorax: Brown; length prothorax, 4.1 mm; length mesothorax, 1.3 mm; length metathorax, 1.3 mm; prothorax glabrous except for some scattered hairs in prozonal region; dorsal surface of pronotum bearing flattened transverse rugae; apical portion of pronotum just posterior to prozonal region with dorsal constriction; prozonal region of pronotum saddle-shaped; mesoscutal fur- row inconspicuous; wings clear, no infus- cated area at apex; trichosores absent; setae on all wing veins; lacking subcosta cross- veins; pterostigma absent in all wings, re- placed by macrosetae; forewing length, 12.0 mm; greatest width, 3.5 mm; second anal vein not forked; with strongly recurved humeral crossvein; two radial cells origi- nating from first radial cell (r,), three radial cells originating from second radial cell (r,); ewo radial cells originating from third radial cell (r;); hindwing length, 10.4 mm; greatest width, 3.2 mm; 3 radial cells orig- inating from first radial cell (r,), two radial cells originating from second radial cell (1) (in one wing, second radial cell divided by a separate vein) (Fig. 7), two radial cells originating from third radial cell (r;); 7—8 graduate crossveins; legs mostly light in VOLUME 108, NUMBER 2 3. Feroseta priscus in Dominican amber. 1. Holotype male. Scale bar = 3.9 mm. 2, Head and prothorax. Note dorsal pronotal constriction (arrow). Scale bar — trichia (arrow) in place of pterostigma. Scale bar = Bisse l= 1.3 mm. 3, Tip of forewing showing macro- 850 jm. Insert shows claws on left hind leg. The smallest claw does not show on this photo. Scale bar = 512 pm. 414 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON +8 Figs. 4-5. Feroseta priscus in Dominican amber. 4, Head showing darkened 6 terminal antennal segments. Scale bar = 550 pm. 5, Femur, tibia and tarsus of foreleg showing major and 2 of 3 minor spines (arrows). Scale bar = 440 wm. color; forecoxa divided into 2 parts; fore- ments, length segments, 1, 435 ym; 2, 86 femur dark, length, 4.0 mm; bearing one pm; 3, 97 wm; 4, 97 wm; 5, 183m; length large central spine (major spine of Lambkin — foreleg claw, 69 jm; arolium absent; length 1986a) (length, 718 jm), 3 minor spines’ mid- and hind leg claws, 154 wm; claws on and 7—8 small spines; foretarsus with 5 seg- mid- and hind legs 4 cleft. VOLUME 108, NUMBER 2 Abdomen: Coloration uniformly light, without mottling; ectoprocts elongate, nar- row, bearing long setae, with well-devel- oped median lobes bearing short, thick se- tae on inner surfaces; length of narrow ter- minal portion of ectoproct, 324 jm; width of narrow terminal portion of ectoproct, 94 j.m; mediuncus with extended terminus. Material examined.—Holotype male in Dominican amber, deposited in the Poinar amber collection maintained at Oregon State University (accession # N-4-16). Etymology.—‘“‘Priscus” is Latin for an- cient, of former times. Diagnosis.—The flattened transverse ru- gae and the constriction on the pronotum are characteristics of several mantispid gen- era, including the genus Entanoneura En- derlein 1910. However, in Entanoneura the pterostigma is chitinized and darkened, there are numerous branches of the radius in both wings and the cubitus bends down and either touches or is fused for a short distance with the anal vein (Enderlein 1910). DISCUSSION The American Mantispinae range from North America to Argentina, with some 119 species in 15 genera reported from South and Central America (Rehn 1939; Welch and Kondratieff 1991; Penny 1977, 1982b; Handschin 1960; Ohl 2004). Lambkin (1986a) stated, “‘the presence of a pteros- tigma is one of the characteristic features of the family”’. Thus, the absence of a scler- otized pterostigma in F. priscus is surpris- ing. Consideration was given to the possi- bility that the portion of the wing bearing the pterostigma had somehow cleared dur- ing the fossilization process. However, oth- er mantispid fossils in amber (see below) possess well-developed pterostigmae and even if some clearing had occurred in the present fossil, one would expect that at least one wing would have had a partial pteros- tigma. However, all wings are equally de- void of both pigmentation and sclerotiza- 415 tion and the region of the pterostigmae con- tains macrosetae. Another unusual character on the fossil is the character and positioning of the fla- gellomeres. In most mantispids, the flagel- lomeres are quite pubescent and closely ap- pressed to one another, however in the fos- sil, the flagellomeres are sparsely hirsute and separated (Fig. 4). Separation of adja- cent flagellomeres is caused by an exten- sion (pedicel) at the base of each flagello- mere. The shape of the terminal flagello- mere in each antenna suggests that the ter- minal and subterminal segments might have fused (Fig. 4). The shape of the male terminalia is used in the systematic placement of the mantis- pids. As far as could be determined, the ec- toprocts of F. priscus are more elongated and pointed than those of any other Amer- ican or even Australian species of Mantis- pinae (Fig. 10) (Enderlein 1910; Penny 1982b; Lambkin 1986a,b). The wing venation is known to be vari- able in this group and this is apparent by the presence of an extra radial cell in one of the hind wings, caused by a vein sepa- rating the second radial cell into two equal cells (Fig. 7). Also, in one forewing, the tip of the RI vein is represented by a row of macrotrichia. Fossil mantidflies are rare. Whalfera ven- atrix (Whalley 1983) of the subfamily Pla- tymantispinae in British amber (quite likely to be an extended deposit of Eocene Baltic amber and roughly the same age) is fairly small (6 mm in length), lacks a recurrent humeral vein in the forewing, has forelegs with only 4 tarsal segments and possesses wing trichosors. All of these characters sep- arate it from F. priscus. A second fossil, Vectispa relicta (Cockerell), was described from a wing fragment and a complete left forewing in Oligocene sedimentary depos- its from the Isle of Wight (Cockerell 1921; Jarzembowski 1980). The presence of a dis- tinct pterostigma separates this fossil, which was placed in the Mantispinae, from F. priscus. Another mantispid in Dominican 416 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 6-10. 1S Feroseta priscus in Dominican amber. 6, Forewing. Scale bar = 1 mm. 7, Hind wing. Note two r, radial cells. Scale bar = 1 mm. 8, Fore tarsus. Scale bar = 114 wm. 9, Anterior portion of prothorax. Scale bar = 512 pm. 10, Dorsal view of terminalia showing elongated lobed ectoprocts and terminal mediuncus. Scale bar = 432 wm. hv = humeral vein; r, = first radial cell; r, = second radial cell; r; = third radial cell; 1A = first anal vein; CuP = cubitus vein; Sc = subcostal vein. amber, which was tentatively assigned to the genus Mantispa, was mentioned and figured in Poinar and Poinar (1999). This specimen, which is in a private collection in Europe, has a well defined pterostigma in both wings, which separates it from F. priscus. Prosagittalata oligocenica Nel and Climaciella henrotayi Nel were described from Oligocene deposits in France. The presence of well-developed pterostigmae in both wings and the wing venation separate these fossils from F. priscus. Other fossil VOLUME 108, NUMBER 2 mantispids listed by Ohl (2004) have wing features that are quite different from F. priscus. As a family, mantispids develop on a wide range of insect larvae in the orders Hymenoptera, Lepidoptera, and Coleoptera. However as far as known, all members of the subfamily Mantispinae, which includes F. priscus, develop only on spider eggs (Redborg 1998). There would have been a wide host selection in the Dominican amber forest since presently some 152 species of spiders in 82 genera and 36 families are known from these deposits (Wunderlich 2004). ACKNOWLEDGMENTS I thank Norm Penny and Roberta Poinar for comments on earlier drafts of this man- uscript and Norm Penny for bringing the author’s attention to certain references. LITERATURE CITED Cockerell, T. D. A. 1921. Fossil arthropods in the Brit- ish Museum. VI. Oligocene insects from Gurnet Bay, Isle of Wight. Annals and Magazine of Nat- ural History 7: 453—480. Draper G., P. Mann, and J. EK Lewis. 1994. Hispaniola. pp. 129-150. In Donovan, S. and T. A. Jackson, eds. Caribbean Geology: An Introduction, The University of the West Indies Publishers’ Associ- ation, Kingston. Enderlein, G. 1910. Klassifikation der Mantispiden nach dem Material des Stettiner Zoologischen Museums. Stettiner Entomologische Zeitung 71: 341-379. Handschin, E. 1960. Zur Revision siid-amerikanischer Mantispiden. Revue Suisse de Zoology 67: 523- 558. Hoffman, K. 2002. Family Mantispidae. Proceedings of the California Academy of Sciences 53: 161— 457. Iturralde-Vinent M. A. and R. D. E. MacPhee. 1996. Age and paleogeographic origin of Dominican amber. Science 273: 1850-1852. Jarzembowski, E. A. 1980. Fossil insects from the Bembridge Marls, Paleocene, of the Isle of Wight, 417 Southern England. Bulletin of the British Museum of Natural History (Geology) 33: 237-293. Lambkin, K. L. 1986a. A revision of the Australian Mantispidae (Insecta: Neuroptera) with a contri- bution to the classification of the family 1. Gen- eral and Drepanicinae. Australian Journal of Zo- ology, Supplementary Series, 116: 1-142. . 1986b. A revision of the Australian Mantis- pidae (Insecta: Neuroptera) with a contribution to the classification of the family 2. Calomantispinae and Mantispinae. Australian Journal of Zoology, Supplementary Series 117: 1-113. Nel, A. 1988. Deux nouveaux Mantispidae (Planipen- nia) fossils de l’?Oligocene du sud- est de la France. Neuroptera International 5: 103-109. Ohl, M. 2004. Annotated catalog of the Mantispidae of the world (Neuroptera). Contributions on En- tomology, International 5: 131—262. Penny, N. D. 1977. Lista de Megaloptera, Neuroptera e Raphidioptera do México, América Central, [llas Caraibos e América do Sul. Acta Amazonica (sup- plemento) 7: 1-61. . 1982a. Review of the generic level classifi- cation of New World Mantispidae (Neuroptera). Acta Amazonica 12: 209-223. . 1982b. Neuroptera of the Amazon Basin. Part 6. Mantispidae. Acta Amazonica 12: 415—463. Poinar, G. O., Jr. 1991. Hymenaea protera sp. n. (Le- guminosae: Caesalpinoideae) from Dominican amber has African affinities. Experientia 47: 1075-1082. Poinar, G. O., Jr. and R. Poinar. 1999. The Amber For- est. Princeton University Press, Princeton, New Jersey. Poinar, G. O., Jr. and M. Mastalerz. 2000. Taphonomy of fossilized resins: determining the biostratinomy of amber. Acta Geologica Hispanica 35: 171-182. Redborg, K. E. 1998. Biology of the Mantispidae. An- nual Review of Entomology 43: 175-194. Rehn, J. W. H. 1939. Studies in North American Man- tispidae (Neuroptera). Transactions of the Ameri- can Entomological Society 65: 237-263. Schlee, D. 1990. Das Bernstein-Kabinett. Stuttgart Beitrage der Naturkunde (C), No. 28, 100 pp. Welch, J. L. and B. C. Kondratieff. 1991. The Mantis- pidae (Neuroptera) of Colorado. Journal of the Kansas Entomological Society 64: 69-76. Whalley, P. 1983. Fera venatrix gen. and sp.n. (Neu- roptera, Mantispidae) from amber in Britain. Neu- roptera International 2: 229—233. Wunderlich, J. 2004. Fossil spiders in amber and copal. Beitrage zur Araneologie 3A,B. 1908 pp. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 418-428 A STUDY OF THE GENUS RHODOBATES RAGONOT (LEPIDOPTERA: TINEIDAE) FROM CHINA HOUHUN LI AND YUNLI XIAO College of Life Sciences, Nankai University, Tianjin, 300071, P. R. China (e-mail: lihouhun @ nankai.edu.cn) Abstract.—We treat seven species of Rhodobates Ragonot from China, of which five are described as new: Rhodobates bifurcatus Li and Xiao, n. sp., Rhodobates triangu- latus Li and Xiao, n. sp., Rhodobates fascespinatus Li and Xiao, n. sp., Rhodobates curvativus Li and Xiao, n. sp., and Rhodobates cupulatus Li and Xiao, n. sp. The female of Rhodobates sinensis Petersen is reported for the first time. Photographs of the adults and illustrations of wing venation, male 8th abdominal segments, and genital struc- tures of all the species except Rhodobates amorphopa (Meyrick) are provided, along with a key to the species. Key Words: Rhodobates Ragonot was proposed in 1895, with Euplocamus laevigatellus Her- rich-Schaffer as its type species. Petersen (1965) reviewed the genus and added six new species from the Palaearctic Region; Gozmany and Vari (1973) described two species from the Afrotropical Region; Pe- tersen (1987) revised the genus, providing a discussion of its phylogenetic relation- ship, and added three new species and one new subspecies; and Robinson and Tuck (1996) synonymized the monotypic genus Phaulogenes Meyrick and transferred its type species to Rhodobates Ragonot. To date, the genus comprises 12 named species and subspecies. Rhodobates is distributed mainly in the Af- rotropical, Palaearctic, and Oriental regions. In China, two species previously were re- corded, R. amorphopa (Meyrick) and R. si- nensis Petersen. Here we describe five new species from Yunnan, Sichuan, Guangdong, and Tianjin, China. All the specimens ex- amined, including the types, are deposited in the Insect Collection, College of Life Scienc- es, Nankai University, Tianjin, China. Lepidoptera, Tineidae, Rhodobates, systematics, new species, China Rhodobates Ragonot, 1895 Rhodobates Ragonot 1895: civ. Type spe- cies: Euplocamus laevigatellus Herrich- Schaffer 1854, by monotypy. Paraplutella Rebel 1901: 163. Type spe- cies: Paraplutella algiricella Rebel, 1900, by monotypy. Chliarostoma Meyrick 1913: 335. Type species: Chliarostoma relecta Meyrick, 1913, by monotypy. Phaulogenes Meyrick 1938: 27. Type spe- cies: Phaulogenes amorphopa Meyrick, 1938, by monotypy. Tineodoxa Amsel 1955: 32. Type species: Myrmecozela tibulella Rebel, 1936, by original designation. Description.—Medium to large sized. Head densely covered with erected narrow lamellar scales. Maxillary palpus five-seg- mented, folded. Labial palpus recurved; second segment with or without sparse bris- tles on outer side, ventral side with scale brush protruded forward and downwards in most species. Antenna about 2/3 length of VOLUME 108, NUMBER 2 forewing, without pecten; each flagellar segment in male basally covered with one annulus of broadly fanlike scales, female with two annuli of narrow lamellar scales; cilia in male about 0.5 times diameter of flagellomere, shorter in female. Forewing dark brown to dark ocherous, R, and R; stalked or separate. Hind wing with M, and M, short stalked or separated. Male genitalia: Uncus more or less con- cave on caudal margin; ganthos with arms joined distally by sclerotized structure; val- vae connected by a complex of transtilla, anellus, and juxta; sacculus folded inwards; anellus usually with lateral side fused with base of valva, ventrally open. Aedeagus usually slender, curved; vesica covered with spinules; cornutus present or absent. Female genitalia: Posterior apophyses about twice length of anterior apophyses. Ostium funnel-shaped in general; ductus bursae elongate, slender; corpus bursae pyr- iform, signum absent. Distribution.—China, Lebanon, Jordan, Iran, Iraq, Turkey, Russia, Spain, Italy, Mal- ta, Greece, Egypt, Morocco, Libya, Algeria, Tunisia, Tanzania, South Africa, and several islands of the Mediterranean Sea. Remarks.—Rhodobates is closest to Cimitra Walker (Hapsiferinae) in appear- ance. It is characterized by the head with erect narrow lamellar scales, gnathos joined by one sclerotized structure, and valvae connected by a specialized complex of tran- stilla, anellus, and juxta. Petersen (1987) transferred Rhodobates from the Myrme- cozelinae to Hapsiferinae, but Robinson and Nielsen (1993) disagreed with this placement and referred it back to Myrme- cozelinae, which we follow in this paper. KEY TO THE CHINESE SPECIES OF RHODOBATES [Because R. amorphopa (Meyrick) is known only from the female, it is not included in the key. ] 1. Gnathos connected by a slender band ...... 2 — Gnathos connected by a small weakly sclero- tized triangular plate (Fig. 23) 419 R. cupulatus Li and Xiao, n. sp. 2. Band connecting gnathos without sclerotized OSES . 5..c oe ORO NNCNONONO o ONOMMONS fos o es oso. 3 — Band connecting gnathos posteriorly with two strongly sclerotized triangular processes (Fig. ZO). So cde R. triangulatus Li and Xiao, n. sp. 3. Sacculus with spines close to apex ........ + = SAcCulusmwATDOUtESpPINES) leila eee =) 4. Valvae strongly curved inwards at about distal ven (eas 2/2) . R. curvativus Li and Xiao, n. sp. SV Alwacmotctiievied (1070210) saan eran een ae Rey cae cote R. fascespinatus Li and Xiao, n. sp. 5. Uncus deeply concave at middle of caudal mar- gin (Fig. 19)... R. bifurcatus Li and Xiao, n. sp. — Uncus with caudal margin almost straight (Fig. DAY Tee ee eta etaie, ats R. sinensis Petersen Rhodobates bifurcatus Li and Xiao, new species (Bigsy ise75, 135 195) 25) Types.—Holotype d, China: Mt. Wei- bao, Weishan County (25°14'N, 100°18’E), Yunnan Province, 2,200 m, 21 vii 2001, leg. Houhun Li & Xinpu Wang, genitalia slide No. XYLO4150. Paratypes: 46 ¢, 15 2, same data as holotype. Diagnosis.—The new species is allied to R. sinensis, but it can be separated from the latter by the presence of two small yellow- ish-white spots on the forewing, one at about % of fold and the second at the distal end of cell, uncus deeply concave at middle of caudal margin, ventral valva with two subtriangular protuberances at % and %4, and the juxta anteriorly bifurcate. In R. sinensis, the forewing is uniformly dark grey, the un- cus is almost straight in caudal margin, the valva ventrally lacks protuberance, and the juxta is anteriorly rounded. Description (Figs. 1, 7).—Wingspan 18.0—25.0 mm. Head dark ocherous. Labial palpus dark ocherous; second segment with O—S5 bristles in outer side, scale brush with inside dark ocherous, outside dark brown. Thorax and tegula dark brown. Forewing and cilia dark brown, with small yellowish- white spot at % of fold and distal end of cell; R, connate with R,;. Hind wing and cilia dark grey, all veins separate. Male 8th abdominal segment (Fig. 13): Tergum somewhat triangular, rounded cau- 420 Figs. 1-6. 5, R. cupulatus. 6, R. sinensis. dally; sternum laterally sclerotized, gradu- ally narrowed toward caudal margin; ante- rior margin heavily sclerotized. Male genitalia (Fig. 19): Uncus with 2— 3 toothlike spines along lateral margin, deeply concave at middle of caudal margin, both ends slightly curved inwards; concave margin each with small heavily sclerotized toothlike spine at ¥;. Gnathos joined in dis- tal 4 by one narrow V-shaped band, curved at 1%; distal % quite sclerotized, ventrally with one small toothlike spine near apex; apex pointed. Valvae with apex somewhat truncate, costa slightly concave at about 4% and '2, ventral margin with subtriangular protuberance at 2 and %4; transtilla extended PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Adults of Rhodobates spp. 1, R. bifurcatus. 2, R. triangulatus. 3, R. fascespinatus. 4, R. curvativus. from lateral side of anellus to % of valvae, its distal 7, expanded toward costa in irreg- ularly triangular shape. Anellus laterally fused with base of costa. Juxta long, nar- row, anteriorly bifurcate, slightly concave at middle posteriorly. Sacculus folded in- wards, extended backward to about % of valvae. Saccus small, somewhat triangular, rounded apically. Aedeagus strongly curved near apex; ductus ejaculatorius about % length of aedeagus, with two slender scler- otized bands inside; cornutus absent. Female genitalia (Fig. 25): Ostium long funnel-shaped, caudal margin concave, broad V-shaped. Ductus bursae with trans- versal folds in posterior 4%. VOLUME 108, NUMBER 2 Figs. 7-12. R. curvativus. 11, R. cupulatus. 12, R. sinensis. Etymology.—The specific name is de- rived from the Latin bifurcatus = bifurcate, in reference to the anteriorly bifurcate juxta. Rhodobates triangulatus Li and Xiao, new species (Bigs. 2,78; 14, 203,26) Types.—Holotype d¢, China: Zhubalong, Batang County (30°00'N, 99°09'E), Sich- 421 Wing venation of Rhodobates spp. 7, R. bifurcatus. 8, R. triangulatus. 9, R. fascespinatus. 10, uan Province, 2,500 m, 10 vii 2001, leg. Houhun Li & Xinpu Wang, genitalia slide No. XYL04152. Paratypes: 2 5, 1 2, same data as holotype; 4 2°, Xiangcheng County (28°55'N, 99°47'E), Sichuan Province, 2,800 mm, 13 vii 2001, leg. Houhun Li & Xinpu Wang. Diagnosis.—The new species resembles R. sinensis, but it is distinguishable by the 422 22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON triangularly concave caudal margin of the uncus, which is almost straight in the latter; the band connecting arms of gnathos bear- ing two small extra sclerotized triangular processes, which are lacking in R. sinensis; and ductus ejaculatorius only about % the length of the aedeagus, but about % length of aedeagus in R. sinensis. Description (Figs. 2, 8).—Wingspan 18.0—23.0 mm. Head dark grey. Labial pal- pus dark ocherous; second segment with O— 3 bristles in outer side, scale brush dark ocherous, sparsely with scattered dark brown scales. Thorax and tegula dark grey. Forewing grey, tinged with dark brown scales; cell with white spot at end; basal distance between R, and R, about 3.0 times that between R, and R,; cilia dark brown. Hind wing and cilia dark grey, all veins separated. Male Sth abdominal segment (Fig. 14): Tergum more or less triangular, distally slightly expanded, caudally blunt; sternum deeply concave at middle, somewhat trian- gular laterally, with anterior margin heavily sclerotized. Male genitalia (Fig. 20): Uncus with caudal margin triangularly concave, both ends slightly curved inwards, somewhat tri- angular; concave margin each with small toothlike spine at %. Gnathos elongate, acutely bent at 4; arms connected by a nar- row V-shaped band at about ¥, the band posteriorly with two extra sclerotized tri- angular processes; distal 7; heavily sclero- tized, pointed at apex. Valvae with apex broad and truncate; costa with basal half slightly sinuate; transtilla extending from lateral side of anellus to center of valva at %, slightly expanded in distal part. Anellus with lateral side fused with base of costa. Juxta narrow from middle to both ends, caudally with a small process at left. Sac- culus slightly folded inwards, extending to about % length of valva; apex obtusely pointed. Saccus small, somewhat bowl- shaped; anterior margin obtusely rounded. Aedeagus curved near apex; ductus ejacu- latorius about % length of aedeagus, with two sclerotized slender bands inside; cor- nutus absent. Female genitalia (Fig. 26): Ostium long funnel-shaped; caudal margin shallowly emarginate, widely V-shaped. Ductus bur- sae with numerous transversal folds in pos- tenor = Etymology.—The specific name is de- rived from the Latin triangulatus = trian- gular, in reference to the band connecting the gnathos posteriorly with two extra scler- otized triangular processes. Rhodobates fascespinatus Li and Xiao, new species (Figs: 3, S5aS; 2 a27)) Types.—Holotype 6, China: Dawuling Natural Reserve, Xinyi County (22°21'N, 110°56’E), Guangdong Province, 1,000 m, 8 viii 2003, leg. Dandan Zhang, genitalia slide No. XYL04176. Paratypes: 2 2, same locality data as holotype, 8~10 viii 2003. Diagnosis.—This new species is similar to R. sinensis, but it can be distinguished by the presence of two small dark brown spots on the forewing, one at % of fold and the other at the distal end of the cell; in R. sinensis the forewing has no spots. In R. fascespinatus, the valvae bear numerous fascicled spines at the apex of the sacculus, the saccus is ill-defined and cornuti consist of six small spines; in R. sinensis, the val- vae lack spines, the saccus is well-devel- oped, and the cornuti are absent. Description (Fig. 3, 9).—Wingspan 15.0—19.0 mm. Head ocherous yellow. La- bial palpus ocherous yellow; second seg- ment without bristle in outer side, scale brush ocherous yellow, outside with scat- tered dark brown scales. Thorax and tegula dark ocherous. Forewing and cilia dark ocherous, scattered with dark brown scales; a small dark brown spot located at % of fold; distal end of cell with a small dim dot; R, and R, short-stalked, basal distance be- tween R, and R, about 5.0 times that be- tween R, and R,,;. Hind wing and cilia grey, all veins separate. Male 8th abdominal segment (Fig. 15): VOLUME 108, NUMBER 2 423 Figs. 13-18. 8th abdominal segments of males of Rhodobates spp. 13, R. bifurcatus. 14, R. triangulatus. 15, R. fascespinatus. 16, R. curvativus. 17, R. cupulatus. 18, R. sinensis. 424 Tergum trapezoidal, lateral side gently arched inwardly; sternum medially con- cave, laterally triangular, anterior margin greatly sclerotized. Male genitalia (Fig. 21): Uncus with caudal margin somewhat straight, median 4% concave, narrowly V-shaped; concave mar- gin distally short spinelike. Gnathos elon- gate, sharply bent at basal 4, with arms joined at distal % by large U-shaped band; distal %4 much sclerotized, ventrally with a small tooth near apex, terminating in a curved tip. Valvae with basal 7 about twice width of distal ¥;, apex broad, truncate; cos- ta convex at 4; transtilla extending from lateral side of juxta to 7% of valva near apex of sacculus, then expanded obliquely out- ward from ¥, to subapex near costal %. Anellus laterally fused with base of valva. Juxta long rectangular, posterior /; greatly sclerotized, bifurcate. Sacculus folded in- wards, with numerous fascicled spines near apex, the spines about /, to 4% times length of valvae; apex reaching about 7 of valvae, broad, somewhat truncate. Saccus poorly defined. Aedeagus bent near apex; ductus ejaculatorius equal in length to aedeagus, with two long and slender sclerotized inte- rior bands; cornuti consisting of six small spines arranged in a line. Female genitalia (Fig. 27): Ostium long funnel-shaped, distally concave, U-shaped. Ductus bursae not delimited from ostium. Etymology.—The specific name is de- rived from the Latin prefix fasce- = fasci- cled, and Latin spinatus = spined, in ref- erence to the sacculus with numerous fas- cicled spines. Rhodobates curvativus Li and Xiao, new species (Bigs. 4. 10s 6122. 23) Types.—Holotype d, China: Mt. Da- dong, Lian County (24°48'N, 112°23’B), Guangdong Province, 650 m, 21 vi 2004, leg. Dandan Zhang, genitalia slide No. XYLO04156. Paratypes: 1 6,5 2, same data as holotype. Diagnosis.—The new species is allied to PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON R. sinensis, but can be differentiated from the latter by the absence of a small tooth at the concave margin of the uncus, which is present in R. sinensis; the cornuti composed of 13-16 small spines arranged in a line, which are absent in R. sinensis; and the shape of the valvae. Description (Figs. 4, 10).—Wingspan 17.0—22.0 mm. Head ocherous yellow. La- bial palpus ocherous yellow; second seg- ment with O—4 bristles on outer side, scale brush with inside ocherous yellow, outside dark brown. Thorax, tegula, forewing, hind wing, and cilia all dark grey; forewing with basal distance between R, and R, about 5.0 times that between R,; and R,; hind wing with all veins separate. Male 8th abdominal segment (Fig. 16): Tergum irregularly trapezoidal, lateral side arched inwardly, posterior margin convex medially; sternum laterally sclerotized, somewhat triangular, deeply concave at middle, anterior margin heavily sclerotized. Male genitalia (Fig. 22): Uncus concave at middle of caudal margin, laterally form- ing two triangular processes. Gnathos with arms linked by medially widened band at about distal %4, curved at about %; distal “4 sclerotized, ventrally with a small toothlike spine near apex, more or less curved dis- tally, ending in point. Valvae very long, wide at base, narrowed to %, greatly bent inwards at about distal 4%; distal % gradually broadening, with apex obtusely rounded; ventral margin with small process at %, which gradually broadens from middle to apex, asymmetrical: left process with apex slightly concave, right process with apex truncate; long digital process at about 7%; transtilla extending from anellus to 7% of valva, broad in distal 7%, somewhat trian- gular. Anellus small, far away from base of valva. Juxta with anterior half somewhat rounded, posterior half somewhat rectan- gular, terminating in two small processes. Sacculus slightly folded inwards, about % as long as valva, with 4—8 medially curved spines close to apex; spines about % length of valvae; apex pointed. Saccus unconspic- VOLUME 108, NUMBER 2 425 Figs. 19-24. Male genitalia of Rhodobates spp. 19, R. bifurcatus. 20, R. triangulatus. 21, R. fascespinatus. 22, R. curvativus. 23, R. cupulatus. 24, R. sinensis. Scale is 0.5 mm except 1.0 mm in Fig. D2 426 uous. Aedeagus sinuate in distal 7; ductus ejaculatorius about equal in length to ae- deagus, with two internal sclerotized elon- gate bands; cornuti composed of 13-16 slightly curved, small spines arranged in a line. Female genitalia (Fig. 28): Ostium with posterior 7, concave V-shaped, anterior ¥, somewhat rectangular. Ductus bursae slightly expanded, sclerotized in distal half. Etymology.—The specific name is de- rived from the Latin curvativus = curved, in reference to strongly curved distal 4% of the valvae. Rhodobates cupulatus Li and Xiao, new species (Figs.'S, 1lel7e223529) Types.—Holotype 6, China: Mt. Jiulong, Ji County (40°0’’N, 117°24’E), Tianjin, 180 m, 10 vi 2004, leg. Houhun L1 et al., gen- italia slide No. XYLO04154. Paratypes: 1 &, same data as holotype; | 9%, Limutai, Ji County, Tianjin, 300 m, 11 vi 2004, leg. Houhun Li et al. Diagnosis.—The new species resembles R. algiricellus (Rebel), but it differs by the following three characters: uncus laterally not folded inward in the new species, con- spicuously folded inward in R. algiricellus; saccus large and broad in the new species, unconspicuous in the latter; aedeagus strongly curved at distal / in R. cupulatus, medially curved in R. algiricellus. Description (Figs. 5, 11).—Wingspan 16.5—24.0 mm. Head dark brown, mixed with grayish white. Labial palpus with in- side grayish white, outside dark brown; sec- ond segment with 3-10 bristles in outer side, scale brush unconspicuous. Thorax, tegula, forewing, hind wing, and cilia all dark brown; forewing with distance be- tween bases of R, and R,; about 2.0 times that between R, and R,; hind wing with all veins separate. Male Sth abdominal segment (Fig. 17): Tergum trapezoid; sternum with posterior margin emarginate, somewhat V-shaped. Male genitalia (Fig. 23): Uncus broad, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with caudal margin concave, broadly V- shaped, forming two triangular processes, caudally with strongly sclerotized tooth at each side. Gnathos relatively short and stout, strongly curved at about 7, some- what U-shaped, arms joined with each oth- er by a small, weakly sclerotized, trian- gular plate; distal %4 sclerotized, curved and pointed. Valvae with apex narrow and obtusely rounded; costa nearly straight; transtilla extending from lateral side of anellus to center of valva “4 from base. Anellus fused with base of costa. Juxta with anterior 7; weakly sclerotized, some- what rectangular; posterior ¥; bifurcate, distally pointed. Sacculus folded inwards, reaching % of valva, obtusely rounded at end. Saccus large and broad, anteriorly rounded. Aedeagus short, curved at distal /;; ductus ejaculatorius about % length of aedeagus; cornutus absent. Female genitalia (Fig. 29): 8th abdomi- nal segment with posterior half of sternum concave. Ostium short, cup-shaped, poste- rior margin slightly convex. Etymology.—The specific name is de- rived from the Latin cupulatus = cup- shaped, in reference to the cup-shaped os- tium. Rhodobates sinensis Petersen, 1987 (Figs. 6, 12; 18; 24, 30) Rhodobates sinensis Petersen 1987: 174. Material examined._4 6, 70 @, Lijiang County (26°52'N, 100°14’E), Yunnan Prov- ince, 2,650 m, 16 vii 2001, leg. Houhun Li and Xinpu Wang. Description (Figs. 6, 12).—Wingspan 19.0-21.0 mm. Male Sth abdominal segment (Fig. 18): As illustrated. Male genitalia (Fig. 24): As illustrated. Female genitalia (Fig. 30): Ostium elon- gate, funnel-shaped; caudal margin slightly concave at middle. Ductus bursae with transversal folds in basal %. Distribution.—China (Yunnan). Remarks.—The female of this species is reported for the first time. Petersen (1987) VOLUME 108, NUMBER 2 Ostium Ductus bursae Corpus bursae 28 Figs. 25-30. 28, R. curvativus. 29, R. cupulatus. 30, R. sinensis. Scale is 1 mm. described the species based on a single male collected by HoGne in 1934. The spec- imens we examined in this study were col- lected from the same locality. Rhodobates amorphopa (Meyrick 1938) Phaulogenes amorphopa Meyrick 1938: DT’ 427 Female genitalia of Rhodobates spp. 25, R. bifurcatus. 26, R. triangulatus. 27, R. fascespinatus. Rhodobates amorphopa (Meyrick): Robin- son and Tuck 1996: 8. Distribution.—China (Yunnan). Remarks.—Meyrick (1938) proposed the genus Phaulogenes based on the female of this species collected from Yunnan. We found no additional specimens of this spe- cies, and we were unable to locate the type. 428 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Based on the original description, this spe- cies can be distinguished from R. bifurcatus which is also described from Yunnan Prov- ince, China, by the forewing pattern and color. ACKNOWLEDGMENTS We truly thank Dr. R. Gaedike (Deutsch- es Entomologisches Institut, Miincheberg) for providing us with helpful literature. The research was supported by the National Natural Science Foundation of China for the Special Program. LITERATURES CITED Amsel, H. 1955. Uber mediterrane Microlepidopteren und einige transcaspische Arten. Bulletin de l'Institut Royal des Sciences Naturelles de Bel- gique Entomologie 31: 1—64. Gozmany, L. A. and L. Vari, 1973. The Tineids of the Ethiopian Region. Transvaal Museum Memoir 18: 1-238. Meyrick, E. 1913. Descriptions of South African Mi- crolepidoptera. Annals of the Transvaal Museum 10: 267-336. . 1938. Microlepidoptera excl. Pyralidae, pp. 1— 29. In Caradja, A. and E. Meyrick, Materialien zu einer Mikrolepidopterenfauna des Ytilingshan- massivs (Provinz Ytinnan). Deutsche Entomolo- gische Zeitschrift “Iris”, herausgegeben vom En- tomologische Verein zu Dresden, Vol. 52. Petersen, G. 1965. Revision der Gattungen Rhodobates Ragonot und Pachyarthra Amsel (Lepidoptera: Tineidae). Beitrage zur Entomologie 15: 87—100. . 1987. Revision der Gattung Rhodobates Ra- gonot (Lepidoptera, Tineidae, Hapsiferinae). En- tomologische Abhandlungen, Staatliches Museum fiir Tierkunde, Dresden 50: 167—190. Robinson, G. S. and E. S. Nielsen, 1993. Tineid Gen- era of Australia (Lepidoptera). CSIRO Publica- tions, Melbourne 344 pp. Robinson, G. S. and K. R. Tuck. 1996. A revisionary checklist of the Tineidae (Lepidoptera) of the Ori- ental Region. Occasional Papers on Systematic Entomology 9: 1—22. Ragonot, E. L. 1895. Microlépidopteres de la Haute- Syrie récoltés par M. Ch. Delagrange et descrip- tions des espéces nouvelles. Bulletin de la Société Entomologique de France 64: 94—109 [Annals de la Société Entomologique de France 64 (appen- dix)]. Paris Rebel, H. 1901 (1900). Neue palaearctische Tineen. Deutsche Entomologische Zeitschrift “‘Iris’’, her- ausgegeben vom Entomologische Verein zu Dres- den 13: 161-188. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 429-442 A REVISED CHECKLIST OF THE STONEFLIES (PLECOPTERA) OF WEST VIRGINIA (USA) DONALD C. TARTER AND CHARLES H. NELSON (DCT) Department of Biological Sciences, 1 John Marshall Drive, Marshall University, Huntington, WV 25755, U.S.A.; (CHN) Department of Biological and Environmental Sciences, 615 McCailie Avenue, The University of Tennessee—Chattanooga, Chattanooga, TN 37403, U.S.A. (e-mail: charles-nelson @utc.edu) Abstract.—Based on the examination of additional collections and published records, the revised checklist of the stoneflies of West Virginia includes 135 species in 42 genera. Eight state records and 399 county records are added to the plecopteran fauna of the state. We have noted range extensions for the following stoneflies: Allocapnia illinoensis Frison; Perlesta cinctipes (Banks); P. frisoni Banks; and P. teaysia Kirchner and Kondratieff. Key Words: Prior to this review, Tarter and Kirchner (1980) reported 106 species of stoneflies comprising 37 genera from West Virginia. Subsequent emendations and taxonomic changes involving seven of these species, Allocapnia indianae Ricker (Stark et al. 1986), Paracapnia opis (Newman) (Stark et al. 1986), Alloperla caudata Frison (Sur- dick 2004), Sweltsa mediana (Banks) (Sur- dick 1985), Acroneuria evoluta Klapalek (Stark and Brown 1991), Perlesta placida (Hagen) (Stark 1989), and Yugus bulbosus (Frison) (Nelson 2001) resulted in their de- letion from the list. Four other species, Tae- niopteryx nivalis (Fitch), Cultus decisus (Walker), Yugus arinus (Frison), and Allo- perla idei (Ricker), the latter first reported by Stewart and Stark (1988), are also de- leted from the list as we have not been able to confirm their presence with the collection of adult material. However, an additional 25 species have been reported in the published literature. Based on the examination of ad- ditional collections, we add eight new state records along with 399 new county records to the plecopteran fauna of West Virginia. Plecoptera, stoneflies, West Virginia, species checklist, drainage basins Thus, a total of 135 species representing 42 genera are recorded for the state. Distribu- tions of stoneflies in the seven major drain- age basins of West Virginia are presented in this paper as well. Range extensions are noted for the following species: Al/locapnia illinoensis Frison (IL, IN, MD, ME, MN, NY, OH, ON, QC, VA, WI) southward; Perlesta cinctipes (Banks) (AR, IA, IL, OH, KS, MO, NE, OK) eastward; P. frisoni Banks (NC, SC, TN, VA) and P. teaysia Kirchner and Kondratieff (MD, PA, VA) westward. Numerous investigators, including Alex- ander and Stewart (1999), Baumann (1974), Claassen (1931), Farmer and Tarter (1976), Frison (1935, 1942), Griffith and Perry (1992), Griffith et al. (1998), Harper and Kirchner (1978), Hissom and Tarter (1976), Illies (1966), Kirchner (1978, 1982), Kirch- ner and Kondratieff (1984, 1985, 1988), Kondratieff and Baumann (2000), Kondra- tieff and Kirchner (1982, 1987, 1988, 1996), Needham and Claassen (1925), Nel- son (2001), Ricker (1949, 1952), Ricker and Ross (1968, 1969, 1975), Ross and 430 Ricker (1964, 1971), Stark (1990), Stark and Gaufin (1976), Stark and Baumann (1978), Stark and Kondratieff (1987, 2004), Stark et al. (1986, 1988), Steele and Tarter (1977), Stewart and Stark (1988, 2002), Stewart et al. (1991), Surdick (1985, 2004), Tarter and Kirchner (1980), Tarter et al. (1975, 1976, 1977), Turner et al. (1996), and Zwick (1973), have reported stonefly records from West Virginia. Janssen (1973) divided the state into seven major drainage basins (Fig. 1). All of the records in this investigation are listed by these seven ma- jor drainage basins and counties (Fig. 2) within the drainage basin. Stonefly speci- mens examined for this study are deposited at the following institutions or agencies: Marshall University (West Virginia Ben- thological Survey), United States Army Corps of Engineers (Huntington District), Compliance Monitoring Lab (Chapmans- ville, WV), West Virginia Department of Agriculture, West Virginia Department of Environmental Protection, and West Virgin- ia University. Collecting data (county, stream, date, collector) are recorded in the West Virginia Benthological Survey at Marshall University. CHECKLIST Below is a checklist of 135 stonefly spe- cies found in West Virginia. For each spe- cies in this checklist the drainage basins are indicated by roman numerals (I—VII), coun- ties are enclosed in parenthesis and the con- tiguous states where they have been report- ed are enclosed by brackets. New county records are indicated with an ‘‘*’’, while new state records are denoted by a ‘‘**’’, We also include an additional 17 species from a group of 58 species found in the immediately adjacent states but not reported from West Virginia (Kondratieff and Bau- mann 2000). These species, indicated by a * + ”’, we consider to have a greater like- lihood of being collected in the future as they have been recorded from counties in the adjacent states that are contiguous with West Virginia. Additional species and tax- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON onomic corrections will likely be incorpo- rated into the West Virginia checklist from ongoing revisionary efforts of S. W. Szczyt- ko on eastern North American Isoperlinae (Perlodidae) and P. P. Harper on eastern North American Leuctridae. GROUP EUHOLOGNATHA Family Capniidae Allocapnia curiosa Frison. Drainage basins iy tie tvV.~ Vv, Vil (Braxton Bayetie: Grant, Greenbrier, Hampshire, *Hardy, *McDowell, *Mercer, *Monroe, *Nich- olas, Pendleton, Pocahontas, Randolph, *Wayne, *Webster) [MD, PA,VA]. A. forbesi Frison. Drainage basin VII (Wayne) [KY, OH]. A. frisoni Ross and Ricker. Drainage basins Il, V, VI, VII (*Boone, Calhoun, Fayette, Greenbrier, Kanawha, Logan, McDowell, Mingo, *Nicholas, *Pocahontas, Preston, Randolph, Ritchie, Tucker, *Wayne, *Webster) [KY, MD, OH, VA]. A. frumi Kirchner. Drainage basins H, V (Fayette, Greenbrier, *Nicholas, Poca- hontas, *Upshur, *Webster) [MD]. A. granulata (Claassen). Drainage basins I, MH, UT V, Vi CCabell, “Greenbrier Hampshire, *Hancock, *Hardy, *Kana- wha, Logan, Mason, Mineral, *Monroe, Pendleton, Pocahontas, Preston, *Put- nam, *Randolph, *Summers, Wood) [KY, MD, OH, PA]. A. harperi Kirchner. Drainage basin V (Greenbrier, *Nicholas, Pocahontas) [MD, PA, VA]. **A. illinoensis Frison. Drainage basin I (*Mason, Ohio River, 31 January 2003, 1 6, R. FE Kirchner) [MD, PA, VA]. A. loshada Ricker. Drainage basins II, V, VIT (McDowell, *Monroe, Randolph) [VA]. A. maria Hanson. Drainage basins II, III (Hardy, Preston, *Randolph) [MD, PA, VAI. A. mystica Frison. Drainage basins I, VI, VII (Cabell, Calhoun,*Mason, Ritchie, *Wayne) [KY, OH, VA]. VOLUME 108, NUMBER 2 431 OF WEST VIRGINIA SHOWING DRAINAGE ANO MAJOR WATERSHEDS Fig. 1. Drainage basins of West Virginia (according to Janssen 1973). I = Ohio River, Il = Monogahela River, IIf = Potomac River, [1V = Little Kanawha River, V = Kanawha River, VI = Guyandot River, VII = Big Sandy River. A. nivicola (Fitch). Drainage basins I-VI son, *Mineral, *Monroe, *Morgan, *Pen- (*Boone, *Braxton, *Cabell, Calhoun, Doddridge, Fayette, *Grant, Greenbrier, Hampshire, *Hardy, Lincoln, Logan, *Mason, McDowell, *Mercer, Mineral, Mingo, *Monongalia, *Morgan, Nicho- las, Pendleton, Pocahontas, Preston, *Ra- leigh, Randolph, *Upshur, *Wayne, *Webster) [KY, MD, OH, VA]. A. ohioensis Ross and Ricker. Drainage ba- sins VI, VII (Cabell, *Wayne) [KY, OH]. A. pygmaea (Burmeister). Drainage basins I-VII (*Braxton, *Cabell, *Greenbrier, *Hampshire, *Hardy, “Kanawha, *Ma- dleton, Pocahontas, *Putnam,*Randolph, *Summers, *Wayne, *Webster) [K Y, OH, PA, VA]. A. recta (Claassen). Drainage basins I-VII (Braxton, *Cabell, Clay, Grant, *Green- brier, Hampshire, *Kanawha, Lincoln, Mason, *Mercer, Mineral, Mingo, Mon- roe, *Nicholas, Pendleton, *Pocahontas, Preston, *Raleigh, Randolph, Roane, Summers, *Wayne, *Webster) [KY, MD, OH, PA,VA\]. . rickeri Frison. Drainage basins I—VII (Barbour, *Braxton, Cabell, Calhoun, 4 rae NICHOLAS a 4 LINCOLN BOONE “Ss FAYETTE ZL RALEIGH N \ WYOMING \ eae \ McDOWELL MERCER Fig. 2. County map of West Virginia. Doddridge, Gilmer, Greenbriar, *Hamp- shire, Kanawha, Mason, Mercer, *Min- eral, Monroe, Ohio, Pendleton, Preston, Putnam, Pocahontas, *Randolph, Ritchie, Roane, *Summers, Wayne) [KY, MD, OH, PA, VA]. A. vivipara (Claassen). Drainage basins I, II, IV, V, VI, VII (Cabell, Calhoun, Dod- dridge, *Hancock, Lewis, Ohio, *Mason, McDowell, Preston, Putnam, Ritchie, Ro- ane, *Wayne) [KY, MD, OH, PA, VA]. A. zola Ricker. Drainage basins II, IV, V, VI (*Braxton, Clay,” Lincoln, <*Mercer, *Nicholas, Pocahontas, Preston, *Ra- leigh, *Randolph, *Webster) [MD, OH, PA, VA]. Paracapnia angulata Hanson. Drainage ba- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sins I, I, V, VI; VIL @ayettesiGrant Greenbrier, Hampshire, *Hardy, Logan, *Morgan, *Nicholas, *Pendleton, Poca- hontas, Randolph, *Summers, Tucker, *Upshur, “Wayne, Webster) [KY, MD, OH, PA, VA]. Family Leuctridae Subfamily Leuctrinae Leuctra alexanderi Hanson. Drainage ba- sins II, II, VI (Hardy, Logan, Tucker) [KY, MD, PA, VA]. L. biloba Claassen. Drainage basins II, III (*Hardy, Pendleton, Tucker) [VA]. L. carolinensis Claassen: Drainage basin II (Tucker) [MD, PA, VA]. VOLUME 108, NUMBER 2 L. duplicata Claassen. Drainage basins II, III, V (*Greenbrier, Pendleton, Tucker) [MD, PA, VA]. L. ferruginea (Walker). Drainage basins II, lil, V, VI, VU (*Hardy, Logan, Mingo, Pendleton, Pocahontas, Preston, Ran- dolph, Tucker, *Wayne) [KY, MD, OH, PA, VA]. L. grandis Banks. Drainage basins IH, V (*Greenbrier, Tucker) [MD, PA]. L. maria Hanson. Drainage basins I, VI (Lincoln, Tucker) [PA]. L. rickeri James. Drainage basin VII (*Wayne) [KY, MD, OH, VA]. L. sibleyi Claassen. Drainage basins II, I], V, VI, VII (Cabell, *Greenbrier, * Hardy, Logan, Mingo, *Pocahontas, Tucker) [KYSMDVOH.PAYV A). L. tenella Provancher. Drainage basin II (Randolph, Tucker) [MD, PA, VA]. L. tenuis (Pictet). Drainage basins III, V, VI (Logan, Mineral, Pocahontas) [KY, MD, OH, PA, VA]. L. triloba Claassen. Drainage basin II (Tucker) [VA]. L. truncata Claassen. Drainage basins I, ll, V, VI (Greenbriar, Lincoln, Pendle- ton, Pocahontas, Tucker, Upshur) [PA, VA]. + L. variabilis Hanson. [MD, PA, VA]. Paraleuctra sara (Claassen). Drainage ba- Sinseell,, Mile Ive. Vs VIVil. (Braxton, *Greenbrier, “Kanawha, *Hardy, Logan, *McDowell, Mingo, *Morgan, *Pendle- ton, *Pocahontas, *Randolph, Tucker, *Wayne) [KY, MD, OH, PA, VA]. Zealeuctra claasseni (Frison). Drainage ba- sin IV (Ritchie) [KY]. Z. fraxina Ricker and Ross. Drainage basin Ill (Hardy) [KY, OH]. Subfamily Megaleuctrinae Megaleuctra flinti Baumann. Drainage ba- sins II, V (Greenbrier, Pocahontas, Tuck- er) [MD, PA, VA]. Family Nemouridae Subfamily Amphinemurinae Amphinemura delosa (Ricker). Drainage basins I-VII (Berkeley, *Brooke, Cabell, 433 Grant, “Hancock, Harrison, Lincoln, Ma- son, Mercer, Mineral, Morgan, Pendleton, Preston, Putnam, Ritchie, Tucker, Tyler, *Upshur, Wayne, Wirt) [KY, MD, OH, PA, VA]. A. nigritta (Provancher). Drainage basins I, Tn Ve Ve Vil (Cabell) Fayette; Hampshire, *Hardy, *Kanawha, *Logan, *Mingo, *Pendleton, Pocahontas, *Ran- dolph; Ritchie, Roane, Tucker, Tyler *Wayne, *Wirt) [KY, MD, OH, PA, VA]. A. wui (Claassen). Drainage basins II, V (*Fayette, Pocahontas, Randolph, Tucker, Webster) [KY, MD, PA, VA]. Subfamily Nemourinae Ostrocerca albidipennis (Walker). Drainage basins II, HI, V (*Greenbrier, Grant, *Hardy, Pocahontas, *Randolph, Tucker) [MD, PA, VA]. O. complexa (Claassen). Drainage basins II, Il], V (*Hardy, Pendleton, Pocahontas, Randolph, Tucker) [MD, PA, VA]. O. prolongata (Claassen). Drainage basins II, II (*Pendleton, Tucker) [PA, VA]. O. truncata (Claassen). Drainage basins II, TE Vie NV 1. (Cabell, “Greenbrier, Hardy, Logan, Pendleton, Pocahontas, Tucker) [KY, MD, OH, PA, VA]. Paranemoura perfecta (Walker). Drainage basins II, III, V (*Greenbrier, *Hardy, Pocahontas, Randolph) [MD, PA, VA]. Prostoia completa (Walker). Drainage ba- sins II, IV, V (Braxton, *Hampshire, *Hardy, *Kanawha, *Pendleton, *“Poca- hontas) [KY, MD, PA, VA]. P. similis (Hagen). Drainage basins IJ, III, V, VI, VU (Braxton, Grant, *Hampshire, *Kanawha, *Lewis, Logan, *Mineral, Mingo, *Pendleton, Pocahontas, *Put- nam, *Wayne) [KY, MD, OH, PA, VA]. Soyedina carolinensis (Claassen). Drainage basins Hl, IV, V, VI (*Braxton, Logan, *Pocahontas, Randolph) [MD, VA]. + §. kondratieffi Baumann and Grubbs. [MD]. S. vallicularia (Wu). Drainage basins II, III, VI, VII (*Hardy, *Lincoln, Logan, *Mc- 434 Dowell, Mingo, *Upshur, Wyoming) [KY, MD, OH, PA, VA]. S. washingtoni (Claassen). Drainage basin II (Pendleton, Tucker) [MD, PA]. Family Taeniopterygidae Subfamily Brachypterinae Bolotoperla rossi (Frison). Drainage basins i TV, V;, Nil (Braxton, Greenbrier *TLewis, *Nicholas, “Pocahontas, Wayne) [PA, VA]. Oemopteryx contorta (Needham and Claas- sen). Drainage basins IH], V (Greenbrier, *Nicholas, *Pocahontas, Raleigh, Tucker, *Upshur, *Webster) [KY, MD, PA, VA]. O. glacialis (Newport). Drainage basins IV, V (Braxton, *Greenbrier, *Monroe, Po- cahontas, *Summers). Strophopteryx appalachia Ricker and Ross. Drainage basins I, I], Ul, IV, V, VU (*Braxton, Hampshire, *Kanawha, *Lewis, *Mason, Mercer, Mingo, *Po- cahontas, Preston, *Putnam, Tucker, *Wayne, *Webster) [MD, PA, VA]. S. fasciata (Burmeister). Drainage basins I, II, U1, IV, V, VII (*Braxton, *Cabell, Jef- ferson, Hampshire, *Hardy, *Greenbrier, *Kanawha, Mason, *Mineral, Mingo, *Monroe, Nicholas, Pendleton, Pocahon- tas, *Putnam, *Randolph, *Summers, Wayne, Webster) [KY, MD, OH, PA, VA]. Taenionema atlanticum Ricker and Ross. Drainage basins II, II, IV, V, VI, VII (Braxton, *Cabell, Fayette, Greenbrier, *Hardy, *Kanawha, *Lewis, Logan, Mer- cer, Mingo, *Nicholas, Pendleton, *Po- cahontas, *Putnam, Randolph, Wayne) [KY, MD, OH, PA, VA]. Subfamily Taeniopteryginae Taeniopteryx burksi Ricker and Ross. Drainage basins I, HI, IV, V, VI, VII (Braxton, Cabell, Clay, Fayette, Hamp- shire, Hardy, Kanawha, Mason, *Miner- al, Nicholas, *Putnam, *“Summers, Tyler, *Wayne, *Webster) [KY, MD, OH, PA, VAI. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON **7T\ lita Frison. Drainage basin I (*Mason, Ohio River, 31 January, 1995, 1 6, R.E Kirchner) [KY, VA]. T. maura (Pictet). Drainage basins II, Hl, IV, V, VI, VU (*Braxton, Cabell, Clay, Fayette, Grant, Greenbrier, Hampshire, *Hardy, Mineral, Mingo, *Monroe, Nich- olas, Pendleton, Pocahontas, Raleigh, *Randolph, *Summers, Wayne, Webster, Wyoming) [KY, MD, OH, PA, VA]. T. metequi Ricker and Ross. Drainage ba- sins I, II, IV, V, VI, VII (*Braxton, *Cab- ell, Clay, Greenbriar, Lewis, *Mason, Nicholas, Pocahontas, *Preston,* Wayne, Webster) [KY, MD, OH, PA, VA]. + T. nivalis (Fitch). [MD, PA]. T. parvula (Banks). Drainage basins I, IV, V, VI (*Braxton, *Cabell, Clay, *Green- brier, Kanawha, Mason, *Monroe, *Nich- olas, *Putnam, *Webster) [K Y, MD, OH, PA, VA]. T. ugola Ricker and Ross. Drainage basins I, , IV, Vv, Vi CBraxton +Cabell- Greenbrier, Mercer, Mingo, Nicholas, *Pocahontas, Preston,*Raleigh, *Ran- dolph, *Wayne, *Webster, *Upshur) [KY, MD, PA, VA]. Family Chloroperlidae Subfamily Chloroperlinae Alloperla aracoma Harper and Kirchner. Drainage basins II, III, IV, V, VI, VII (*Braxton, *Fayette, Logan, Mingo, Pen- dleton, Tucker) [MD, PA]. + A. atlantica Baumann. [MD, OH, PA, VA]. + A. banksi Frison. [VA]. A. biserrata Nelson and Kondratieff. Drain- age basins III, V (Grant, Hardy, Pendle- ton, Pocahontas) [MD, PA, VA]. A. chloris Frison. Drainage basins I, HI, V (*Hardy, Pendleton, Pocahontas, *Ran- dolph) [KY, MD, OH, PA, VA]. A. concolor Ricker. Drainage basin V (*Greenbrier, Nicholas, Pocahontas) [PA]. +A. idei (Banks). [KY, OH, VA]. A. imbecilla (Say). Drainage basins V, VII VOLUME 108, NUMBER 2 (Pocahontas, Summers, *Wayne, Web- ster) (KY, MD,"OH) PA, VA]. A. petasata Surdick. Drainage basin V (Po- cahontas) [OH, PA, VA]. A. usa Ricker. Drainage basins II, IV, V, VI, VII (Braxton, *Fayette, *Kanawha, Lo- gan, Mingo, Pocahontas, Randolph, Tucker) [KY, MD, OH, PA, VA]. Haploperla brevis (Banks). Drainage basins I-VII (Braxton, *Fayette, *Grant, Hardy, Hampshire, *Jefferson, *Kanawha, Lo- gan, Mason, *Mineral, Mingo, *Morgan, *Nicholas, Pendleton, Pocahontas, Pres- ton, Randolph, *Roane, *Summers, iucker Wayne) “[IK°Y,’ MID, ‘OH, PA, VA]. Rasvena terna (Frison). Drainage basin V (*Kanawha, Pocahontas) [PA, VA]. Suwallia marginata (Banks). Drainage ba- sins II, III, V (Greenbrier, Pendleton, *Pocahontas, Randolph, Tucker) [MD, PAs “WAT. Sweltsa lateralis (Banks). Drainage basins Il, Ul, 1V, V (Braxton, *Fayette, Green- brier, Hampshire, Hardy, *Kanawha, Nicholas, Pendleton, Preston, Pocahon- tas, Randolph, Tucker) [MD, OH, PA, VAI. S. naica (Provancher). Drainage basin V (Pocahontas) [PA, VA]. S. onkos (Ricker). Drainage basins I —VII (Braxton, Fayette, Greenbrier, Logan, Monongalia, Nicholas, Pendleton, Poca- hontas, *Preston, Randolph, Tucker, Wayne, *Wirt) [KY, MD, OH, PA, VA]. S. palearata Surdick. Drainage basin III (Hardy, Mineral, Pendleton) [MD,VA]. S. pocahontas Kirchner and Kondratieff. Drainage basins IV, V (Braxton, Green- briar, Pocahontas) [MD]. Subfamily Paraperlinae Utaperla gaspesiana Harper and Roy. Drainage basin V (Pocahontas) [PA]. Group SYSTELLOGNATHA Family Peltoperlidae Subfamily Peltoperlinae 435 Peltoperla arcuata Needham. Drainage ba- sinsa i mi IVs Vv; VI, Vili (Braxton, *Grant, Greenbrier, “Hampshire, *Hardy, Logan, Mingo, *Monongalia, *Nicholas, Pocahontas, Tucker) [MD, OH, PA, VA]. P. tarteri Stark and Kondratieff. Drainage basins III, V, VI (Fayette, *Hardy, *Ka- nawha, Wyoming) [VA]. + Tallaperla anna (Needham and Smith). [VA]. T. maria (Needham and Smith). Drainage basins I, II, V (*Grant, Greenbrier, *Jef- ferson, *Mineral, *Morgan, Nicholas, *Pendleton, *Pocahontas, Raleigh, *Summers, *Tucker, Webster) [MD, PA, VAI. Family Perlidae Subfamily Acroneurinae Acroneuria abnormis (Newman). Drainage basins I-VII (*Barbour, Berkeley, Boone, Braxton, Fayette, Grant, Greenbrier, Hampshire, *Hardy, Jackson, *Jefferson, Lincoln, Logan, Marion, *Marshall, *Mercer, Mineral, Mingo, Monongalia, Monroe, Morgan, Nicholas, Pendleton, *Pleasants, Pocahontas, Preston, *Put- nam, Randolph, *Ritchie, Summers, *Taylor, Tucker, Upshur, Wayne, Web- Sten) [IK Y, MD OH SPAwmV A: A. arenosa (Pictet). Drainage basin Il (Berkeley, *Hardy, *Mineral) [MD, PA, VAI. + A. arida (Hagen). [PA]. A. carolinensis (Banks). Drainage basins I —VII (Barbour, *Berkeley, Boone, Brax- ton, \*Brooke, Clay.) Fayette, *Grant, Greenbrier, Hampshire, Hancock, *Har- dy, Kanawha, Lewis, Logan, *McDow- ell, Mercer, *Mineral, *Monongalia, Mingo, Morgan, Nicholas, Pendleton, *Pleasants, Pocahontas, Preston, Putnam, *Raleigh, Randolph, Summers, Tucker, *Tyler, Upshur, Wayne, Webster, Wetzel, Wyoming) [KY, MD, OH, PA, VA]. A. filicis Frison. Drainage basins II, II, IV, V, VII (Braxton, Greenbrier, *Jefferson, Mingo, Randolph, *Ritchie, *Summers) [KY¥oMD: OHYPA, VAi: 436 A. frisoni Stark and Brown. Drainage basins I, If, Ul, 1V, V (*Gilmer, *Greenbrier, *Hardy, *Jackson, *Mason, *Monongal- ia, *Nicholas, Pocahontas, *Putnam, *Randolph, *Ritchie, Summers, *Tyler) [KY, MD, OH, PA, VA]. A. internata (Walker). Drainage basins II, Ill, IV, V (*Barbour, Berkeley, Braxton, Clay, *Gilmer, Greenbrier, “Hardy, Mer- cer, Mineral, Monongalia, Morgan, Po- cahontas, Preston, Randolph, “Summers, Webster) [KY, VA]. A. kirchneri Stark and Kondratieff. Drain- age basins H, V, VI (Cabell, Nicholas, Preston) [KY, VA]. +A. kosztarabi Kondratieff and Kirchner. [VA]. A. lycorias (Newman). Drainage basins I— VII (Barbour, Berkeley, Boone, Braxton, Greenbrier, Grant, Lewis, Lincoln, Mon- ongalia, Monroe, *Nicholas, *Pendleton, *Pleasants, Pocahontas, Preston, Ran- dolph, Ritchie, Roane, Summers, Tucker, Wayne, Webster, Wood, Wyoming) [KY, MD ORE PAS VA. A. perplexa Frison. Drainage basins I, I, IV, V, VI, VII (Barbour, Doddridge, Gil- mer, Greenbrier, *Jackson, Lincoln, Mar- ion, Mason, Mingo, Monongalia, *Nich- olas, *Pleasants, Pocahontas, *Summers, *Tyler, Upshur, Wayne, Wirt, *Wood) [KY, OH, PA]. +Attaneuria ruralis (Hagen). [MD, OH, PA, VA] Eccoptura xanthenes (Newman). Drainage basins I—-VII (*Barbour, Berkeley, *Greenbrier, *Hardy, Jackson, Lincoln, Logan, Mingo, Monongalia, Nicholas, Pocahontas, *Raleigh, Ritchie, Roane, *Summers, Wayne) [KY, MD, OH, PA, VAI. Hansonoperla appalachia Nelson. Drain- age basin V (Greenbrier, Nicholas) [MD, aN WAN, H. hokolesqua Kondratieff and Kirchner. Drainage basin VII (Wayne) [KY]. **Perlesta cinctipes (Banks). Drainage ba- sin II (*Monongalia, no locality data, 1 6,6 2, L. Butler) [OH]. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON P. decipiens (Walsh). Drainage basins I—-VII (Braxton, *Cabell, “Clay; ‘Greenbrier, Jackson, Jefferson, Hampshire, *Mason, *Mineral, Monongalia, Pendleton, Pleas- ants, Pocahontas, Preston, Putnam, Ran- dolph, *Ritchie, Tyler, Wayne, Wirt) [KY, OH, PA, VA] **P. frisoni Banks. Drainage basin III (*Pendleton, Big Run of North Fork of South Branch of Potomac River, 29 July 1978, 1 3d, R. E Kirchner) [VA]. **P. nelsoni Stark. Drainage basin V (*Po- cahontas, Tea Creek of Williams River, 6 August 1977, 1 ¢, R. E Kirchner) [MD, VA]. ** P| teaysia Kirchner and Kondratieff. Drainage basins I, Il, V (*Kanawha, no locality data, 17 June1988, 1 3d, 1 2, D. C. Tarter; *Mason,; Ohio’ River «18 June1998, 135, 1 2, R. E Kirchner; *Ran- dolph, Shavers Fork, 19 September 1980, ) ¢; Di €: Farter) (MD, PAS WAI: Perlinella drymo (Newman). Drainage ba- sins I, Hf, I, V (Cabell, Hampshire, *Ma- son, Putnam, Tucker) [K Y, MD, OH, PA, VA]. P. ephyre (Newman). Drainage basins III, V (*Fayette, Hampshire) [KY, MD, OH, PAS VAI. Subfamily Perlinae **A gnetina annulipes (Hagen). Drainage basins III, V, (*Fayette, New River, 10 June 1978, 1 6,-L. Ke “Evans;-+Hamp- shire, Cacapon River of Potomac River, 16 July 1980, 2 2, BG. Kondtatefi, *Summers, New River 10 July 1978, 1 6, R. E Kirchner, Bluestone River, 22 June 1977, 1 5 1 2, R.E Kirchner, Blue- stone River, 2!" June49305ias, leek E Kirchner) [MD, VA, PA]. A. capitata (Pictet). Drainage basins I, III, IV, V, (Braxton, Clay, Grant, Greenbrier, Hardy, Mercer, Monroe, Nicholas, Poca- hontas, Randolph, Summers) [KY, MD, OH, PA, VA]. A. flavescens (Walsh). Drainage basin III (*Berkeley, Sleepy Creek, 24 July 1975, 2 2, L. Miller, *Hampshire, Romney, 15 VOLUME 108, NUMBER 2 June 1976, 1 2, L. Miller, *Hardy, no locality data, 19 July 1975, 1 2, L. Mill- er, * Mineral, Burlington, 10 July 1979, 2 2, L. Miller) [KY, MD, OH, PA, VA]. Neoperla choctaw Stark and Baumann. Drainage basin III (Berkeley) [PA]. N. clymene (Newman). Drainage basins III, V (*Hampshire, Summers) [VA, PA]. + N. robisoni Poulton and Stewart. [MD, PA]. N. stewarti Stark and Baumann. Drainage basin III (Berkeley, Hampshire) [KYY, MD, OH, PA, VA]. Paragnetina immarginata (Say). Drainage basins I, II, I, IV, V (*Braxton, Clay, *Grant, *Greenbrier, Hardy, Mineral, Nicholas, Pendleton, *Pleasants, Poca- hontas, Preston, Randolph, Tucker, Web- ster) [KY, MD, PA, VA]. P. media (Walker). Drainage basins II, Il, EV VARY Le (Braxton (\Clayen* Fayette, *Greenbrier, Lewis, Mercer, Mingo, *Monongalia, Monroe, Nicholas, Pendle- ton, Pocahontas, Preston, Randolph, Summers, Tucker, Upshur, Wayne, Web- ster) (CYS MD; OHSPA; VA]. Family Perlodidae Subfamily Isoperlinae Clioperla clio (Newman). Drainage basins alls ALE, We WN, eV (Cabell), “Hardy; Jackson, Logan, Mason, McDowell, Mer- cer, Mingo, Monongalia, Nicholas, Pen- dleton, Pocahontas, Putnam, Raleigh, Randolph, Summers, *Tucker, Wayne, Webster)) (KY, MD; OH; PA, VA]. Isoperla bilineata (Say). Drainage basins I, II], VI (*Cabell, Jackson, Lewis, Mason, Monongalia, Pocahontas, Ritchie) [KY, OH, PA, VA]. I. burksi Frison. Drainage basins I, VII (Jackson, Mason, *Wayne) [KY, MD, OH, VA]. I. cotta Ricker. Drainage basins III, VII (Mingo, Pendleton, Pocahontas). dicala Frison. Drainage basins II, V (*Kanawha, Randolph, Summers) [MD, PA, VA]. > 437 I. gibbsae Harper. Drainage basin VII (Wayne) [MD]. I. holochlora (Klapalek). Drainage basins I-VII (Hardy, Jackson, Logan, McDow- ell, Mingo, Pocahontas, Randolph, Rit- chie, Summers, Webster) [KY, MD, PA, VAI. I. lata Frison. Drainage basins II, HI, IV, V (*Braxton, Greenbrier, Nicholas, Pendle- ton, Pocahontas, Randolph, Webster) [RATeV AY: + I. major Nelson and Kondratieff. [VA]. I. marlynia Needham and Claassen. Drain- age basin V (Mercer) [KY, PA, VA]. + I. montana (Banks). [PA]. I. namata Frison. Drainage basins I-VII (*Braxton, Calhoun, Gilmer, Harrison, Jackson, Lewis, Logan, Mercer, Mingo, Monroe, Nicholas, Pendleton, Pocahon- tas, Putnam, Randolph, Summers, Web- ster) IKYS MD, OH, PAs-VAI: I. orata Frison. Drainage basins I, II, III, V (*Braxton, “Hampshire, Jackson, Mon- ongalia, Pocahontas, Randolph, Ritchie, Tucker) [OH, PA, VA]. I. richardsoni Frison. Drainage basins V, VII (Mingo, Pocahontas) [KY, PA]. I. signata (Banks). Drainage basin V (*Summers) [OH, PA, VA]. I. similis (Hagen). Drainage basins II, III, IV, V, VI, VI (Greenbrier, *Hardy, Lin- coln, McDowell, Monongalia, Morgan, Nicholas, Pendleton, *Pocahontas, Ran- dolph, Tucker, Webster, Wood) [KY, MD, PA, VA]. I. slossonae (Banks). Drainage basin II (Tucker) [PA, VA]. I. transmarina Newman. Drainage basins I, II, IV, V, VI, VII (Cabell, Calhoun, *Fay- ette, Gilmer, Jackson, Lewis, Marion, Mason, McDowell, Mercer, Monongalia, Monroe, Nicholas, *Pocahontas, Preston, Putnam, Randolph, Ritchie, Summers, Webster, *Wood) [KY, PA, VA]. Subfamily Perlodinae +Cultus decisus decisus (Walker). [PA]. C. verticalis (Banks). Drainage basins II, V 438 (Harrison, Kanawha, Nicholas, Pocahon- tas, Randolph, Webster) [PA, VA]. Diploperla duplicata (Banks). Drainage ba- sins I, I, IV, V, VI, VII (*Braxton, Cab- ell, Fayette, Greenbrier, Jackson, Kana- wha, Lewis, Mason, Monongalia, Mon- roe, *Nicholas, *Pocahontas, Putnam, Randolph, Ritchie, *Upshur, Wayne) [MD, PA, VA]. D. kanawholensis Kirchner and Kondra- tieff. Drainage basins II, IV, V (Braxton, Lewis, *Mercer, *Nicholas, Pocahontas) [VA]. + D. morgani Kondratieff and Voshell. [VA]. D. robusta Stark and Gaufin. Drainage ba- sins I, I, IV, V, VI, VII (*Braxton, Cab- ell, Calhoun, Fayette, Greenbrier, Jack- son, Kanawha, Lewis, Lincoln, Logan, Mason, Mingo, Monongalia, Monroe, Nicholas, Pleasants, *Pocahontas, Put- nam, *Raleigh, Randolph, *Upshur, Wayne, Webster, Wirt) [KY, MD, OH, PA, VA: Helopicus subvarians (Banks). Drainage basins II, IV, V, VI (*Braxton, *Cabell, *Kanawha, *Lewis, *Mercer, Nicholas, *Putnam, Randolph, Summers) [KY, PA, VA]. Isogenoides hansoni (Ricker). Drainage ba- sins I, HI, IV, V (*Braxton, Greenbrier, *Lewis, Nicholas, *Pendleton, Randolph, Tucker, Webster) [MD, PA, VA]. Malirekus hastatus (Banks). Drainage ba- sins IT, II, IV, V, VI, VII (Braxton, Clay, Fayette, Greenbrier, Kanawha, Logan, McDowell, Mercer, *Mineral, Mingo, Nicholas, Pendleton, Pocahontas, Pres- ton, *Raleigh, Randolph, *Summers, Tucker, *Upshur, Webster) [K Y, VA]. +Malirekus iroquois Stark and Szczytko. [MD, PA]. Remenus bilobatus (Needham and Claas- sen). Drainage basins II, HI, V, VII (*Berkeley, *Hardy, *Kanawha, Mingo, *Morgan, Tucker, *Wayne, Webster) [KY, MD, OH, PA, VA]. +Yugus arinus (Frison). [VA]. Y. kirchneri Nelson. Drainage basins H, III, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON IV, V, VI, VII (*Braxton, Calhoun, *Fay- ette, Greenbrier, *Hardy, Kanawha, Lo- gan, Mingo, Nicholas, *Pendleton, Po- cahontas, *Randolph, Tucker, *Wayne, Webster) [MD, PA, VA]. Family Pteronarcyidae Pteronarcys biloba Newman. Drainage ba- sins II, HI, IV, V (Braxton, Clay, Green- brier, Hardy, Mineral, Monongalia, *Monroe, Nicholas, Pendleton, Pocahon- tas, Preston, Randolph, Tucker, Webster) [KY, MD, OH, PA, VA]. P. comstocki Smith. Drainage Basins II, II], IV, V (*Braxton, Grant, *Greenbrier, *TLewis, Mineral, Monongalia, Nicholas, Pendleton, Pocahontas, Randolph, Web- ster) [KY, PA, VA]. P. dorsata (Say). Drainage Basins III, V (Clay, Hampshire, Jefferson, *Pocahon- tas, *Putnam) [KY, MD, OH, PA, VA]. P. proteus Newman. Drainage Basins II, III, IV, V, VI, VII (Boone, Braxton, Clay, Fayette, *Grant, Greenbrier, “Hampshire, Hardy, *Jefferson, Kanawha, Lewis, *Logan, *Mineral, Mingo, Monongalia, *Morgan, “Monroe, Nicholas, Pendleton, Pocahontas, Preston, *Raleigh, Ran- dolph, Summers, Tucker, *Upshur, Web- Ster) [KY, MID; "OH, PA; VAI DISCUSSION Presence/absence of the stonefly species for the seven drainage basins was analyzed using a computerized unweighted pair- group arithmetic average clustering of the mean character difference (Swofford 2002). The results (Fig. 3) indicate two large clus- ters one consisting of drainage basins II, III, and V and the other composed of I, IV, VI, and VII. This, in part, may reflect the larger number of species exhibited by drainage basins belonging to the former cluster. It also may reflect habitat differences between an ordered trellised drainage system that characterizes drainage basin III, the eastern half of drainage basin II, as well as the ex- treme eastern edge of drainage basin V, and the haphazard drainage system that char- VOLUME 108, NUMBER 2 Fig. 3. 439 Vi Vil Dendrogram resulting from computerized unweighted pair-group arithmetic average clustering of the mean character difference of presence/absence data of Plecoptera species from the seven drainage basins in West Virginia. I = Ohio River, If = Monogahela River, III = Potomac River, [V = Little Kanawha River, V = Kanawha River, VI = Guyandot River, VII = Big Sandy River. acterizes drainage basins I, IV, VI, VI, the western half of drainage basin II, and much of drainage basin V (Jansson 1973). The following 15 stonefly species were found in all drainage basins (I-VI) of West Virginia: Acroneuria abnormis, A. caroli- nensis, A. lycorias, Allocapnia nivicola, A. pygmaea, A. recta, A. rickeri, Amphine- moura delosa, Amphinemura nigritta, Ec- coptura xanthenes, Haploperla brevis, Iso- perla holochlora, I. namata, Perlesta deci- piens, and Sweltsa onkos. The highest per- centage (76.3) of the stonefly species was found in drainage basin V, while the lowest percentage (28.1) was noted in drainage ba- sin I (industrialization and farming). The distribution pattern (drainage basins I], III, IV, V, VI, VID) of Zsoperla similis, Para- leuctra sara, Malirekus hastatus, Paragne- tina media, Pteronarcys proteus, Taenione- ma atlanticum, Taeniopteryx maura and Yugus kirchneri appears to indicate sensi- tivity to industrial pollution but greater tol- erance to acid mine pollution. The distri- bution of Clioperla clio displays a pattern closely related with the areas of Pennsyl- vania sandstone found in the state. Ten of the 135 stonefly species in West Virginia were restricted to drainage basin V, while seven species were restricted to drainage basins III and six were restricted to drain- age basin II. Four or fewer stonefly species were restricted to drainage basins I, IV, VI, and VII. Collected from 37 counties, the perlid Acroneuria carolinensis was the most widely distributed stonefly species in the state. All 55 counties of West Virginia recorded stonefly representatives. ACKNOWLEDGMENTS We are indebted to the following individ- uals who loaned specimens or helped with identifications and records: J. Bailey, Wild- life Biologist I], West Virginia Department of Environmental Protection; L. Butler, West Virginia University; J. L Earle, Penn- sylvania Department of Environmental Pro- tection; S. A. Grubbs, Western Kentucky 440 University; R. W. Hood, US Geological Survey, Denver; R. E Kirchner, US Army Corps of Engineers, Huntington District; B. C. Kondratieff, Colorado State University; L. Miller, West Virginia Department of Ag- riculture; D. R. Smith, Systematic Ento- mology Laboratory, USDA, Washington, D.C.; and C. White, Compliance Monitor- ing Lab. We appreciate the suggestions from the pre-publication reviews of the manuscript from D. Chaffee, Forest Biolo- gist Consultant, Richmond, KY, R. E Kirch- ner, US Army Corps of Engineers, Huntington District, and M. S. Schorr and G. S. Van Horn, Department of Biological and Environmental Sciences, the University of Tennessee at Chattanooga. We also ac- knowledge the helpful comments of an anonymous reviewer. Special thanks to M. Shafer for typing the paper. LITERATURE CITED Alexander, K. D. and K. W. Stewart. 1999. Revision of the genus Suwallia Ricker (Plecoptera: Chlo- roperlidae). Transactions of the American Ento- mological Society 125: 185—250. Baumann, R. W. 1974. 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Mala iiittles Re Ey Korchnes Wa D> \Wat- kins, R. G. Farmer and D. Steele. 1975. Distri- bution of pteronarcid stoneflies in West Virginia 442 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Insecta; Plecoptera). Proceedings of the West Vir- Sarracenia purpurea L. (Sarraceniaceae). Pro- ginia Academy of Science 47: 79-85. ceedings of the Entomological Society of Wash- Turner, T.S., J. L. Pittman, M. E. Poston, R. L. Peter- ington 98: 119-121. son, M. Mackenzie, C. H. Nelson, and R.M. Duf- Zwick, P. 1973. Insecta: Plecoptera. Phylogenetisches field. 1996. An unusual occurrence in West Vir- System and Katalog. Das Tierreich, 94. Walter de ginia of stoneflies (Plecoptera) in the pitcher-plant, Gruyter and Company, Berlin. I-XXXI, 1—465. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 443—466 A REVIEW OF THE SPECIES OF THRIPS LINNAEUS, 1758 (THYSANOPTERA: THRIPIDAE) FROM AFRICA, EUROPE, AND THE MEDITERRANEAN REGION DaAvip A. NICKLE Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, Beltsville Agricultural Research Center—West, Bldg. 005, 10300 Baltimore Avenue, Beltsville, MD 20705-2350, U.S.A. (e-mail: dnickle@ sel.barc.usda.gov) Abstract.—A total of 130 species of thrips occurring in Africa, Europe, and the Med- iterranean region were intercepted by U.S. agricultural quarantine officers in shipments of plants and cut flowers at the various ports of entry in the United States from 1983 to 1999. Of the 24 most commonly intercepted species encountered by port identifiers during this period, 10 were species of the genus Thrips Linnaeus (7. tabaci Lindeman, T. fus- cipennis Haliday, 7. major Uzel, T. vulgatissimus (Haliday), 7. meridionalis Priesner, T. flavus Schrank, 7. atratus (Haliday), T. simplex (Morison) , T. nigropilosus Uzel, and T. australis (Bagnall)). This paper provides information on all 100 species of Thrips from Africa, Europe, and the Mediterranean region necessary for completion of Part III of a guide to the identification of thrips entering the U.S.A. from those regions. It is designed primarily to aid the identification capabilities of the U.S. Department of Agriculture, Animal and Plant Health Inspection Service (USDA, APHIS) identifiers at U.S. ports-of- entry, but those interested in thrips in general will also benefit from this information. Key Words: The genus Thrips Linnaeus, 1758, is the largest of the thripine genera, with 267 spe- cies worldwide (Mound and Kibby 1998), 100 of them from Europe, Africa, and the Mediterranean region (Table |). The genus has not as yet been revised on a worldwide basis, but New World species were first re- vised by Gentile and Bailey (1968) with 31 species; Jacot-Guillarmod (1975) upwardly revised this number, in part by revalidating names synonymized by Gentile and Bailey, and provided information on hosts and spe- cies distributions. Since then, several spe- cies previously assigned to the genus Tae- niothrips Amyot and Serville, 1845, were transferred to the genus Thrips when the ge- nus was redefined to include species pos- thrips, pests of flowers, Europe, Africa, Thrips, Taeniothrips sessing ctenidia on abdominal tergites V to VIII wherein the location of ctenidia on VIII is anteromesad of the spiracle (O’ Neill 1972; Mound et al. 1976; Bhatti 1978; 1980; Schliephake and Klimt 1979; Bhatti and Mound 1980; Mound and Palmer 1981; Mound and Walker 1982; Mound and Houston 1987). Nakahara (1994) provided an excellent review of the New World spe- cies of Thrips, defining characters, charac- terizing species with descriptions and an identification key, listing distributions, and commenting on hosts, biology, and other items of information. Much of the infor- mation in this paper is derived from the comprehensive works on the European fau- na of Thrips by Schliephake and Klimt 444 (1979) and Priesner (1964). While perhaps most of the European species are described (though in fact little is known about many of them), the African fauna is poorly known, and many species probably as yet have not been discovered. While preparing Part III (see Nickle 2002, 2004 for Parts I and II) of a series of papers which is intended to facilitate iden- tification of thrips species commonly inter- cepted at U.S. ports on a wide range of ag- ricultural commodities from these regions, I had intended to cover only the 23 most common species encountered by port iden- tifiers. But because of the magnitude and unwieldy amount of information on the ge- nus Thrips in these regions, I decided to review the entire regional fauna, so that Part III will be more useful for determining Thrips species at the various U.S. ports. This paper provides preliminary infor- mation about the Thrips fauna from Africa, Europe, and the Mediterranean region, their pest status, and likelihood of interceptions at U.S. ports-of-entry. It provides an overall picture of the regional species of the genus. Of these, only 23 species of Thrips will be covered in Part III, since over the course of more than 20 years, they have been the only species so far encountered. MATERIALS AND METHODS The original citation, list of synonyms, type locality, [location of the type], distri- bution (by country), and host records (often not complete) of each species found in Af- rica, Europe, and the Mediterranean region are reported below. It also discusses the likelihood of encountering the species in commerce by port identifiers in the United States. Degrees of likelihood consisted of the following: very unlikely (either known only from type material, are very rarely col- lected, or have limited distribution and a single known host); unlikely (have a broad- er distribution, but limited number of hosts, usually not found on commercial hosts); possible (has a broad distribution, with at least some hosts of commercial export val- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ue); probable (has been intercepted at least occasionally, has a broad distribution with a number of commercially exportable hosts); and very probable (is on the list of the 23 most commonly intercepted species of Thrips from the above regions). Table 1 tabulates the most recent list of Thrips species found in Africa, Europe, and the Mediterranean region and summarizes their distribution in the regions of interest. It does not report on additional distributions (e.g., Oriental Region, Australasian Re- gion), although numerous species in this list have been transported to those regions through commerce. Table 2 lists only the more likely species to be encountered at U.S. ports-of-entry. In this table, some of the primary characters used to separate se- lected Thrips species are provided in matrix form to separate these species. More de- tailed information on these species will be undertaken in Part III of the identification guide (Nickle, in progress). Character values in Table 2 are encoded as follows: # ant. segs.: number of antennal segments (either 7 or 8 in number); acc. se- tae on abd. sterns.: accessory setae (besides marginal setae) on abdominal sternites. 0 = absent, 1 = present; acc. setae on pleurt. III— IV: absent = O, present = 1; # setae on lat. marg. terg. IT; actual number of setae on lat- eral margins of abdominal tergite II (either 3 or 4); comb on terg. VIIT: refers to devel- opment of microtrichia across distal margin of tergite VII, absent = 0, microtrichia de- veloped only laterally, absent medially = 1, comb of microtrichia complete across entire margin = 2; # distal setae on forewing: ac- tual number of setae on forevein of forewing distad of interruption of setae along vein; usually 3 in number but larger in number in several species and forevein without any in- terruption in setal arrangement in 7. aus- tralis; wing coloration: clear or unpigment- ed = O, lightly shaded either brown or gray = 1, darkly pigmented = 2. RESULTS Most of the information in the following treatment is based on the works of Gentile VOLUME 108, NUMBER 2 445 Table 1. Species of Thrips in Europe, the Mediterranean, and Africa. Areas of distribution are indicated with an ‘‘x’’. Establishment or occurrence of any of these species in the United States or South America also are indicated by an **x’’, but these distributions are likely introductions from either Europe, Africa, or the Mediter- ranean region. Superscripted numbers refer to actual states in the United States where a particular species has become established. Species Eur. Med. Afr. US. S.Am. acaciae Trybom, 1911 X albopilosus Uzel, 1895 alectorolophi Oettinger, 1953 alni Uzel, 1895 angusticeps Uzel, 1895 ~ eK OK armeniacus Pelikan, 1973 x asparagi zur Strassen, 1968 X asper Haliday, 1852 asperulae Jordan, 1888 assimilis Bagnall, 1913 atratus Haliday, 1836 aurantithoracis Hood, 1932 X Fal fal cal fal oe * australis Bagnall, 1915 X x X xe i banicus zur Strassen, 1968 benseleri Frauenteld, 1867 brevicornis Priesner, 1920b brevisetosus Trybom, 1911 x calcaratus Uzel, 1895 cereolus Oettingen, 1944 conferticornis Priesner, 1922 corymbiferarum Haliday, 1836 crassicornis Bagnall, 1923 dentatus Solowiow, 1924 difficilis Priesner, 1920a dilitatus Uzel, 1895 discolor Haliday, 1836 disjunctus Oettingen, 1944 dubius Priesner, 1927 ebneri Karny, 1914 euphorbiae Knechtel, 1923 eT oo oo oo os euphorbiicola Bagnall, 1924 exilicornis Hood, 1932 flavus Schrank, 1776 fulvipes Bagnall, 1923 funebris Bagnall, 1923 fuscipennis Haliday, 1836 grossulariae Haliday, 1836 herricki Bagnall, 1923 hispidipennis Hood, 1932 hoodi Priesner, 1938 incognitus Priesner, 1914 ~ eK KK OK inopinatus zur Strassen, 1963 italicus (Bagnall, 1926) Juniperinus Linnaeus, 1758 ~ KK K MK Pal kikuyuensis Trybom, 1911 Xx klapaleki Uzel, 1895 x leucadophilus Priesner, 1936 X lini Ladureau, 1878 Xx 446 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Species Eur. Med. Afr. U:s: S.Am. a lividus Haliday, 1836 x major Uzel, 1895 mx mancosetosus (Maltbaek, 1928) Xx mareoticus (Priesner, 1932) Xx mariae Cotte, 1924 Ke medialis Oettingen, 1951 ax mediterraneus Priesner, 1934 Xx menyanthidis Bagnall, 1923 meridionalis Priesner, 1926 Xx x microchaetus Karny, 1920 minutissimus Linnaeus, 1758 xe me montivagus Priesner, 1934 nigropilosus Uzel, 1895 Xx ? X Ke obscuricornis Priesner, 1920b Xx origani Priesner, 1926 oryzophagus Rondani, 1873 Xx palmi Karny, 1925 ? Ke ? paludosus Bagnall, 1913 Xx paluster Reuter, 1899 Xx panousei zur Strassen, 1968 Xx paucisetosus Priesner, 1927 pelikani Titschack, 1962 pennatus zur Strassen, 1968 persicae Haliday, 1836 mm PS PM OP OPS physapus Linnaeus, 1758 phytolaccae Priesner, 1951 pillichi Priesner, 1927 pilosissimus Priesner, 1927 ponticus Knechtel, 1965 mM mM RO poultoni Bagnall, 1933 praetermissus Priesner, 1920b Ke priesneri Hood, 1932 pusillus Bagnall, 1926 quadridentatus Trybom, 1910 ranunculi Schrank, 1781 robustus Priesner, 1920a roepkei Doeksen, 1953 sambuci Heeger, 1854 simplex (Morison, 1930) spadix Hood, 1932 spoliatus Priesner, 1964 mm mK OS Pal tabaci Lindeman, 1889 Xx x x Ko xe tarfayensis zur Strassen, 1968 tenellus Trybom, 1912 tenuisetosus Knechtel, 1923 timidus Priesner, 1926 trehernei Priesner, 1927 urticae Fabricius, 1781 uzelianus Priesner, 1926 validus Uzel, 1895 variegatus von Gleichen, 1764 Po lo 9 os a a VOLUME 108, NUMBER 2 447 Table 1. Continued. Species Eur. Med. Afr. US. S.Am viminalis Uzel, 1895 — vulgatissimus Haliday, 1836 yee ' atratus: Connecticut, Maine, Maryland, Michigan, New Jersey, New York, Oregon, Vermont, Washington. ? australis: California, Hawaii. 3 calcaratus: Michigan, New York, Wisconsin. + discolor: Connecticut, New York, Pennsylvania, Wisconsin. 5 flavus: New York. © fuscipennis: Illinois, New York, Oregon, Utah, Washington. 7 herricki: New Jersey, New York. 8 nigropilosus: California, Georgia, Illinois, Indiana, lowa, New Jersey, New York, South Dakota, Virginia, Washington. ° palmi: Florida. '0 physapus: Illinois, Michigan, New Jersey, New York, Utah. 'l pilosissimus: Colorado. "> simplex: All states. '3 tabaci: All states. '4 trehernei: California, Colorado, Connecticut, District of Columbia, Georgia, Idaho, Illinois, Indiana, Iowa, Kansas, Maine, Maryland, Michigan, Missouri, Montana, Nebraska, New Jersey, New York, Ohio, Oregon, Utah, Virginia, Washington, Wisconsin, West Virginia, Wyoming. 'S yalidus: Idaho. 'e ypulgatissimus: Alaska, Arizona, California, Colorado, Georgia, Idaho, Illinois, Maine, Montana, New Jersey, New Mexico, Nevada, Oregon, South Dakota, Utah, Washington, Wyoming. and Bailey (1968), Jacot-Guillarmod (1975), Mound et al. (1976), Nakahara (1994), Priesner (1964), and Schliephake and Klimt (1979). Information on most of these species is also based on 436 slide- mounted specimens of these species in the National Thrips Collection, National Mu- seum of Natural History, Smithsonian In- stitution, located at Beltsville, Maryland. Of the 100 species listed, only 35 are likely to be intercepted in commerce at U.S. ports. These include the 23 most often encoun- tered species, although several species not as yet intercepted are widely distributed and found on many of the same commodities that the commonly encountered species oc- cur. Thrips acaciae Trybom, 1910 Thrips acaciae Trybom, 1910: 161. Type locality. Kalahari: Pfanne Kooa. [Swed- ish Museum of Natural History, Stock- holm]. Synonyms: Thrips hirtiventris Hood, 1932. Distribution.—(Africa) Botswana; South Africa; Tanzania. Hosts.—Acacia caffra, A. horrida, A. ni- gricens, A. sp., Athanasia acerosa, Caesal- pinia sp., Cassine burkeana, Citrus spp., Combretum sp., Cyprerus articulatus, Dal- bergia armata, Elyonurus argenteus, Indi- gofera hilaris, Lessertia brachystachya. Likelihood of interception.—Possible; occasionally on commodities, including greenhouse plants. Thrips albopilosus Uzel, 1895 Thrips albopilosus Uzel, 1895: 190. Type locality. Bohemia: Hradec Kralove. [Vi- enna Museum? ]. Synonyms: Thrips albopilosus var. obscur- icornis Priesner, 1920b. Distribution.—(Europe) Albania, Aus- tria, Czech Republic, England, France, Georgia, Germany, Latvia, Poland, Scot- land, Serbia, Slovakia, Russia, Ukraine. (North America) USA [Utah]. Hosts.—Humulus lupulus, Juniperus communis, Phyteuma Populus alba, Quercus robur, Trifolium pratense, wild roses, Veronica spuria, various green- house plants. spicatum, 448 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. A character matrix for species of Thrips from Europe, the Mediterranean, and Africa most likely to be encountered at U.S. ports-of-entry. Character values are encoded as follows: # ant. segs.: number of antennal segments (either 7 or 8 in number); acc. setae on abd. stern.: accessory setae (besides marginal setae) on abdominal sternites, 0 = absent, | = present; acc. setae on pleurt. III-IV: accessory seta on pleurotergites, 0 = absent, 1 = present; # setae on lat. marg. terg. IJ: actual number of setae on lateral margins of abdominal tergite II (either 3 or 4); comb on terg. VIII: refers to development of microtrichia across distal margin of tergite VIII, 0 = absent, | = microtrichia developed only laterally, absent medially, 2 = comb of microtrichia complete across enter margin; # distal setae on forewing: actual number of setae on forevein of forewing distad of interruption of setae along vein; usually 3 in number but larger in number in several species and forevein without any interruption in setal arrangement in 7. australis; wing coloration, actual color range from yellow to brown. Accessary Accessary Comb of # Setae on Setae on # Setae Michro- # Distal Antennal Abd. Pleurt. on Lateral trichia on Setae on Wing Species Segs. Sternites III-IV Marg. Terg II Terg. VII Forewing Coloration angusticeps 7 | 0 3-4 |—2 6-9 pale brown atratus 8 0 l 3 (rarely 4) 2 6-9 brown australis 8 | | 4 | complete brown brevicornis 7 0 3 2 3 pale yellow flavus 7 0 0 4 2 3 pale yellow fulvipes 7 0 | 3) 3} pale yellow fuscipennis 7 0 0) 4 | 3 light brown italicus 8 | l 3 2 complete light brown major vi O 0 3 l 3 light yellow meridionalis 8 l l 3 2 3) gray nigropilosus 7 O O 3 2 3 pale gray palmi 7 0 0 4 2 3} pale yellow physapus 8 1 O 3 2 3 gray or brown pillichi 7 l 0 3 l 3 yellow simplex 8 | O 3 2. complete grayish brown tabaci 7 O O 3 2 4—5 (rare 6) pale yellow trehernei 7 O 0 4 (rarely 3) l 3 pale brown vulgatissimus 8 | | 3) O-1 3 pale yellow Likelihood of interception.—Possible; Distribution.—(Europe) Austria, Czech widely distributed and occasionally on Republic, England, Finland, Germany, Ita- commodities, including greenhouse plants. — ly, Poland, Romania, Slovakia, Switzerland, Ukraine. (Asia) Trancaucasia. Hosts.—Alnus incana, A. glutinosa, As- Thrips alectorolophi Oettingen, 1953: 164. tragalus alpinus, Betula verrucose, Carpi- Type locality. Germany: der Helme bei Martinsried (Harz). [Deutsches Entomo- logisches Institut]. Thrips alectorolophi Oettingen, 1953 nus betulus, Fagus orientalis, F. silvatica, Larix sp., Prunus sp., Salix caprea, S. pur- purea. Distribution Germany. Likelihood of interception.—Possible; Hosts.—Alectorolophus minor. widely distributed and occasionally on Likelihood of interception—Very un- commodities. likely; known only from type material. Thrips alni Wel 1895 Thrips angusticeps Uzel, 1895 Thrips alni Uzel, 1895: 189. Type locality. Thrips angusticeps Uzel, 1895: 191. Type Bohemia: Opocno. [Vienna Museum’?]. locality. Bohemia. [Vienna Museum’? ]. Synonyms: Thrips alni var. humuli Priesner, Synonyms: Achaetothrips loboptera Karny, 1920b; Thrips alni var. insperata Pries- 1908; Bagnallia asemus Williams, 1913; ner, 925: Thrips ebneri Karny, 1914. VOLUME 108, NUMBER 2 Distribution.—(Europe) Austria, Canary Islands, Czech Republic, Denmark, Eng- land, Finland, France, Georgia, Germany, Hungary, Lithuania, Netherlands, Poland, Scotland, Spain, Sweden, Switzerland, Ukraine, former Yugoslavia, Wales. (Afri- ca) Egypt, Morocco. (Mediterranean) Pal- estine [Israel], Persia [Iran, Iraq]. Hosts.—Numerous plants; a pest on flax. Likelihood of interception.—Probable; both widely distributed and found on many hosts, including commodities; a pest on flax. Thrips armeniacus Pelikan, 1973 Thrips armeniacus Pelikan, 1973: 34. Type locality. Armenia: Lake Sevan, NW shore near road to Yerevan, 1,950 m. [Vienna Museum? ]. Distribution.—(Europe) Armenia: Lake Sevan. (Asia) Trancaucasia. Hosts.—Low plants. Likelihood of interception.—Very un- likely. Thrips asparagi zur Strassen, 1968 Thrips asparagi zur Strassen, 1968: 48. Type locality. Southern Morocco: Cap Rhir, nw Agadir [Senckenberg Museum, Frankfort am Main]. Distribution.—Southern Morocco. Hosts.—Asparagus spp. Likelihood of interception.—Very un- likely; known only from type material; found on north African Asparagus. Thrips asper Haliday, 1852 Thrips asper Haliday, 1852: 1109. Type lo- cality—England [??]. Distribution.—England, Hungary. Hosts.—Unknown. Likelihood of interception.—Very un- likely; very little is known of this species. Thrips asperulae Jordan, 1888 Thrips asperulae Jordan, 1888: 568. Type locality. Germany: G6ttingen [??]. 449 Distribution.—Germany. Hosts.—Asperula odorata. Likelihood of interception.—Very un- likely; very little is known of this species. Thrips assimilis Bagnall, 1913 Thrips assimilis Bagnall, 1913: 294. Type locality. Tunis: Sousse [The Natural His- tory Museum, London]. Distribution.—Tunisia. Hosts.—Unknown. Likelihood of interception.—Very un- likely; very little is known of this species. Thrips atratus Haliday, 1836 Thr.[ips] atrata Haliday, 1836: 447. Type locality. Britain [Unknown]. Synonyms: Physopus atrata var. adusta Uzel, 1895; Taeniothrips atratus var. lon- gicornis Priesner, 1926; Ceratothrips britteni Bagnall, 1914. Distribution.—(Europe) Albania, Aus- tria, Azores, Czech Republic, Denmark, England, Estonia, France, Georgia, Ger- many, Hungary, Ireland, Italy, Lithuania, Netherlands, Poland, Romania, Russia, Scotland, Serbia, Slovakia, Spain, Sweden, Switzerland, Turkey, Ukraine, former Yu- goslavia. (Mediterranean) Cyprus. (North America) Canada (Ontario), USA [Con- necticut, Maine, Maryland, Michigan, New Jersey, New York, Oregon, Vermont, Wash- ington]. Hosts.—Numerous flowers, especially those of Caryophyllaceae, Labiatae, and Compositae. Likelihood of interception.—Probable; in- tercepted frequently on a variety of flowers. Thrips aurantithoracis Hood, 1932 Thrips aurantithoracis Hood, 1932: 135. Type locality. Cameroons [National Mu- seum of Natural History, Washington, DC]. Distribution.—Cameroon. Hosts.—flowers of daisy. Likelihood of interception.—Very un- likely; limited distribution and hosts. 450 Thrips australis (Bagnall 1915) Isoneurothrips australis Bagnall, 1915: 592; Bailey, 1957: 182; Jacot-Guillar- mod, 1974: 844. Type locality. Western Australia: Mundaring Weir, Darling Range, Perth [The Natural History Mu- seum, London]. Thrips australis (Bagnall): Bhatti, 1980: IN Thrips spinosus Hood: Johansen, 1974: 34 (misidentification) Synonyms: Anomalothrips amygdali Mor- gan, 1929. Distribution.—(Africa) Egypt, Kenya, Madagascar, Malawi, Morocco, Rhodesia, South Africa. (Europe) Canary Islands, Spain, Turkey. (Mediterranean) Cyprus, Palestine (Israel). (North America) Barba- dos, USA [California, Hawaii]. (South America) Brazil. (Asia) Japan. (Australasia) Australia, New Zealand, Tasmania. Hosts.—Acacia cyanophylla; A. pulchel- la; A. sp.; Adenocarpus foliosus villosus; Adenostoma sparsifolium, Buddleia japon- ica; Crataegus sp.; Dryandra floribunda; Eucalyptus leucoxylon, E. sp.; Goodenia sp.; Laurestina sp.; Leucadendron daph- noides; Leucaena glauca; Linum usitatissi- mum; Metrosideros sp.; Pinus canariensis; Pittosporum undulatum; Podocarpus ma- crophylla; Prunus amygdallus; Psidium guayava; Reseda odorata; Sonchus conges- tus; S. oleraceus; Tropaeolum majus; arum lily; blackberry, dahlia; Indian tea; orange; rose. Likelihood of interception.—Probable; intercepted frequently on a variety of flow- ers and commodities. Thrips banicus zur Strassen, 1968 Thrips banicus zur Strassen, 1968: 52. Type locality. Southern Morocco: Bou-Guejouf im Djebel Bani, s. Taidalte (Cercle de Goulimine) [Senckenberg Museum, Frankfort am Main]. Distribution.—Morocco. Hosts.—Gymnocarpos decander. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Likelihood of interception.—Very un- likely; limited distribution and hosts. Thrips banseleri Frauenfeld, 1867 Thrips banseleri Frauenfeld, 1867: 800. Type locality. Austria [Vienna Muse- um?]. Distribution.—Austria. Hosts.—Zea mays (in greenhouses). Likelihood of interception.—Very un- likely; limited distribution and hosts. Thrips brevicornis Priesner, 1920b Thrips brevicornis Priesner, 1920b: 59. Type locality. Austria: Ober6sterreich, al- pine region [Priesner Coll., Linz, Aus- tria]. Distribution.—Austria. Hosts.—variety of flowers. Likelihood of interception.—Probable; has been intercepted repeatedly on flowers from Europe. Thrips brevisetosus Trybom, 1911 Thrips kikuyuensis subsp. brevirostristris Trybom, 1911: 10. Type locality. British East Africa: escarpment [Stockholm Mu- seum]. Distribution.—Kenya, Congo. Hosts.—Yellow Eupatorium-like flow- ers. Likelihood of interception.—Very un- likely; known only from type locality, not known to occur on commodities. Thrips calcaratus Uzel, 1895 Thrips calcaratus Uzel, 1895: 195. Type lo- cality. Bohemia [Vienna Museum?]. Distribution.—(Europe) Austria, Czech Republic, Denmark, England, Finland, France, Georgia, Germany, Hungary, Italy, Romania, Scotland, Switzerland, Ukraine. (North America). USA [Michigan, New York, Wisconsin]. Hosts.—Tilia europaea, T. platyphyllos, T. vulgaris. Likelihood of interception.—Unlikely; VOLUME 108, NUMBER 2 although widely distributed, it has been found mainly on Tilia spp. Thrips cereolus Oettingen, 1944 Thrips cereolus Oettingen, 1944: 42. Type locality. Germany: Landsberg (Warthe). Distribution.—Germany. Hosts.—forest meadow. Likelihood of interception.—Very un- likely; known only from type material. Thrips conferticornis Priesner, 1922 Thrips conferticornis Priesner, 1922: 93. Type locality. Austria: Heratinger-See bei Ibm, Ober6sterreich [Priesner Coll., Linz, Austria]. Distribution.—Austria, Czech Republic, Finland, Germany, Hungary, Italy, Slovak- ia, Ukraine. Hosts.—Leucanthemum sp., Nasturtium sp., Ranunculus sp., meadow, turf. Likelihood of interception.—Unlikely; not intercepted as yet. Thrips corymbiferarum Haliday, 1836 Thrips corymbiferarum Haliday, 1836: 449. Type locality. Ireland: Glassnevin, near Dublin [The Natural History Museum, London]. Distribution.—Ireland. Hosts.—Flowers of Corymbiferae with a white border in botanical garden. Likelihood of interception.—Very un- likely; known only from type material. Thrips crassicornis Bagnall, 1923 Thrips crassicornis Bagnall, 1923: 59. Type locality. England: Newton Abbot, Devon- shire [The Natural History Museum, London]. Distribution.—England, France, Germa- ny, Romania. Hosts.—Euphorbia amygdaloides, Gal- ium cruciatum. Likelihood of interception.—Very un- likely; rare in collections. Thrips dentatus Solowiow, 1924 Thrips dentatus Solowiow, 1924: 25. Type locality. Russia: Gorki [Unknown]. Distribution.—Russia. Hosts.—Blossoming weeds. Likelihood of interception.—Very un- likely; known only from type material. Thrips difficilis Priesner, 1920a Thrips difficilis Priesner, 1920a: 75. Type locality. Austria: Grunberg, Oberoster- reich [Priesner Coll., Linz, Austria]. Distribution.—Austria, England, Estonia, France, Germany, Netherlands, Scotland, Ukraine. Hosts.—Several species of Salix. Likelihood of interception—Very un- likely; never intercepted and known only from material on Salix. Thrips dilitatus Uzel, 1895 Thrips dilitatus Uzel, 1895: 202. Type lo- cality. Bohemia [Vienna Museum? ]. Distribution.—Austria, Czech Republic, Denmark, England, Estonia, Finland, France, Germany, Netherlands, Russia, Scotland, Slovakia, Sweden, Switzerland, Ukraine. Hosts.—Euphrasia brevipila, E. frigida, E. nemorosa, E. occidentalis, Pedicularis palustris, P. sylvatica, Rhinanthus minor. Likelihood of interception.—Unlikely; not as yet intercepted. Thrips discolor Haliday, 1836 Thrips discolor Haliday, 1836: 449. Type locality. Britain [Unknown]. Synonyms: Thrips nubilans Hood, Thrips pallens Haliday, 1836. 1941; Distribution.—(Europe) Austria, Czech Republic, Denmark, England, Finland, Ger- many, Netherlands, Poland, Romania, Scot- land, Serbia, Slovakia, Sweden, Switzer- land, Ukraine. (North America) USA [Con- necticut, New York, Pennsylvania, Wiscon- sin]. 452 Hosts.—Ranunculus repens, a variety of other plants and grasses. Likelihood of interception.—Unlikely; not as yet intercepted. Thrips disjunctus Oettingen, 1951 Thrips disjunctus Oettingen, 1951: 56. Type locality. Poland: Swiec (Schwetz) [Un- known]. Distribution.—Poland. Hosts.—Unknown. Likelihood of interception.—Very un- likely; known only from type material. Thrips dubius Priesner, 1927 Thrips dubius Priesner, 1927: 58. Type lo- cality. not specified, either Austria or Hungary [Priesner Coll., Linz, Austria]. Distribution.—(Europe) Austria, Czech Republic, France, Germany, Hungary, Slo- vakia, Transcaucasia. (Mediterranean) Cy- prus, Turkey. Hosts.—Euphorbia cyparissias, turf. Likelihood of interception.—Unlikely; not found on general commodities. Thrips ebneri Karny, 1914 Thrips ebneri Karny, 1914: 54. Type local- ity. not specified [Priesner Coll.?, Linz, Austria]. Distribution.—(Europe) France, Sicily. (Africa) Morocco, Tunisia. (Mediterranean) Cyprus. Hosts.—Calendula arvensis, C. fulgida, Centaurea jacea. Likelihood of interception.—Unlikely; not found on general commodities. Thrips euphoribiae Knechtel, 1923 Thrips euphoribiae Knechtel, 1923: 75. Type locality. Rumania: Canrana (Vlas- ca) [Knechtel Coll., Bucharest]. Synonyms: Thrips euphorbiae f. adusta Priesner, 1927. Distribution.—Austria, Czech Republic, England, Germany, Hungary, Poland, Ro- mania, Slovakia, Ukraine. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hosts.—Euphorbia spp. Likelihood of interception—vVery un- likely; although widely distributed, known to occur only on species of Euphorbia. Thrips euphorbiicola Bagnall, 1924 Thrips euphorbiicola Bagnall, 1924: 115. Type locality. England: New Forest, Hampshire [The Natural History Muse- um, London]. Replacement name _ for Thrips euphorbiae Bagnall, 1924 [not T. euphorbiae Knechtel, 1923]. Synonyms: Thrips euphorbiella Bagnall, WG le Distribution.—England, France. Hosts.—Euphorbia amygdaloides, E. characias, E. dendroides, E. paralias. Likelihood of interception.—Very un- likely; limited distribution and not found on general commodities. Thrips exilicornis Hood, 1932 Thrips exilicornis Hood, 1932: 130. Type locality. S. Nigeria: Ibandan [sic] [Na- tional Museum of Natural History, Wash- ington, DC]. Distribution.—Nigeria, Sierra Leone, Tanzania, Uganda. Hosts.—Coffea robusta, Dolichos lablab, Ipomoea batatas, Melia azedarach, Musa sapientum, kingiramuti, mango, matovu, plaintain. Likelihood of interception.—Possible; found on a variety of agricultural commod- ities. Thrips flavus Schrank, 1776 Thrips flavus Schrank, 1776: 31. Type lo- cality. Austria [Unknown]. Synonyms: Thrips melanopa Schrank, 1781; Thrips flavidus Bagnall, 1916; Thrips flavus var. flavosetosus Priesner, 1919; Physothrips flavidulus Bagnall, 1923; Thrips flavus var. obscuricornis Priesner, 1927; Thrips clarus Moulton, 1928a; Thrips flavus var. kyotoi Moulton, 1928a; Thrips nilgiriensis Ramakrishna, VOLUME 108, NUMBER 2 1928; Thrips luteus Oettingen, 1935; Taeniothrips sulfuratus Priesner, 1935. Distribution.—(Europe) Austria, Azores, Czech Republic, Denmark, England, Fin- land, France, Germany, Hungary, Ireland, Italy, Netherlands, Poland, Romania, Rus- sia, Scotland, Slovakia, Sweden, Switzer- land. (Asia) Taiwan, China, Korea, Philip- pines, Transcaucasia, Turkistan. (Africa) Malawi. (North America) USA [New York]. Hosts.—Many plants. Likelihood of interception.—Probable; has a broad distribution and has been found on many hosts; intercepted frequently from various countries and on various hosts and commodities. Thrips fulvipes Bagnall, 1923 Thrips fulvipes Bagnall, 1923: 59. Type lo- cality. England: Shotover, Oxfordshire [The Natural History Museum, London]. Distribution.—Austria, Czech Republic, Denmark, England, France, Germany, Hun- gary, Romania, Scotland, Slovakia, Swe- den, Switzerland. (Asia) Transcaucasia. Hosts.—Brachypodium sylvaticum, Cor- ylus sp., Mercurialis perennis. Likelihood of interception.—Probable; has been found on various hosts from Eu- rope. Thrips funebris Bagnall, 1924 Thrips funebris Bagnall, 1924: 274. Type locality. England: Henigstbury, Hamp- shire [The Natural History Museum, London]. Distribution.—England, Germany. Hosts.—Carex vulpina, Triglochin mar- itima. Likelihood of interception.—Very un- likely; limited distribution and on non-eco- nomically important hosts. Thrips fuscipennis Haliday, 1836 Thrips fuscipennis Haliday, 1836: 448. Type locality. Britain [Unknown]. 453 Synonyms: Thrips fuscipennis f. drabae Priesner, 1927; Thrips meledensis Karny, 1907. Distribution.—(Europe) Albania, Aus- tria, Czech Republic, Denmark, England, Estonia, Finland, France, Germany, Hun- gary, Ireland, Italy, Lithuania, Netherlands, Poland, Romania, Russia, Scotland, Serbia, Slovakia, Sweden, Switzerland, Turkey, Wales, former Yugoslavia. (Mediterranean) Sardinia. (Asia) China, Trancaucasia. (North America) Canada [British Columbia, Quebec], USA [Illinois, New York, Oregon, Utah, Washington]. Hosts.—Plants of various families. Likelihood of interception.—Very prob- able; a common species frequently inter- cepted in ports on a variety of hosts. Thrips grossulariae Haliday, 1836 Thrips grossulariae Haliday, 1836: 448. Type locality. Britain. [?]. Distribution.—Britain, Italy. Hosts.—Flowers of gooseberry. Likelihood of interception—Very un- likely; not known to occur on commodities. Thrips herricki Bagnall, 1926 Thrips herricki Bagnall, 1926: 183. Type locality. Unknown [The Natural History Museum, London]. Synonyms: Thrips veratri Hood, 1927; Thrips veratri Herrick, 1927. Distribution.—(Europe) Czech Republic, France, Slovakia, Spain. (North America) USA [New Jersey, New York]. Hosts.—Veratrum album, V. viride. Likelihood of interception—Very un- likely; not as yet intercepted and known only from material on highly restricted host. Thrips hispidipennis Hood, 1932 Thrips hispidipennis Hood, 1932: 122. Type locality. Tanganyika Territory: Dar- es-Salam. [Unknown]. Distribution.—Tanzania. Hosts.—Flowers of Jasmine. 454 Likelihood of interception.—Very un- likely; known only from type material. Thrips hoodi Priesner, 1938 Thrips hoodi Priesner, 1938: 352. Type lo- cality. Belgischer Congo [Musée Royal de l'Afrique Centrale, Tervuren, Bel- gium]. Distribution.—Congo. Hosts.—Unknown. Likelihood of interception.—Very un- likely; known only from type material. Thrips incognitus Priesner, 1914 Thrips incognitus Priesner, 1914: 259. Type locality. Austria: Umgebung Graz (Sterermark) [Priesner Coll., Linz, Aus- trial]. Distribution.—Austria, Czech Republic, Hungary, Poland, Romania, Slovakia, Ukraine. Hosts.—Galium cruciatum, grass. Likelihood of interception.—Very un- likely; seldom collected in nature. Thrips inopinatus zur Strassen, 1963 Thrips inopinatus zur Strassen, 1963: 523. Type locality. Germany: Frankfurt am Main, Sandgebiet von Schwanheim [Senckenberg Museum, Frankfurt am Main]. Distribution.—England, Germany, Neth- erlands. Hosts.—Solanum dulcamara. Likelihood of interception.—Very un- likely; seldom collected. Thrips italicus (Bagnall, 1926) Taeniothrips italicus Bagnall, 1926: 650. Type locality. Italy: Portici (Napoli) [The Natural History Museum, London]. Thrips italicus: Revalid status, zur Strassen 1995S. Synonyms: Euthrips annulata Karny, 1907; Taeniothrips sodalis Bagnall, 1996; Tae- niothrips arcangelii Cappelletto, 1933. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Distribution.—Albania, England, France, Italy, Turkey, Ukraine, former Yugoslavia. Hosts.—Bellis sp.; Citrullus vulgaris; Er- ica verticillata; Euphorbia wulfenti; Ficus sp.; Genista sp.; Photinia elliptica; Quercus ilex; Ranunculus sp.; Robinia sp.; Scabiosa sp.; Solanum dulcamara. Spartium sp.; Spasticum sp. Likelihood of interception.—Probable; occasionally intercepted from various com- modities. Thrips juniperinus Linnaeus, 1758 Thrips juniperinus Linnaeus, 1758: 457. Type locality. Not specified [Unknown]. Synonyms: Thrips junipericola Morison, 1949; Thrips carpathicus Knechtel, 1948. Distribution.—Austria, Czech Republic, England, France, Germany, Romania, Scot- land, Slovakia, Transcaucasia. Hosts.—Juniperus communis, J. nana, J. sp. Likelihood of interception.—Unlikely; limited host range, although commonly col- lected in Europe. Thrips kikuyuensis Trybom, 1911 Thrips kikuyuensis Trybom, 1911: 6. Type locality. British East Africa: Escarpment [Stockholm Museum]. Synonyms: Thrips spadix var. brevipes Hood, 1932. Distribution.—Cameroon, Congo, Ken- ya, Tanzania, Uganda. Hosts.—Coffea arabica, C. sp., Ipomoea sp., Lablab niger, Mangifera indica, Musa sapientum, yellow Eupatorium-like flowers, small thistle sp., plaintain. Likelihood of interception.—Unlkely; although found on some commodities, this species has not as yet been intercepted in U.S. ports. Thrips klapeleki Uzel, 1895 Thrips klapeleki Uzel, 1895: 203. Type lo- cality. Bohemia [Vienna Museum?]. Distribution.—(Europe) Austria, Bel- gium, Czech Republic, England, France, VOLUME 108, NUMBER 2 Germany, Italy, Netherlands, Scotland, Slo- vakia. (Asia) Transcaucasia. Hosts.—Dactylorchis fuchsii fuchsii, D. incarnata incarnata, D. maculata ericeto- rum, Gymnadena conopsea, Orchis incar- nata, O. maculata, O. morio, O. sambucina. Likelihood of interception.—Unlikely; not common, and as yet has not been inter- cepted in U.S. ports. Thrips leucadophilus Priesner, 1936 Thrips leucadophilus Priesner, 1936: 91. Type locality. Sudan: Ghubshan [Priesner Coll.?, Linz, Austria]. Distribution.—Sudan. Hosts.—Leucas nubica. Likelihood of interception.—Very un- likely; known only from type material. Thrips lini Ladureau, 1878 Thrips lini Ladureau, 1878: 953. Type lo- cality. France [Unknown]. Synonyms: Thrips linarius Uzel, 1895. Distribution.—(Europe) Austria, Czech Republic, France, Germany, Italy, Nether- lands, Poland, Romania, Russia, Scotland, Serbia, Slovakia, Ukraine. (Africa) North Africa. (Asia) Korea. Hosts.—Pest on Linum occasionally on other plants. Likelihood of interception.—Possible; has a broad distribution and is found on several commodities; a pest on Linum usi- tatissimum. usitatissimum, Thrips lividus Haliday, 1836 Thrips lividus Haliday, 1836: 449. Type lo- cality. Britain [Unknown]. Distribution.—Britain. Hosts.—Ulex europaea. Likelihood of interception.—Very un- likely; rare in collections. Thrips major Uzel, 1895 Thrips major Uzel, 1895: 179. Type local- ity. Bohemia [Vienna Museum’?]. Synonyms: Thrips major var. adusta Uzel, 455 1895; Thrips major var. gracilicornis Uzel, 1895; Thrips major var. corticina Priesner, 1925; Thrips major var. saro- thamni Priesner, 1927; Thrips major vat. banatica Knechtel (7. litt.) Priesner, 1927; Thrips major var. banatica f. dorsima- culata Priesner, 1927; Thrips major vat. banatica f. ustulata Priesner, 1927; Phy- sothrips rosaceae Moulton, 1936. Distribution.—(Europe) Albania, Aus- tria, Czech Republic, Dalmatia, Denmark, England, Estonia, Finland, Germany, Hun- gary, Ireland, Italy, Lithuania, Poland, Ro- mania, Russia, Scotland, Slovakia, Sweden, Transcaucasia, Wales. (Mediterranean) Cy- prus, Palestine (= Israel). (Africa) Algeria, Canary Islands, Madeira, Mallorca, Moroc- co. (Asia) Mongolia. Hosts.—Large variety of flowers. Likelihood of interception.—Very prob- able; a common species frequently inter- cepted in ports on a variety of hosts. Thrips mancosetosus Titschack, 1954 Priesneria mancosetosus Maltbaek, 1928: 11. Nom. nud. Thrips mancosetosus (Maltbaek) Titschack, 1954: 344. Type locality. Denmark”: Haderslev [Unknown]. Distribution.—Austria, Denmark, Ger- many. Hosts.—Cirsium oleraceum, Gentiana germanica. Likelihood of interception.—Unlikely: Seldom collected. Thrips mareoticus (Priesner 1932) Stenothrips mareoticus Priesner, 1932: 10. Type locality. Egypt: Mariut [Priesner Coll., Linz, Austria]. Thrips mareoticus (Priesner): Speyer, 1935: 49. Synonyms: Thrips quadrisetosus Hood, 1932. Distribution.—(Europe) Italy. (Mediter- ranean) Cyprus, Palestine (= Israel), Tur- key. (Africa) Egypt, Morocco. 456 Hosts.—Achillea santolina, Chrysanthe- mum coronarium, C. macrocarpum = au- reum, C. segetum, Ferula communis typica, Matricaria sp., Plantago sp., Reichardia tingitana, Olive, peach, yellow marigold. Likelihood of interception.—Unlikely; although more broadly distributed and with a greater variety of hosts than some species, this species is uncommon. Thrips mariae Cott, 1924 Thrips mariae Cotti, 1924: 2. Type locality. France: Cimiez (Nice) [Unknown]. Distribution.—France. Hosts.—Celtis australis. Likelihood of interception.—Very un- likely; rare, known only from type material. Thrips medialis Oettingen, 1951 Thrips medialis Oettingen, 1951: 55. Type locality. Not specified [Unknown]. Distribution.—Poland, Lithuania. Hosts.—Plants in meadows. Likelihood of interception.—Very un- likely; limited distribution, uncommon, not found on economically important hosts. Thrips mediterraneus Priesner, 1934 Thrips mediterraneus Priesner, 1934: 276. Type locality. Egypt [Priesner Coll., Linz, Austria]. Distribution.—Egypt. Hosts.—Phlomis floccosa, barley. Likelihood of interception.—Very un- likely; limited distribution, uncommon, not found on economically important hosts. Thrips menyanthidis Bagnall, 1923 Thrips menyanthidis Bagnall, 1923: 126. Type locality. England: Patterdale, West- morland [The Natural History Museum, London]. Distribution.—Albania, Czech Republic, England, Estonia, Finland, Hungary, Slo- vakia. Hosts.—Menyanthes trifoliata, Passiflo- ra sp., Pedicularis palustris. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Likelihood of interception.—Unlhkely; seldom encountered in nature. Thrips meridionalis (Priesner, 1926) Taeniothrips meridionalis Priesner, 1926: 301. Type locality. Unknown [Priesner Coll., Linz, Austria]. Thrips meridionalis (Priesner): Bhatti 1978: 301. Distribution.—(Africa) African Mediter- ranean coast. (Europe) Albania, France, Georgia, Greece, Italy, Romania, Russia, Turkey, former Yugoslavia. (Mediterra- nean) Cyprus, Palestine (Israel), Iran, Iraq, Lebanon. Hosts.—Achillea sp.; Anthemis maritima; Atriplex portulacoides; Brassica sp.; Citrus sp.; Erica verticillata; Fontanesia_ philly- reoides;, Frankenia laevis; Fraxinus ornus; Helichrysum arenarium; Tris germanica; Matthiola sinuata; Prangos ferulacea; Ra- phanus sp.; Robinia pseudoacacia; Rosa sp.; Sambucus ebulus; Scilla sp.; Sinapis sp.; Styrax officinalis; Tamarix sp.; Ulex sp.; Viburnum sp.; Gramineae. Likelihood of interception.—Very prob- able; often intercepted. Thrips microchaetus Karny, 1920 Thrips microchaetus Karny, 1920: 27. Type locality. Anglo-Egyptian Sudan: Kororak [Unknown]. Distribution.—Egypt, Kenya, Sudan. Hosts.—Aerva tomentosa, Coffea arabi- ca, Nitraria sp., Pulicaria crispa, Triumfet- ta flavescens, grape vine, yellow compos- ites. Likelihood of interception.—Unlikely; seldom encountered species; litthe com- merce with countries within range. Thrips minutissimus Linnaeus, 1758 Thrips minutissimus Linnaeus, 1758: 457. Type locality. Europe [Unknown]. Synonyms: Bagnallia variabilis Williams, 1913; Thrips minutissimus var. obscura Coesfeld, 1898; Thrips evestigatus Oet- tingen, 1944. VOLUME 108, NUMBER 2 Distribution.—(Europe) Austria, Czech Republic, Denmark, England, Finland, France, Germany, Hungary, Italy, Lithua- nia, Netherlands, Poland, Romania, Scot- land, Serbia, Slovakia, Sweden, Switzer- land, Wales. (Mediterranean) Cyprus, Cri- mea, Palestine (= Israel). Hosts.—various flowers, young leaves. Likelihood of interception.—Possible; broad distribution and found on a variety of hosts; has been encountered numerous times at U.S. ports. buds, and Thrips montivagus Priesner, 1923 Thrips montivagus Priesner, 1923: 83. Type locality. Upper Austria: Dachsteingebier (nahe der Grobsteinhtitte) [Priesner Coll.?, Linz, Austria]. Distribution.—Austria, France, Romania, Switzerland, Ukraine. Hosts.—Found in alpine flowers. Likelihood of interception.—Very un- likely. Not found on any commodities. Thrips nigropilosus Uzel, 1895 Thrips nigropilosus Uzel, 1895: 198. Type locality. Bohemia [Vienna Museum’. Synonyms: Thrips lactucae Beach, 1896; Thrips nigropilosus f. pilosissimus Pries- nem 922: Distribution.—(Europe) Albania, Aus- tria, Czech Republic, Denmark, England, Finland, France, Germany, Hungary, Lith- uania, Netherlands, Poland, Romania, Rus- sia, Scotland, Slovakia, Sweden, Switzer- land, Turkey. (Africa) Egypt. (Asia) Japan, South Korea. (Oceana) Hawaii, Fiji. (Aus- tralasia) Australia. (North America) USA [California, Georgia, Illinois, Indiana, Iowa, New Jersey, New York, South Dakota, Vir- ginia, Washington]. Hosts.—Various flowers, mainly com- posites. Likelihood of interception.—Probable; Widely distributed and found on numerous flowers transported in commerce. 457 Thrips obscuricornis Priesner, 1920b Thrips obscuricornis Priesner, 1920b: 57. Type locality. Austria Ober6sterreich [Priesner Coll., Linz, Austria]. Distribution.—Austria, Denmark, Fin- land, Germany, Hungary, Netherlands, Ro- mania, Turkey. Hosts.—Carlina vulgaris, Centaurea sol- stitialis, C. sp., Cirsium erysithales, Orchis incarnata, Taraxacum officinale. Likelihood of interception.—Unlikely; uncommon. Thrips origani Priesner, 1926 Thrips origani Priesner, 1926: 272. Type lo- cality. Hungary [The Natural History Museum, London]. Distribution.—Austria, Czech Republic, England, France, Germany, Hungary, Ma- deira, Romania, Slovakia. Hosts.—Origanum vulgare. Likelihood of interception.—Very un- likely; uncommon. Thrips oryzophagus Rondani, 1871 Thrips oryzophagus Rondani, 1871: 29. Type locality. Italy [Unknown]. Distribution.—Italy. Hosts.—Rice. Likelihood of interception.—Very un- likely; uncommon. Thrips palmi Karny, 1925 Thrips palmi Karny, 1925: 54. Type local- ity. Indonesia [Unknown]. Synonyms: Thrips clarus Moulton, 1928a; Thrips leucadophilus Priesner, 1936; Chloethrips aureus Ananthakrishnan and Jagadish, 1967; Thrips gracilis Anan- thakrishnan and Jagadish, 1968. Distribution.—(Africa) Sudan. (Asia) Bangladesh, Hong Kong, India, Indonesia, Japan, Malaysia, Pakistan, People’s Repub- lic of China, Philippines, Singapore, Tai- wan, Thailand. (Australasia) Guam, New Caledonia, Samoa, Wallis Island. (North America) Antigua, Barbados, Dominican 458 Republic, Guadeloupe, Martinique, Puerto Rico, Saint Lucia, Trinidad, USA [Florida]. (South America) Venezuela. Hosts.—Brassica campestris, B. Juncea, Camellia thea, Citrus medica, Cymbidium bicolor, Cypripedium sp., Dendrobium gra- tiosissimus, Fabia vulgaris, Goodlaea sp., Gossypium herbaceum, Habernaria sp., Lathyrus sp., Pyrus malus, Sesamum indi- cum, Striga sp. Likelihood of interception.—Probable, but not from Europe, Mediterranean or Af- rican regions; may be intercepted in mate- rial channeled into Netherlands hub. Thrips paludosus Bagnall, 1913 Thrips paludosus Bagnall, 1913: 235.Type locality. England: Weston-on-the Green, Oxfordshire [The Natural History Muse- um, London]. Distribution.—Czech Republic, England, Poland, Slovakia. Hosts.—Erica tetralix, Fagus silvatica, sedge. Likelihood of interception.—Unlikely; not common and not found on commodities being transported from Europe. Thrips paluster Reuter, 1899 Thrips paluster Reuter, 1899: 58. Type lo- cality. Finland: Munduksa [Helsinki Mu- seum]. Distribution.—England, Finland, Mora- via, Norway, Ukraine. Hosts.—Pedicularis palustris, marsh plants. Likelihood of interception.—Very un- likely; from a region not usually involved in commerce with U.S. and not found on any commodities. some Thrips panousei zur Strassen, 1968 Thrips panousei zur Strassen, 1968: 55. Type locality. Southern Morocco: Aouinet-Torkoz (Cercle de Goulimi- ne)[Senckenberg Museum, Frankfort am Main]. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Distribution.—Morocco. Hosts.—Ephedra alata alenda, Pituran- thus chloranthus. Likelihood of interception.—Very un- likely; from a region not usually involved in commerce with U.S. and not found on any commodities. Thrips paucisetosus Priesner, 1927 Thrips paucisetosus Priesner, 1927: 57. Type locality. Not specified [Priesner Coll., Linz, Austria]. Distribution.—Austria, Czech Republic, Denmark, Finland, Hungary, Slovakia. Hosts.—Echium sp., Centaurea sp. Likelihood of interception.—Very un- likely; from a region not usually involved in commerce with U.S. and not found on any commodities. Thrips pelikani Titschack, 1962 Thrips pelikani Titschack, 1962: 24. Type locality. Greece [Type Lost?]. Distribution.—Greece. Hosts.—Hieracium sp. Likelihood of interception.—Very un- likely; from a region not usually involved in commerce with U.S. and not found on any commodities. Thrips pennatus zur Strassen, 1968 Thrips pennatus zur Strassen, 1968: 2. Type locality. Madeira: Rabagal [Senckenberg Museum, Frankfort am Main]. Distribution.—Madeira. Hosts.—Leontodon sp. Likelihood of interception.—Very un- likely; from a region not usually involved in commerce with U.S. and not found on any commodities. Thrips persicae Haliday, 1836 Thrips persicae Haliday, 1836: 146. Type locality. Britain [Unknown]. Distribution.—Britain. Hosts.—Peach. VOLUME 108, NUMBER 2 Likelihood of interception.—Very un- likely; rare. Thrips physapus Linnaeus, 1758 Thrips physapus Linnaeus, 1758: 457. Type locality. Not specified [Unknown]. Synonyms: Thrips fusca Miiller, 1776; Thrips flavicornis Reuter, 1879; Thrips physapus var. adusta Uzel, 1895; Thrips f. annulata Karny, 1907; Thrips physapus var. flavescens Priesner, 1919; Thrips physapus var. quadrisetosus Priesner, 1923; Thrips physapus f. brunnea Ishida, 1936; Thrips collinus Oettingen, 1951. Distribution.—(Europe) Albania, Aus- tria, Czech Republic, Denmark, England, Estonia, Finland, France, Germany, Greece, Hungary, Ireland, Italy, Lithuania, Nether- lands, Norway, Poland, Romania, Russia, Scotland, Serbia, Slovakia, Sweden, Swit- zerland, Turkey, Wales, former Yugoslavia. (Mediterranean) Persia (Iraq, Iran). (Africa) Sudan, Morocco. (Asia) Japan. (North America) Canada (British Columbia), USA [Hlinois, Michigan, New Jersey, New York, Utah]. Hosts.—Numerous flowers, especially composites. Likelihood of interception.—Probable; intercepted frequently on a variety of flow- ers. Thrips phytolaccae Priesner, 1951 Thrips phytolaccae Priesner, 1951: 256. Type locality. Turkey: Diizce [Unknown]. Distribution.—Turkey. Hosts.—Glycyrrhiza sp., Phytolacca americana, P. sp. Likelihood of interception.—Very un- likely; limited distribution and no econom- ically important hosts. Thrips pillichi Priesner, 1924 Thrips pillichi Priesner, 1924: 2. Type lo- cality. Hungary: Simontornya [Priesner Coll., Linz, Austria]. Synonyms: Thrips f. fallaciosa_ Priesner, 459 1924; Thrips f. hiemalis Priesner, 1925; Thrips kerschneri Priesner, 1927. Distribution.—Austria, Czech Republic, England, France, Germany, Hungary, Neth- erlands, Norway, Romania, Spain, Turkey, Ukraine. Hosts.—Achillea_ millefolium, Alopecu- rus pratensis, Anthemis arvensis, Arrhen- atherum elatius, Bidens tripartitus, Camilla sp., Centaurea jacea f. pannonica, Ceras- tium tomentosum, Chrysanthemum leucan- themum, C. vulgare, C. sp., Cichorium in- tybus, Cuscuta europaea, Euphorbia cypar- issias, E. palustris, Hieracium umbellatum, Isatis tinctoria, Lotus corniculatus, Matri- caria chamomilla, Medicago sativa, Ranun- culus sp., Senecio jacobaea, S. sylvaticus, Solidago virgaurea, Sonchus palustris, Sor- bus torminalis, Tanacetum vulgare, Vicia cracca. Likelihood of interception.—Probable. Although seldom intercepted at U.S. ports, this widely distributed species found on many hosts is likely to be encountered at U.S. ports. Thrips pilosissimus Priesner, 1922 thrips prlosissmmus Pricsner, 1922--92: Type locality. Not specified [Priesner Coll.?, Linz, Austria]. Distribution.—(Europe) Austria, Germa- ny, Hungary, Ukraine. (North America) USA [Colorado]. Hosts.—Chrysanthemum spp., Hiera- cium pilosella, turf grass. Likelihood of interception.—Unlikely; seldom collected. Thrips ponticus Knechtel, 1965 Thrips ponticus Knechtel, 1965: 134. Type locality. Rumania: Rayon Medgidia, Re- gion Dobrogea [Unknown]. Distribution.—Romania. Hosts.—Achillea setacea, Brassica rapa, Carduus leiophyllus, Chrysanthemum mil- lefolium, Conium maculatum, Marrubium peregrinum, Matricaria chamomilla. 460 Likelihood of interception.—Very un- likely; limited distribution, seldom collect- ed: Thrips poultoni Bagnall, 1933 Thrips poultoni Bagnall, 1933: 656. Type locality. Canary Islands: Las Palmas [The Natural History Museum, London]. Synonyms: Thrips canariensis Priesner, 1933: Distribution.—Canary Islands, southern Morocco. Hosts.—Polyphagous. Likelihood of interception.—Unlikely; little commerce with countries within dis- tribution of this species. Thrips praetermissus Priesner, 1920b Thrips praetermissus Priesner, 1920b: 58. Type locality. Austria: Klaus, Ober6ster- reich [National Museum of Natural His- tory, Washington, DC]. Distribution.—(Europe) Austria, Czech Republic, England, Slovakia, Ukraine. (Asia). Mongolia. Hosts.—Forest turf, flowers mainly of composites. Likelihood of interception.—Unlikely; seldom collected. Thrips priesneri Hood, 1932 Thrips priesneri Hood, 1932: 136. Type lo- cality. Cameroons [National Museum of Natural History, Washington, DC]. Distribution.—Cameroon. Hosts.—Coffea robusta, Melia azedar- ach, flowers of an unidentified shrub. Likelihood of interception—Very un- likely; rare in collections. Thrips pusillus Bagnall, 1926 Thrips pusillus Bagnall, 1926: 112. Type locality. W. Africa: Gold Coast, Aburi [The Natural History Museum, London]. Synonyms: Thrips meliaefloris Hood, 1932 Distribution.—Ghana, Nigeria. Hosts.—Canna sp., Coffea robusta, Me- lia azedarach, Strophanthus gratus, lemon. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Likelihood of interception.—Unlhkely; seldom collected. Thrips quadridentatus Trybom, 1910 Thrips quadridentatus Trybom, 1910: 164. Type locality. Kalahari: Pfanne Kooa [Berlin Museum? ]. Distribution.—South Africa. Hosts.—Acacia karroo. Likelihood of interception.—Very un- likely: Limited distribution and not found on any commodity. Thrips ranunculi Schrank, 1781 Thrips ranunculi Schrank, 1781: 299. Type locality. Austria [Unknown]. Distribution.—Austria. Hosts.—Ranunculus sp. (Gmplied from name). Likelihood of interception.—Very un- likely; known only from type material. Thrips robustus Priesner, 1920a Thrips robustus Priesner, 1920a: 76. Type locality. Austria [Priesner Coll., Linz. Austria]. Synonyms: Thrips robustus var. pyrenaica Bagnall, 1926. Distribution.—Andorra, Austria, Finland, France, Germany (?), Italy, Romania, Spain, Ukraine. Hosts.—Campanula sp., Cerastium sp., Gentiana austriaca, G. clusii, G. kochiana, G. lutea, G. nana, G. pyrenaica, G. verna, Pedicularis palustris. Likelihood of interception.—Unlkely; not common. Thrips roepkei Doeksen, 1953 Thrips roepkei Doeksen, 1953: 169. Type locality. Holland: Scheveningen [Un- known]. Distribution.—Netherlands. Hosts.—Solanum nigrum. Likelihood of interception.—Very un- likely; known only from type material. VOLUME 108, NUMBER 2 Thrips sambuci Heeger, 1854 Thrips sambuci Heeger, 1854: 215. Type lo- cality. Austria .[Vienna Museum?]. Synonyms: Thrips nigra Williams, 1916. Distribution.—Austria, Czech Republic, Denmark, England, Finland, France, Ger- many, Hungary, Italy, Lithuania, Nether- lands, Poland, Romania, Scotland, Slovak- ia, Switzerland, Ukraine. Hosts.—Cornus sanguinea, Sambucus nigra, S. racemosa. Likelihood of interception.—Unlikely; seldom collected. Thrips simplex (Morison, 1930) Physothrips simplex Morison, 1930: 12. Type locality. Not specified [Unknown]. Taeniothrips simplex (Morison): Steele, 19352 33: Thrips simplex (Morison): Bhatti, 380. Synonyms: Taeniothrips gladioli Moulton and Steinweden, 1931; Physothrips spi- ranthidis plurisetae Girault, 1933. 1969: Distribution.—(Europe) Albania, Aus- tria, Czech Republic, Denmark, England, Estonia, Finland, France, Germany, Greece, Hungary, Ireland, Italy, Lithuania, Nether- lands, Norway, Poland, Romania, Russia, Scotland, Serbia, Slovakia, Sweden, Swit- zerland, Turkey, Wales, former Yugoslavia. (Africa) Sudan, Morocco. (Asia) Japan, Persia [Iraq, Iran]. (North America) Canada [British Columbia], USA [Illinois, Michi- gan, New Jersey, New York, Utah]. Hosts.—Numerous flowers, especially composites. Likelihood of interception.—Probable; intercepted frequently on a variety of flow- ers. Thrips spadix Hood, 1932 Thrips spadix Hood, 1932: 131. Type lo- cality. Cameroons [National Museum of Natural History, Washington, DC]. Distribution.—Cameroon, Congo. Hosts.—Unidentified shrub, unidentified small weed, unidentified small thistle. 461 Likelihood of interception.—Very un- likely; rare, from countries seldom involved in commerce with U.S. Thrips spoliatus Priesner, 1964 Thrips spoliatus Priesner, 1964: 98. Type locality. Hungary [Priesner Coll.?, Linz, Austria]. Distribution.—Hungary. Hosts.—Forest turf. Likelihood of interception.—Very un- likely; known only from type material. Thrips tabaci Lindeman, 1889 Thrips tabaci Lindeman, 1889: 72. Type lo- cality. Basarabia [Unknown]. Synonyms: 7. adamsoni Bagnall, 1923; T. communis Uzel, 1895; T. flava var. ob- soleta Uzel, 1895; T. bremnerii Moulton, 1907; 7. dorsalis Bagnall, 1927; T. hol- oleucus Bagnall, 1914; Limothrips allii Gillette, 1893; T. bicolor Karny, 1907; T. communis var. pulla Uzel, 1895; Parath- rips uzeli Karny, 1907; T. debilis Bagnall, 1923; T. frankeniae Bagnall, 1926; T. seminiveus Girault, 1926; 7. shakespearei Girault, 1926; T. indigenus Girault, 1929. Distribution.—Cosmopolitan. Hosts.—Large number of plants; pest of onions, tobacco, cotton, vector of tomato spotted wilt virus and related forms. Likelihood of interception.—Extremely high; this is one of the most frequently in- tercepted species in U.S. ports and is found on a wide variety of hosts. Thrips tarfayensis zur Strassen, 1968 Thrips tarfayensis zur Strassen, 1968: 59. Type locality. Southern Morocco: Hassi Zehar (Prov. Tarfaya) [Unknown]. Distribution.—Morocco. Hosts.—Pulicaria crispa. Likelihood of interception—Very un- likely; known only from type material. Thrips tenellus Trybom, 1912 Thrips tenellus Trybom, 1912: 6. Type lo- cality. South Africa: Stanford (sic) Hill, 462 Natal [Naturhistoriska Museum, Géote- borg, Sweden]. Distribution.—South Africa. Hosts.—Nephelium litchi; under a stone. Likelihood of interception.—Very un- likely; known only from type material. Thrips tenuisetosus Knechtel, 1923 Thrips tenuisetosus Knechtel, 1923: 203. Type locality. Rumania: Chisinau [Kne- chtel Coll.?]. Distribution.—France, Romania, Turkey, Ukraine. Hosts.—Bellis perennis, Bupleurum sp.?, Isatis glauca (or I. tinctoria?), Lamium pur- pureum, Senecio vulgaris, Taraxacum dens- leonis, Senecio vulgaris, Cruciferae. Likelihood of interception.—Unlikely; an uncommon species. Thrips timidus Priesner, 1926 Thrips timidus Priesner, 1926: 267. Type lo- cality. Hungary: Simontornya [Priesner Coll., Linz, Austria]. Distribution.—Germany, Ukraine. Hosts.—Stachys recta, turf. Likelihood of interception.—Unlikely; an uncommon species not found on com- modities. Hungary, Thrips trehernei Priesner, 1927 Thrips trehernei Priesner, 1927: 356. Type locality. Canada [Priesner Coll., Linz, Austria]. Synonyms: Thrips taraxaci Moulton, 1936b; Thrips hukkineni Priesner, 1937. Distribution.—(Europe) Austria, Czech Republic, England, Estonia, Finland, France, Germany, Greece, Hungary, Lithu- ania, Poland, Russia, Slovakia, Sweden, Turkey. (Asia) Mongolia. (North America) Canada (British Columbia, Ontario), USA [California, Colorado, Connecticut, District of Columbia, Georgia, Idaho, [linois, In- diana, Iowa, Kansas, Maine, Maryland, Michigan, Missouri, Montana, Nebraska, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON New Jersey, New York, Ohio, Oregon, Utah, Virginia, Washington, Wisconsin, West Virginia, Wyoming]. (Oceania) Ha- wail. Hosts.—Various flowers, especially com- posites. Likelihood of interception.—Probable; though not commonly intercepted, this spe- cies is routinely found on a wide variety of flowers and has been intercepted at various U.S. ports recently. Thrips urticae Fabricius, 1781 Thrips urticae Fabricius, 1781: 397. Type locality. Not specified [Lost?]. Distribution.—(Europe) Austria, Bessa- rabia, Czech Republic, Denmark, England, Finland, Germany, Hungary, Poland, Ro- mania, Slovakia, Switzerland, Ukraine. (Asia) Manchuria. Hosts.—Urtica dioica, other plants in lit- erature probably casual. Likelihood of interception.—Unlikely; though widely distributed in Europe, this species has not as yet been encountered at U.S. ports. Thrips uzelianus Priesner, 1926 Thrips uzelianus Priesner, 1926: 273. Type locality. Hungary: Simontornya. [Pries- ner Coll., Linz, Austria]. Distribution.—Czech Republic, Hunga- ry, Slovakia, Switzerland, Ukraine. Hosts.—Urtica dioica, Helichrysum ar- enarium. Likelihood of interception.—Unlikely. Thrips validus Uzel, 1895 Thrips validus Uzel, 1895: 183. Type lo- cality. Bohemia [Vienna Museum? ]. Synonyms: Thrips fuscaMoulton, 1936b; Thrips adusta Uzel, 1895; Thrips adusta var. nigra Uzel, 1895; Thrips longicollis Uzel, 1895; Thrips collinus Oettingen, ns Syle Distribution.—(Europe) Austria, Czech Republic, Denmark, England, Estonia, Fin- VOLUME 108, NUMBER 2 land, France, Germany, Hungary, Italy, Lithuania, Netherlands, Poland, Romania, Russia, Serbia, Slovakia, Sweden, Switzer- land, Ukraine, former Yugoslavia. (Asia) Manchuria. (North America) U.S.A. (Ida- ho). Hosts.—Flowers, especially composites. Likelihood of interception.—Unlikely; not commonly collected. Thrips variegatus von Gleichen, 1764 Thrips variegatus von Gleichen, 1764: 22. Type locality. Not specified [Unknown]. Distribution.—Germany. Hosts.—Linaria sp. Likelihood of interception.—Very un- likely; rare. Thrips viminalis Uzel, 1895 Thrips viminalis Uzel, 1895: 182. Type lo- cality. Bohemia [Vienna Museum’. Distribution.—Albania, Austria, Czech Republic, Dalmatia, England, Finland, France, Georgia, Germany, Hungary, Italy, Netherlands, Poland, Romania, Russia, Ser- bia, Slovakia, Sweden, Switzerland, Ukraine. Hosts.—Salix spp; occasionally on other trees and shrubs. Likelihood of interception.—Unlikely; an uncommon species. Thrips vulgatissimus Haliday, 1836 Thrips vulgatissima Haliday, 1836: 447. Type locality. Bohemia [Vienna Muse- um?]. Synonyms: Euthrips alpina Karny, 1908; Taeniothrips lemansis Treherne, 1924; Physopus pallipennis Uzel, 1895; Tae- niothrips vulgatissimus americanus Moulton, 1929; Taeniothrips vulgatissi- mus atricornis Priesner, 1926. Distribution.—Austria, Czech Republic, England, Finland, Denmark, France, Geor- gia, Germany, Greece, Hungary, Iceland, Ireland, Italy, Netherlands, Poland, Roma- nia, Russia, Scotland, Slovakia, Spain, 463 Sweden, Switzerland, former Yugoslavia. (North America) Canada (Alberta, British Columbia, Manitoba, New Brunswick, Newfoundland, Northwest Territories, Nova Scotia, Ontario, Quebec, Saskatchewan, Yukon Territory), Greenland, U.S.A. (Alas- ka, Arizona, California, Colorado, Georgia, Idaho, Illinois, Maine, Montana, New Jer- sey, New Mexico, Nevada, Oregon, South Dakota, Utah, Washington, Wyoming). Hosts.—From many varieties of plants. Likelihood of interception.—Very Prob- able; a commonly intercepted species. ACKNOWLEDGMENTS I express sincere appreciation to Joseph Cavey and Peter Touhey, USDA/APHIS, Hyattsville, MD, for providing information for the databases, Sueo Nakahara (retired) and Douglass R. Miller, SEL, USDA, and Susan Broda, USDA/APHIS, Baltimore, MD, for their comments in reviews of the manuscript. LITERATURE CITED Ananthakrishnan, T. N. and A. Jagadish. 1967. Studies on the genus Chloethrips Priesner from India. Zoologischer Anzeiger 178: 375-388. . 1968. Studies of species of the genus Thrips from India II. Deutsche Entomologische Zeit- schrift 15: 359-365. Bagnall, R. S. 1913. Brief descriptions of new Thy- sanoptera. Annals and Magazine of Natural His- tory (12)8: 290-299. . 1915. Brief descriptions of new Thysanoptera. Annals and Magazine of Natural History (15)8: 588-597. . 1916. Brief descriptions of new Thysanoptera. VII. Annals and Magazine of Natural History. Se- ries 8, 17: 213—223. . 1923. 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Commonwealth of Australia Coun- cil of Scientific and Industrial Research Pamphlet 54, 59 pp. Titschack, E. 1962. Thysanoptera XV. Neufunde aus unserum Arbeitsgebiet. Bombus 97: 24. Treherne, R. C. 1924. Thysanoptera known to occur in Canada. Canadian Entomologist 56: 82-88. Trybom, E 1910. Ergebnisse einer Forschungsreise im westliche und zentralen Siid Afrika. Denkschriften der Medicinisch-Naturwissenschaftliche Gesell- schaft zu Jena 16(4): 164, Pl. IIc, figs. 28, 29. 1911. Physapoden, Gesemmelt von der Schwedischen Expedition nach dem Brittischen Ostafrika. Arkiv for Zoologi 7: 1—12. . 1912. Physapoden aus Natal und dem Zulu- lande. Arkiy f6r Zoologi 7: 1—59. Uzel, H. 1895. Monographie der Ordnung Thysanop- tera. K6niggratz, BhGmen. 472 pp. Von Gleichen, X. 1764. Das Neueste aus dem Reiche de Pflanzen, p. 22, Pl. 21, figs. 6, 7. Williams, C. B. 1913. Records and descriptions of British Thysanoptera. Journal of Economic Biol- ogy 8: 216—230. . 1916. Biological and systematic notes on Brit- ish Thysanoptera. Entomologist 49: 221—227, 243—245, 275-284. Zur Strassen, R. 1963. Thrips inopinatus n. sp. aus Deutschland. Senckbergischen Biologie 44: 523— ile Zur Strassen, R. 1968. Okologische und zoogeogra- phische Studien tiber die Fransenfliigler-Fauna (Ins., Thysanoptera) des stidliche Marokko. Ab- handlungem der Senckbergischen Naturforschen- den Gesellschaft 515: 1-125. Zur Strassen, R. 1995. Taxonomische Neurungen bei terebranten Fransenfliiglern der westlichen Pa- laarktis (Thysanoptera: Terebrantia: Aeolothripi- dae, Thripidae). Mitteilung international Entomo- logische Verhandlung, Frankfort am Main 20: 87— 102. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 467-473 A NEW SPECIES OF THE BITING MIDGE GENUS DASYHELEA KIEFFER (DIPTERA: CERATOPOGONIDAE) FROM THE BAHAMAS WILLIAM L. GROGAN, JR. AND JULIA A. WIENERS Department of Biological Sciences, Salisbury University, Salisbury, MD 21801, U.S.A. (e-mail: wlgrogan @salisbury.edu) Abstract.—Dasyhelea morrisoni Grogan and Wieners, new species, is described and illustrated from small islets near Staniel Cay, Exuma Cays, Bahamas, and compared with other similar congeners. This small, new species is most closely related to the Nearctic species, D. brevicornis Waugh and Wirth and together they form the Dasyhelea brevi- cornis group (new species group). Key Words: The biting midges of the genus Dasyhe- lea Kieffer are typically small to very small flies that inhabit a wide variety of wet hab- itats worldwide except for the Antarctic re- gion. Ten extinct species and nearly 470 ex- tant species have been described (Borkent and Wirth 1997), but, undoubtedly, many more species still await discovery. Despite their common name, females of all species of Dasyhelea have reduced, vestigial, ma- lelike mandibles without teeth and do not take blood meals, but, like their male coun- terparts, they obtain sugar and water from nectar of flowers (Downes 1969). The lar- vae are aquatic and inhabit a variety of freshwater ecosystems where they usually graze on algae and morphologically resem- ble the larvae of many species of Chiron- omidae. Through the kindness of Lloyd Morrison (the Center for Medical, Agricultural, and Veterinary Entomology, USDA-ARS, Gainesville, FL; current address: U.S Na- tional Park Service, Department of Biology, Missouri State University, Springfield, MO), we have had the opportunity to ex- amine several species of small Diptera that he collected with shallow pan traps from Diptera, Ceratopogonidae, Dasyhelea, biting midges, new species, Bahamas several very small, low islets near Staniel Cay, Exuma Cays, Bahamas. Morrison con- ducted biogeographical and ecological stud- ies of selected flora and fauna in the Ba- hamas for several years (e.g., Morrison 1997, 1998a, b, 2002a, b, 2003), but only recently collected these flies during field work in December 2002 and May 2003. Preliminary examination of these dipterans revealed three species of Chironomidae, the largest of which we tentatively identified as males of Clunio marshalli Stone and Wirth, a species that was originally described from Florida (Stone and Wirth 1947). Subse- quently, we sent all of these chironomids to Peter Cranston, who (in Jitt.) confirmed our earlier identification of the males of C. mar- shalli and determined that the other speci- mens belonged to two different genera. One specimen was a female of Chironomus Meigen that was too badly damaged to identify to species, whereas the other spec- imens Cranston identified as Mesosmittia sp. The most common flies collected by Morrison were a very small species of Das- yhelea, and, after comparing these to other congeners from the New World, we con- cluded that it is undescribed. Herein, we de- 468 scribe and illustrate adults of both sexes of this previously unknown species and pro- vide details about its habitat on the small islands from which it was collected as well as the larger, surrounding islands in the Ex- uma Cays, Bahamas. For general terminology of Ceratopogon- idae, see Downes and Wirth (1981); for an overview of North American Dasyhelea, see Waugh and Wirth (1976). Terms for general structures including genitalia and wing venation follow those in the Manual of Nearctic Diptera by McAlpine (1981) with recent modifications of wing veins proposed by Szadziewski (1996). All spec- imens examined were collected in 70% eth- anol and subsequently mounted onto micro- scope slides in phenol-balsam using the methods described by Wirth and Marston (1968). Measurements and other data are presented as mean, minimum-maximum values, followed by sample size. Dasyhelea morrisoni Grogan and Wieners, new species (Figs. 1-8) Diagnosis.—A small species of Dasyhe- lea of the brevicornis group that is distin- guished from all New World congeners by the following combination of characters: small size (male wing length 0.57—0.63 mm; female wing length 0.59-0.70 mm); short costa (costal ratio of males 0.44—0.48; of females 0.47—0.50); wing membrane with sparse macrotrichia, absent in cell cua, and anal cell of males and cell cua, of fe- males; halter brown; male genitalia very short with sternite 9 produced beyond apex of aedeagus; aedeagus crescent-shaped, very short, heavily sclerotized; female fla- gellum with flagellomeres 1—12 very short (antennal ratio 0.70—0.82), only flagello- mere 13 considerably longer than broad; and spermatheca single, very small, globu- lar with short neck. Male.—Head: Dark brown. Eyes pubes- cent, narrowly separated. Antennal flagel- lum (Fig. 1) with separate flagellomeres, all PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON lightly sculptured; flagellomeres 11—13 only slightly longer than flagellomere 10, antennal ratio 0.48 (0.42—0.51, n = 41); plume of sensilla chaetica moderately dense, extending beyond flagellomere 12. Palpus light brown, 5-segmented; segment 3 with 2—3 small capitate sensilla on mesal surface; palpal ratio 2.75 (2.33-3.40, n = 42). Mandible vestigial, without teeth. Tho- rax: Dark brown; scutellum light brown with 2 submarginal and 4 more centrally located setae. Legs dark brown, tarsi lighter brown, darkest on tarsomeres 5; claws small, bearing a short, basal seta, nearly straight proximally with bent, entire tips. Wing (Fig. 6) hyaline, membrane covered with dense microtrichia, macrotrichia sparsely scattered on distal half, along length of most veins and most of wing mar- gin, absent in cell cua, and anal cell; radial cells obliterated; base of vein M, not con- nected to M,. wing length 0.60 (0.57—0.63, n = 41) mm; costa short, costal ratio 0.46 (0.44—0.48, n = 41); anal lobe poorly de- veloped. Halter stem dark brown, knob light brown. Abdomen: Dark brown. Gen- italia as in Fig. 8. Sternite 9 produced slightly beyond apex of aedeagus, mem- brane non-spiculate; tergite 9 broad, taper- ing slightly distally to rounded caudal mar- gin, ventral portion of apex covered with fine setae, apicolateral processes short, rounded with single apical seta. Gonocoxite moderately short, not quite extending to ba- ses of apicolateral processes, surface with scattered large, coarse setae; gonostylus shorter than gonocoxite, tapering and curved on distal half to rounded tip. Ae- deagus very short, reduced to a narrow crescent-shape, heavily sclerotized. Para- meres separate; heavily sclerotized; basal arm straight; distal portion recurved with moderately slender, rounded tips. Female.—Slightly larger, similar to male with the following notable sexual differenc- es. Head: Antennal flagellum (Fig. 2) with only flagellomere 13 distinctly longer than proximal 12; flagellomeres with basal whorl of moderately long sensilla chaetica; sculp- VOLUME 108, NUMBER 2 469 f iy 4 a —— pl gg gn Ti 8 Figs. 1-8. Dasyhelea morrisoni. 1, Flagellum of male. 2, Flagellum of female. 3, Palpus of female. 4, Female genitalia. 5, Spermatheca. 6, Wing of male. 7, Wing of female. 8, Male genitalia. 470 turing present only on flagellomere 13, poorly developed; antennal ratio 0.76 (0.70—0.82, n = 31). Palpus (Fig. 3) with segment 3 moderately broad; palpal ratio 2.63 (2.17—3.00, n = 30). Thorax: Legs, including tarsi lighter brown, darkest on tar- someres 5; claws more curved than those of male, tips entire. Wing (Fig. 7) broader than male, with more macrotrichia on distal half of wing membrane and in anal cell; wing broader, slightly longer than male, wing length 0.63 (0.59—0.70, n = 30) mm; costa slightly longer than male, costal ratio 0.48 (0.47—0.50, n = 30). Abdomen: Ligh- ter brown. Genitalia as in Fig. 4. Sternite 8 bearing 4—5 large setae anterior to caudal margin, caudal margin with sclerotized M- shaped marking; sternite 9 entire, vaginal lumen ovoid; sternite 10 with pair of widely spaced sub-basal setae and posterior pair of narrowly separated setae. Spermatheca (Fig. 5) single, very small, globular with short neck; length (including neck) 0.029 (0.026—0.032, n = 30) mm, breadth 0.036 (0.023—0.029, n = 20) mm. Distribution.—Known only from the type locality, several small islets near Stan- iel Cay, Exuma Cays, Bahamas. Types.—Holotype 6; allotype @; para- types, 36 6, 25 2, Bahamas, small islands near Staniel Cay, Exuma Cays, 16-V-2003, L. Morrison, pan trap; 5 ¢, 5 2 paratypes with same data except collected 12-VII- 2002. The holotype, allotype and most paratypes are deposited in the National Mu- seum of Natural History, Washington, D. C. Other paratypes are deposited in the Cana- dian National Collection of Insects, Ottawa, Florida State Collection of Arthropods, Gainesville, Museo de La Plata, Argentina, Collection of Arthropods with Medical Im- portance of the Instituto Nacional of Diag- nostico and Referencia Epidemiologicos, Mexico City, Instituto Oswaldo Cruz, Rio de Janeiro, Brazil, and Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica. Etymology.—We are pleased to name this new species in honor of Lloyd Morri- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON son, who collected the entire type series on the small, low, rocky islands during field work in the Exuma Cays, Bahamas. Discussion.—This new species is appar- ently most closely related to D. brevicornis Waugh and Wirth (1976) from Maryland, New York, and Nova Scotia, the females of which also have an antennal flagellum with the proximal 12 flagellomeres very short with only flagellomere 13 being distinctly longer than broad. Females of D. brevicor- nis differ from those of D. morrisoni in be- ing slightly larger (wing length 0.74—0.87 mm), the wing membrane covered with dense macrotrichia that are present in all cells, the medial portion of the genital scler- otization on the caudal margin of sternite 9 is arrowhead-shaped and the spermatheca is larger with a very long, slender neck. The male of D. brevicornis is unknown; how- ever, we believe that these two species are members of the same species group, which we recognize here as the Dasyhelea brevi- cornis group (new species group). Among the circum-Caribbean species of Dasyhelea, the only species that might be confused with D. morrisoni is D. pygmaea (Williston) which was described from only three female specimens from St. Vincent (Lesser Antilles). Females of this species from the southeastern Caribbean differ from those of D. morrisoni in having a much nar- rower wing with dense macrotrichia on the wing membrane of all cells (Williston 1896). Another New World species that might be confused with D. morrisoni is the recently described D. huasteca Huerta and Ibanez-Bernal, which is known only from two females from San Luis Potosi, Mexico. Females of this Mexican species differ from those of D. morrisoni in having an antennal flagellum with all flagellomeres longer than broad, a more slender wing with denser ma- crotrichia on the wing membrane of all cells, an abdomen with very long cerci, a small crescent-shaped sclerotization near the caudal margin of sternite 9 and a larger (length 0.04 mm), subspherical spermathe- VOLUME 108, NUMBER 2 ha i Fig. 9. 471 Photograph of small island near Staniel Cay, Exuma Cays, Bahamas, from which a portion of the type series of Dasyhelea morrisoni was collected. Photo by Lloyd Morrison. ca with a very small, rudimentary 2nd sper- matheca (Huerta and Ibanez-Bernal 1999). The very small male genitalia of D. mor- risoni are somewhat reminiscent of those of D. forsteri Grogan and Wirth from the Sol- omon Islands in that the aedeagus of this south Pacific species is also short, crescent- shaped and is covered by an extension of sternite 9. The male of this south Pacific species is only known from the allotype from San Cristobal Island and differs from D. morrisoni in being even smaller (wing length 0.55 mm), having a wing membrane nearly devoid of macrotrichia, exhibiting more massive parameres, and possessing shorter gonocoxites and gonostyli (Grogan and Wirth 1981). Bionomics.—Dasyhelea morrisonit was abundant on very small (usually = 200 m7’) islands (Fig. 9) in the Central Exuma Cays, Bahamas. These small islands are com- posed of marine limestone and were either barren of terrestrial vegetation or supported one or two of three plant species: Sesuvium portulacastrum L., Rhachicallis americana (Jacq.), or Conocarpus erectus L. These are very low-lying islands (< 1.5 m) and, al- though they are emergent at all but possibly the highest tides, they are frequently wet due to salt spray and waves breaking over them. A diversity of arthropod species was found on these small islands, including rep- resentatives of 10 different orders. A total of 658 individuals of D. morrisoni were collected in pan traps and pitfall traps on 10 small islands, and summed over two 24 h sampling intervals, one in December 2002 and one in May 2003. This species was the most abundant arthropod sampled on the small islands, representing >90% of all dipterans and >75% of all insects (Mor- rison 2005). Currently, little is known of the ecology and natural history of D. morrisoni. Larger islands, with higher plant species diversities 472 (Morrison 1997, 2003), are located within several hundred to several thousand meters of the smaller islands, but, these larger is- lands have not yet been sampled for small Diptera. It is unknown whether D. morri- soni completes its life cycle on the small islands or disperses to or from larger is- lands. Because adult females of Dasyhelea are incapable of taking blood meals, it is possible that they can mate and lay a clutch of eggs without even feeding on nectar (Downes 1969). Alternatively, adults of this new species may feed on flowers on nearby, larger islands before engaging in reproduc- tive activities. They may then congregate on the smaller islands, which likely serve as ‘swarming markers’ for males who cop- ulate with females as they enter their all- male swarms. In the case of this new spe- cies, it is unknown whether their eggs are deposited on the smaller or larger islands, but, we suspect that it is more likely the latter case since the larger islands contain more food resources for their developing larvae. ACKNOWLEDGMENTS This article presents results of an under- graduate research project at Salisbury Uni- versity by JAW who prepared all of the il- lustrations, mounted most of the specimens onto slides, and recorded all meristic data. We gratefully acknowledge support of pub- lication costs from the Department of Bio- logical Sciences, Henson School of Sci- ence, Salisbury University. We especially thank Lloyd Morrison for making his midge collections available to us for scientific study, providing the photo of the small is- lands from which they were collected, some of the literature and other information per- taining to the biogeography and ecology of the Exuma Cays, Bahamas. We also thank Peter Cranston, University of California, Davis for his identifications of the Chiron- omidae. Thanks are also extended to Steve Murphree and an anonymous individual for reviewing an earlier draft of the manuscript. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Borkent, A. and W. W. Wirth. 1997. World species of biting midges (Diptera: Ceratopogonidae). Bulle- tin of the American Museum of Natural History, No. 233, 257 pp. Downes, J. A. 1969. The swarming and mating flight of Diptera. Annual Review of Entomology 14: 271-298. Downes, J. A. and W. W. Wirth. 1981. Chapter 28. Ceratopogonidae, pp. 393-421. In McAlpine, J. F, B. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth and D. M. Wood, eds. Manual of Ne- arctic Diptera, Vol. 1. Agriculture Canada Mono- graph, 27, 264 pp. Grogan, W. L., Jr. and W. W. Wirth. 1981. New species of biting midges from the Solomon Islands (Dip- tera: Ceratopogonidae). Pacific Insects 23: 93- 100. Huerta, H. and S. Ibanez-Bernal. 1999. A new species of Dasyhelea Kieffer (Diptera: Ceratopogonidae) and new records of biting midges from the state of San Luis Potosi, Mexico. Proceedings of the Entomological Society of Washington 101: 496— 502. McAlpine, J. EF 1981. Chapter 2. Morphology and ter- minology—adults, pp. 9-63. In McAlpine, J. F, B. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth and D. M. Wood, eds. Manual of Ne- arctic Diptera, Vol. 1. Agriculture Canada Mono- graph, 27, 264 pp. Morrison, L. W. 1997. The insular biogeography of small Bahamian cays. Journal of Ecology 85: 441-454. . 1998a. A review of Bahamian ant (Hymenop- tera: Formicidae) biogeography. Journal of Bio- geography 25: 561-571. . 1998b. The spatiotemporal dynamics of in- sular ant metapopulations. Ecology 79: 1135-— 1146. . 2002a. Island biogeography and metapopula- tion dynamics of Bahamian ants. Journal of Bio- geography 29: 387-394. . 2002b. Determinants of plant species richness on small Bahamian islands. Journal of Biogeog- raphy 29: 931-941. . 2003. Plant species persistence and turnover on small Bahamian islands. Oecologia 136: 51— 62. . 2005. Arthropod diversity and allochthonous- based food webs on tiny oceanic islands. Diversity and Distributions 11: 517-524. Stone, A. and W. W. Wirth. 1947. On the marine midg- es of the genus Clunio Haliday (Diptera, Tendi- pedidae). Proceedings of the Entomological So- ciety of Washington 49: 201-224. Szadziewski, R. 1996. Biting midges from Lower Cre- taceous amber of Lebanon and Upper Cretaceous VOLUME 108, NUMBER 2 Siberian amber of Taimyr (Diptera: Ceratopogon- idae). Studia Dipterologica 3: 23-86. 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. 473 Williston, S. W. 1896. On the Diptera of St. Vincent (West Indies). Transactions of the Entomological Society of London. 1896: 253-446. 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. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 474-475 NOTE Platystoma seminationis (Fabricius) (Diptera: Platystomatidae) Introduced to the United States Platystoma seminationis (Fabricius), na- tive to Europe, has been observed four times in the last three years in northern II- linois by the junior author and is apparently now established in the United States. It was photographed on three occasions (USA: II- linois: Chicago, North Park Village Nature Center, 1S Sun 2003; KR. (Curtis, “slide #1112407; Chicago, Bunker Hill Forest Re- serve, on grapevine, 2004, R. Curtis, #290911, 290919 (Fig. 1); and Glenview, Kennicott Grove, 15 Jul 2005, R. Curtis, #5010887) and one male (deposited in Na- tional Museum of Natural History, Smith- sonian Institution, Washington, DC) was collected at Illinois: Skokie, 8 Jun 2005, R. Curtis. Six subspecies of P. seminationis have been recognized (see Sods 1984). The cap- Beale on right. tured male keys to the typical subspecies in the key of Hennig (1945). Platystoma sem- inationis differs from most Platystomatidae known from the Americas in having mostly dark brown wings with small hyaline spots. It differs from species of Amphicnephes Loew, which have predominantly dark wings, in lacking a complete subapical hy- aline band or a large posteroapical hyaline area and in the having the frons, most of the thorax, and dorsum of the abdomen densely grayish microtrichose, with numer- ous small, shiny black spots or irregular ar- eas (Amphicnephes species are less densely and more evenly microtrichose). According to Sods (1984), P. semina- tionis 1s widely distributed in northern, cen- tral and eastern Europe, occurring in most countries except Norway and Sweden and Platystoma seminationis, Chicago, Bunker Hill Forest Reserve, 2004. Male on left, pair in copula VOLUME 108, NUMBER 2 the Mediterranean region. It seems likely to become widespread in North America. The biology of Platystoma is poorly known, but most species, like the majority of Platysto- matidae, are probably saprophages as lar- vae. Platystoma seminationis was reared from a fungus, P. euphorbiinum Enderlein from rotting roots, and P. lugubre Robi- neau-Desvoidy from humus soil beneath fallen wood and from a World War II cem- etery (Ferrar 1987). LITERATURE CITED Ferrar, P. 1987. A Guide to the Breeding Habits and Immature Stages of Diptera Cyclorrhapha. Ento- monograph 8. E. J. Brill/Scandinavian Science Press, Leiden & Copenhagen, 907 pp. 475 Hennig, W. 1945. 45. Platystomidae. /n Lindner, E., ed. Die Fliegen der palaearktischen Region, 5: 56 PP- Sods, A. 1984. Family Platystomatidae (Platystomi- dae), pp. 38-45. In Sods, A. and L. Papp, eds., Catalogue of Palaearctic Diptera, Vol. 9 Micro- pezidae—Agromyzidae. Akadémiae Kiad6, Bu- dapest, 460 pp. Allen L. Norrbom and Robert Curtis. (ALN) Systematic Entomology Laboratory, PSI, Agriculture Research Service, U.S. De- partment of Agriculture, % National Mu- seum of Natural History, MRC-168, Wash- ington, DC 20013-7012, U.S.A. (e-mail: an- orrbom@ sel.barc.usda. gov); (RC) 2131 W. Sunnyside Ave., Chicago, IL 60625-1609, U.S.A. (e-mail: robirdman@ aol.com) PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 476—477 NOTE Elatophilus oculatus (Drake and Harris) (Hemiptera: Heteroptera: Anthocoridae) Attracted to Pheromone of Matsucoccus Cockerell (Hemiptera: Sternorrhyncha: Matsucoccidae) in Arizona Mendel et al. (2004) published an inter- esting account of Anthocoridae (Hemiptera: Heteroptera) and Hemerobiidae (Neurop- tera) attracted to the sex pheromones of scale insects of the genus Matsucoccus Cockerell (Hemiptera: Sternorrhyncha: Matsucoccidae). Pheromone-baited traps were placed in the Canary Islands, China, Crete, Cyprus, France, ISreal, Portugal, Spain, Turkey, Ukraine, and the United States (Arizona, Massachusetts, and New York). This note deals with the Anthocori- dae attracted to baited traps in the United States. Elotopilus inimicus (Drake and Harris) was described from New York in 1926. Lussier (1965) wrote a thesis on this antho- corid and its relationship with Matsucoccus resinosae Bean and Goodwin (= M. mat- sumurae Kumana) found on Pinus resinosa Ait. Despite this paper, Kelton (1976) felt that P. banksiana Lamb, was its main host. Lattin and Stanton (1993) discussed E. in- imucus as a predator of the red pine scale and reviewed the literature on this relation- ship (Doane 1965, Dross 1985, Mendel et al. 1991). Mendel et al. (2004) reported that E. inimica was attracted to a pheromone of Matsucoccus feytaudi Ducasse and M. mat- sumurae Kuwana in Massachusetts and New York. Mendel et al. (2004) cited a species of Elatophilus as being attracted to the pher- omone of the exotic Matsucoccus feytaudi at Lake Mary, Coconono County, Arizona, in August, 2001, stating that Pinus ponde- rosa Dougl. ex Laws was the host tree in- volved (note: This tree is the subspecies Pi- nus ponderosa scropulorus Engelm. in that area). I have collected nymphs of Elatophi- lus oculatus (Drake and Harris) from Pinus ponderosa scropulorum on May 9, 2001 at Williams, Arizona, approximately 40 miles west northwest of Lake Mary. Williams, Arizona is the type locality of E. oculatus. Only specimens of the aphid genus Cinara sp. were taken with the bugs by beating. The Matsucoccus would be under the bark or in needle sheaths and thus not readily visible. This is the dominant tree through- out much of the southern Rocky Mountains. Further, I have identified specimens of E. oculatus in the Canadian National Collec- tion from Mt. Lemmon, Arizona, on P. ponderosa scropulorum. Many other spec- imens were identified from six localities in Colorado, including my collection of the bug on P. ponderosa scropulorum at Estes Park, Colorado, all in the Rocky Mountain area of the state. A few specimens from this material had been taken from Pinus mon- ticola Dougl. ex D. Don, P. flexilis James, P. contorta latifolia Engelm., and Pseudot- suga taxifolia (Lamb.) Britton. These spec- imens were collected from late July to early September in Arizona and Colorada. No other species of Elatophilus is known from Pinus ponderosa scropulorum in Arizona or Colorado. A related species, Elatophilus pullus Kelton and Anderson, occurs from Oregon north to British Columbia on P. ponderosa Doug]. ex Laws. Thus, it is clear that the species of Elatophilus cited by Mendel et al. (2004) as being attracted to the pheromone of Matsucoccus feytaudi should be regarded as E. oculatus. It seems reasonable that E. oculatus also will feed on a native species of Matsucoccus besides aphids early in the season. My thanks to Z. Mendel and associates, T. J. Henry, an anonymous reviewer, and L. Parks for manuscript preparation. VOLUME 108, NUMBER 2 LITERATURE CITED Doane, C. C. 1965. The red pine scale. Connecticut Agricultural Experimental Station, Circular 207 (revised). 7 pp. Drake, C. J. and H. M. Harris. 1926. Notes on Amer- ican Anthocoridae with descriptions of new forms. Proceedings of the Biological Society of Wash- ington 39: 33—46. Drooz, A. T. 1983. Insects of eastern forests. U.S.D.A., Forest Service, Miscellaneous Publications No. 1426, 608 pp. Kelton, L. A. 1976. The genus Elatophilus Reuter in North America with descriptions of two new spe- cies (Heteroptera; Anthocoride). The Canadian Entomologist 108: 631—634. Lattin, J. D. 2000. Minute pirate bugs (Anthocoridae), pp. 607—636. In Schaefer, C. W. and A. R. Panizzi, eds. Heteroptera of economic importance. CRC Press, Boca Raton, Florida, 828 pp. Lattin, J. D. and N. L. Stanton. 1993. Taxonomic and biological notes on North American species of Elataphilus Reuter (Hemiptera: Heteroptera: An- 477 thocoridae). Journal of the New York Entomolog- ical Society 101: 88—93. Lussier, S. J. 1965. A study of Elatophilus inimucis D. & H. (Hemiptera: Anthocoridae) and its role in the natural control of the red pine scale Matsu- coccrus resinoseae B. and G. (Homoptera: Mar- garodidae). Master’s thesis. University of Massa- chusetts, 42 pp. Mendel, Z., E. Carmi, and H. Podoler. 1991. Relations between the genera Matsucoccus (Hemiptera: Margarodidae) and Elatophilus (Hemiptera: An- thocoridae) and their significance. Annuals of the Entomological Society of America 84: 502— 507. Mendel, Z., A. Fabrene, and E. Dunkelblu. 2004. Kai- romonal attraction of predatory bugs (Heteroptera: Anthocoridae) and brown lacewings (Neuroptera: Hemerobiidae) to sex pheromones of Matsucoccus species (Hemiptera: Matsucoccidea). Biological Control 30: 134-140. John D. Lattin, Department of Botany and Plant Pathology, Oregon State Univer- sity, Corvallis, OR 97331-2902, U.S.A. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 478-479 NOTE Deletion of Paratriphleps pallida (Reuter) from the United States Fauna (Hemiptera: Heteroptera: Anthocoridae) Brachysteles pallidus Reuter (1884) was described from St. Jean (St. John) and St. Thomas islands (now in the Virgin Islands). Uhler (1886) reported this speices from “S.St.”’>. (Southern States) without detail. Uhler (1894a) ceited this species from St. Vincent and from Grenada (1894b). Van Duzee (1917) repeated the citation of Uhler (1886) and reported B. pallidus from the “Southern States.”’ Barber (1939) reported this species (as Paratriphleps pallidus) from Florida, Jamaica, Puerto Rico, and the Virgin Islands. Herring (in Bacheler and Baranowski 1975) stated that the Paratri- phleps from southern Florida was Paratri- phleps laeviusculus Champion rather than P. pallidus, and reported that P. pallidus also occurs in South America, based on specimens from Peru in the U.S. National Museum. It appears that the range of P. pal- lidus extends from Jamaica through Puerto Rico, the Virgin Islands to St. Vincent, Gre- nada, and into South America, at least to Peru. Henry (1988) reported P. pallidus from Florida and southern United States following Van Duzee (1917) and Barber (1939). Henry and Froeschner (1992) al- tered the specific name from P. pallidus to P. pallida, correcting its gender. Lattin (1999) referred to the clarification of the status of P. laeviusculus and P. pallida by Herring in Bacheler and Baranowski (i975): Paratriphleps laeviusculus Champion (1900) was described from Panama, with an excellent illustration of the adult, and has been reported from Honduras (Drake and Harris 1926), Nicaragua (Carpentero et al. 1997), and introducted into Florida (Bach- eler and Baranowski 1975). According to Herring (in Bacheler and Baranowski 1975), the records from Peru by Hambleton (1944) and Wille (1951) should be referred to P pallida (Reuter). Based upon the above evidence, Paratriphleps pallida (Reuter, 1884) is removed from the list of species of Anthocoridae from America north of Mexico. My acknowledgements to D. L. Carpen- tero, T. J. Henry, and the late J. L Herring for their efforts on the anthocorids, and L. Parks for manuscript preparation. LITERATURE CITED Bacheler, J. S. and R. M. Baranowski. 1975. Paratri- phleps laeviusculus, a phytophagous anthocorid new to the United States (Hemiptera: Anthocori- dae). The Florida Entomologist 58: 157—163. Barber, H. G. 1939. Insects of Porto Rico and the Vir- gin Islands. New York Academy of Sciences 14: 263-441. Champion, G. C. 1897-1901. Insecta: Rhynchota (He- miptera—Heteroptera). Vol. II. Jn Godman and Salvin, eds. Biologia Centrail-Americana. London 1900: 305-344. Carpentero, D. L. J. M. Maes, and M. D. Coscar6n. 1997. Catalogo de los Anthocoridae (Heteroptera) de Nicaragua. Revista Nicaraguensis de Entomo- logia. No. 41: 23-28. Drake, C. J. and H. M. Harris. 1926. Notes on Amer- ican Anthocoridae with descriptions of new forms. Proceedings of the Biological Society of Wash- ington 39: 33—46. Hambleton, E. J. 1944. Heliothis virescens as a pest of cotton, with notes on host plants in Peru. Journal of Economic Entomology 37: 660—666. Henry, T. J. 1988. Family Anthocoridae, pp. 12—28. 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, 958 Pp. 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 Unit- ed States.”” Proceedings of the Entomological So- ciety of Washington 94: 263-272. Lattin, J. D. 1999. Bionomics of Anthocoridae. Annual Review of Entomology 44:207—231. Reuter, O. M. 1884. Monographia Anthocoridarum Or- bis Terrrestris. Helsingforsiae, 204 pp. (also Acta VOLUME 108, NUMBER 2 Societatis Scientiarum Fennicae 14(1885): 558-— 578). Uhler, P. R. 1886. Checklist of the Hemiptera: Heter- optera of North: America. Brooklyn Entomolical Society, Brooklyn, New York. 29 pp. + 3 pp. in- dex. . 1894a. A list of the Hemiptera—Heteroptera of the families, Anthocoridae and Ceratocombidae collected by Mr. H. H. Smith on the island of St. Vincent, with descriptions of new genera and spe- cies. Proceeding of the Zoological Society of Lon- don 1893: 156—160. . 1894b. On the Hemiptera—Heteroptera of the island of Grenada, West Indies. Proceedings of the Zoological Society of London 1894: 167-224. 479 Van Duzee, E. P. 1917. Catalogue of the Heniptera of America north of Mexico excepting the Aphidi- dae, Coccidae, Aleurodidae. University of Cali- fornia Publications, Technical Bulletins, Entomol- ogy 2, xvi + 902 pp. Wille, J. E. 1951. Biological control of certain cotton insects and the application of new organic insec- ticides in Peru. Journal of Economic Entomology 44: 12-18. John D. Lattin, Department of Botany and Plant Pathology, Oregon State Univer- sity, Corvallis, OR 97331-2902, U.S.A. PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 480-481 BooK REVIEW Caterpillars of Eastern North America. A Guide to Identification and Natural His- tory, by David L. Wagner. 496 pages, 24 line illustrations, 1,200+ color photos, 5” X 8”; ISBN 0-691-12144-3. $29.95 paperback. Princeton University Press. Publication date: 24 August 2005. As someone who looks at caterpillars on a daily basis as part of work-related chores, I think I have as great an appreciation for this life stage as nearly anyone. However, the larvae I examine usually are washed- out, lifeless creatures from glass vials with alcohol that bear no resemblance to the beauties in Wagner’s new caterpillar book. What an eye-opener it is to get a glimpse of these gorgeous animals in living postures and vivid colors! Because the majority of the lives of many butterflies and moths is spent in the larval stage, it is remarkable that it has taken so long to recognize the value of caterpillar guides such as this book and the previous two USDA-funded publi- cations complied by Wagner et al. (i.e., Cat- erpillars of Eastern Forests and Geome- troid Caterpillars of Northeastern and Ap- palachian Forests). It is inexplicable that although the larval stages of Lepidoptera are responsible for such significant econom- ic damage to forest, ornamental. and agri- cultural plants, eliciting the need for mil- lions of dollars in control measures, many of which result in ecological catastrophes, that this life stage would be known so much less than adults for most species. Go figure. The caterpillar book by Wagner is divid- ed into three parts: Introductory Sections, Family and Species Accounts, and an un- titled third part that includes typical back- material (e.g., glossary, literature, indices). The introductory part provides brief over- views of caterpillar morphology, life cycles, collecting, rearing, natural enemies, photog- raphy, classification, and other general top- ics. The section includes a few half-tones of morphological features in support of the text and a couple photos of collecting tech- niques. The second part is the meat, with over 700 beautiful color images of live lar- vae organized by family in a sequence sim- ilar to that of the Check List of the Lepi- doptera of America North of Mexico (Hodges et al. 1983). Family coverage ranges from Limacodidae to Arctiidae— virtually all of the so-called macrolepidop- tera. Anyone who has ever hunted for cat- erpillars will appreciate the tremendous in- vestment in time it must have taken to dis- cover all the species shown, and those with an interest in natural history photography will appreciate the skill and patience it must have taken to amass this outstanding col- lection of excellent images. The third part of the book is comprised of a glossary, sev- eral pages of bibliographic references, an index of food plants, and an index to the animals and other subjects. In the Family and Species Accounts, most caterpillar images are accompanied by a standard half page (more or less) of text that includes recognition, occurrence, com- mon food plants, and remarks. The text in each of these is rich in pearls ranging from interesting notes on distribution, to varia- tion in color and/or pattern, to current tax- onomic status. For example, I had no idea the walnut sphinx is now Amorpha juglan- dis! [I would have guessed Amorpha jug- landis is a shrub in the Fabaceae with wal- nut-shaped leaves.] In my opinion, one of the most useful features of the species ac- counts is the small insert of an adult moth or butterfly with most of the caterpillars shown. It really drives home what the cat- erpillar will develop into—not just the name of the animal that has to be looked up elsewhere. The only thing initally a little odd about the images of adults is that sphinx moths are the same size as hairstreak butterflies, but I can get used to that. What VOLUME 108, NUMBER 2 a great companion this book will make to Covell’s moth book (1984). Actually, Wag- ner’s book requires the companionship of no other book, field guide, or manual—it is entirely self-contained and field-ready. One thing that is a little unclear to me is the taxonomic coverage of the book. Al- though the scope is primarily macrolepi- doptera, the first three families for which images of larvae are presented (1.e., Lima- codidae, Megalopygidae, and Zygaenidae) are traditionally part of the microlepidop- tera, a category of convenience not of clas- sification. I suspect that these families are included because they are among the larger mircolepidoptera. Also, they may have been included because their caterpillars are likely to be seen by the avid caterpillar-watcher/ hunter. There are, however, many pyraloids and some other microlepidoptera that may be encountered just as commonly—it must be the size thing. Actually, the absence of Pyraloidea and other microlepidoptera can- not help but leave the user wondering if there will be a sequel .... Little Caterpil- lars of Eastern North America? As a micro- lepidopterist, I am incredibly envious of those who study macrolepidoptera because they can now identify most of their moths from larvae observed in the field without having to bring them home to rear. And they have at their fingertips larval features that may help resolve the systematic place- ment of troublesome taxa. 481 In general, the images of caterpillars and adults in this book are superb, the layout is attractive and easy to use, and the small- size format allows it to slip easily into a backpack for use in the field. I strongly rec- ommend this guide to anyone interested in Lepidoptera, but it also should find a place on the bookshelf of anyone interested in natural history, plant-insect interactions, or management of Lepidoptera pests (macros, anyway). It also will be very handy for any- one with inquisitive children (of any age) that pose that frequently asked question— What will it turn into? LITERATURE CITED Covell, C. V. 1984. A Field Guide to the Moths of Eastern North America. Houghton Mifflin Co., Boston, Massachusetts, 496 pp. Hodges, R. W., T. Dominick. D. R. Davis, D. C. Fer- guson, J. G. Franclemont, E. G. Munroe, and J. A. Powell, eds. 1983. Check List of the Lepidop- tera of America North of Mexico (Including Greenland). E. W. Classey, Ltd. and The Wedge Entomological Research Foundation, London, 284 PP- John W. Brown, Systematic Entomology Laboratory, Plant Sciences Institute, Agri- cultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 168, Washington, DC 20013-7012, U.S.A. (e-mail: jbrown@ sel.barc.usda. gov) PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 482-486 MEMBERSHIP LIST OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The previous list was published in January 2003 with 418 members; the present list contains 364 members from every state in the union except Delaware, Idaho, Maine, Massachusetts, Minnesota, Oklahoma, Rhode Island, South Dakota, and Vermont. The largest representation is in Maryland (53), followed by the District of Columbia (39), California (24), Florida (22), and Texas (13). The figures from the Washington, D.C. area are slightly skewed since some members receive their Proceedings at office addresses. Five of the Canadian provinces are represented and 22 other countries, located on six continents. The format used in this list follows essentially that of the 2003 list. Names of Honorary Members (7) are capitalized, those of Emeritus Members (24) are italicized, and Life Members (26) are distinguished by an asterisk following the year they joined the Society. Dates of election to Honorary or Emeritus status are entered in parentheses. In 1988 Ms. Louise M. Russell was elected Honorary Member. In 1998 Ms. Russell was elected Hon- orary President and Dr. Ronald W. Hodges was elected Honorary Member. In 1999 Dr. Donald M. Anderson and Dr. William E. Bickley were elected Honorary Members. In 2002 Dr. Paul J. Spangler and Dr. Manya B. Stoetzel were elected Honorary members. In 2003, after Ms. Russell resigned as Honorary President, Dr. Anderson was elected to the position. In 2005 Dr. Krombein passed away. Later in 2005 Dr. David R. Smith was elected Honorary Member. I thank Dave Smith for his careful reading of the list—and for his corrections—and Mike Pogue for answering questions concerning membership data. Any corrections to the list can be sent to the Membership Secretary by e-mail (williamsh@si.edu) or at the address listed inside the front cover of this issue. Corrections will be read at the next meeting of the Society, and will be published in the Proceedings by the Recording Sec- retary. Holly B. Williams, Membership Secretary Abrahamson, W. G. 1997 Pennsylvania Adams, M. S. 1983 New York Adamski, D. 1984 District of Columbia Adler, P. H. 1986 South Carolina Ahlstrom, K. R. 1992 North Carolina Aitken, T. G. H. 1957 (1984) Connecticut ANDERSON, D. M. 1954 (1999) New York Anderson, L. D. 1944 (1989) California Armitage, B. J. 1983 Ohio Arnaud, P. H., Jr. 1955 California Athanas, M. M. 2001 Maryland Bahr, S. M., II 2004 Texas Baicher, V. V. 2001 Oregon Baixeras, J. 2001 SPAIN Baker, C. A. A. 2005 District of Colum- bia Baker, G. T. 1987 Mississippi Ball, G. E. 1948 Alberta Balogh, G. J. 1994 Michigan Barber, K. N. 1985 Ontario Barnes, J. K. 1978 Arkansas Barrows, E. M. 1976 District of Colum- bia Baumann, R. W. 1972 Utah Baumgardner, D. E. 1992 Texas Beauzay, P. B. 2003 North Dakota Becker, V. O. 1987* BRAZIL Bellinger, R. G. 1972 South Carolina Berry, R. L. 1972 Ohio Bicha, W. 1981 Tennessee BICKLEY, W. E. 1949* (1999) Maryland VOLUME 108, NUMBER 2 Bilby, P. J. 1993 New Jersey Bilyj, B. 1998 Ontario Blank, S. M. 2002 GERMANY Blom, P. E. 1986 Washington Borkent, A. 1988 British Columbia Bowles, D. E. 1993 Texas Brailovsky, H. 1996 MEXICO Brake, I. 2006. District of Columbia Brambila, J. 2004 Florida Branham, M. A. 1998 Ohio Broda, S. 1991 Maryland Brodel, C. EF 1991 Florida Brou, V. A. 1985 Louisiana Brown, B. V. 1993 California Brown, J. W. 1997 District of Columbia Brown, R. L. 1979 Mississippi Bueno-Soria, J. 1977 MEXICO Bunyard, B. A. 2003 Wisconsin Burger, J. FE 1972. New Hampshire Burns, J. M. 1975* District of Columbia Burrows, W. L. 1983 West Virginia Butler, L. 1966 West Virginia Byers, G. W. 1984 Kansas Calabrese, D. 1997 Maryland Calmasur, O. 2004 TURKEY Carlson, R. W. 1970* Maryland Carpenter, T. L. 2005 Maryland Carroll, J. EF 1977 Maryland Cave, R. D. 1977 Florida Cervantes-Peredo, L. 2003 MEXICO Chaboo, C. S. 2001 New York Choi, S.-W. 2002 SOUTH KOREA Choi, W.-Y. 2003 Illinois Clark, W. E. 1975 Alabama Cline, A. R. 2003 Arkansas Codella, S. G. 2001 New Jersey Cohen, E. A., Jr. 1999 Maryland Connor, E. F 1990 California Contreras-Ramos, A. 1986 MEXICO Cook, J. L. 1996 Texas Cooper, K. W. 1955 California Coovert, G. A. 1996 Ohio Coulson, J. R. 1961 Virginia Courtney, G. W. 1985 Iowa Covell, C. V., Jr. 1971 Kentucky Cross, H. E 1954 Georgia Currie, D. C. 1999 Ontario 483 Darling, D. C. 1981 Ontario Darsie, R. E 1949 Florida Davis, D. R. 1961 District of Columbia Davis, L. R., Jr. 1992 Florida Deans, A. R. 2003 Florida Deeming, J. C. 1974* WALES Deitz, L. L. 1982 North Carolina Dennis, S. 1976 Colorado Deyrup, M. A. 1979 Florida Dorchin, N. 2006 Pennsylvania Dozier, H. L. 1952* South Carolina Droege, S. 2004 Maryland Drummond, R. O. 1954 (1987) Texas Duarte, M. 2003 BRAZIL Durkin, P. 1999 District of Columbia Earle, E. I. 2004 Pennsylvania Eckerlin, R. P. 1990 Virginia Epstein, M. E. 1994 California Erwin, T. L. 1972 District of Columbia Espinasa, L. 2004 Virginia Estes, J. 2004 Texas Etnier, D. A. 1999 Tennessee Evans, W. G. 1957 (1994) Alberta Evenhuis, N. L. 1980 Hawaii Favret, C. R. 2003 Illinois Fee, EK D. 1983 Pennsylvania Fisk F. W. 1968 (1988) Florida Flint, O. S., Jr. 1961 Virginia Flowers, R. W. 1994 Florida Flynn, D. 2005 North Carolina Foard, T. 2005 Maryland Foote, B. A. 1958 Ohio Foottit, R. G. 2002 Ontario Foster, G. A. 1999 Maryland Frank, J. H. 1994 Florida Freidberg, A. 1979 ISRAEL Furth, D. G. 1994 District of Columbia Gagné, R. J. 1966* Maryland Gaimari, S. D. 1995 California Gates, M. 2003 District of Columbia Gelhaus, J. K. 1989 Pennsylvania Glaser, J. D. 1988 Maryland Gordon, R. D. 1968 North Dakota Gordon, S. W. 1998 Maryland Gorham, J. R. 1974 (1995) District of Columbia 484 Grissell, E. E. 1979 Arizona Grogan, W. L. 1997 Maryland Guilbert, E. 2000 FRANCE Gullan, P. 2004 California Halbert, S. E. 1989 Florida Hall, J. BR 2002 District of Columbia Hamilton, S. W. 1982 Tennessee Hanson, P. 1985 COSTA RICA Hansson, C. 1985 SWEDEN Harbach, R. E. 1972 UNITED KING- DOM Harlan, H. J. 1988 Maryland Harris, S. C. 1979 Pennsylvania Harrison, B. A. 1976 North Carolina Harrison, T. L. 1993 Illinois Hastriter, M. W. 1998 Utah Heimdal, D. 2003 Iowa Hellenthal, R. A. 2003 Indiana Henry, T. J. 1975 District of Columbia Heppner, J. B. 1974 Florida Heraty, J. M. 1986 California Hespenheide, H. A., III 1981 California Hevel, G. E 1970 District of Columbia Heyn, M. W. 2002 Florida Hight, S. 1990 Florida Hilton, D. FJ, 1990* Quebec Hinojosa-Diaz, I. 2005 Kansas HODGES, R. W. 1960* (1998) Oregon Hoebeke, E. R. 1980 New York Hoffman, K. M. 1986 California Honda, J. 2004 California Howden, H. EF 1948 Ontario Huang, Y.-M. 1968 District of Columbia Husband, R. W. 1972 Michigan Irwin, M. E. 1976 Illinois Ivie, M. A. 1984 Montana Jimenez, H. H. 1994 California Jimenez-Guarda, P. 1999 CHILE Johnson, N. E 1980 Ohio Joseph, S. R. 1957 Maryland Kaster, C. H. 1979 Michigan Kawahara, A. Y. 2004 Maryland Kefter Sy 1993 Wirsinia Kennedy, J. H. 1995 Texas Kimsey, L. S. 1994 California PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Kingsolver, J. 1963 (1992) Florida Kinkorova, J. 2004. CZECH REPUBLIC Kirchner, K. 2004 Maryland Kirchner, R. EF 1981 West Virginia Kittle, PR D. 1975 Alabama Knight, J. B. 2004 Nevada Knutson, L. V. 1963* ITALY Konstantinov, A. 1997 District of Colum- bia Kondratieff, B. C. 1992 Colorado Korch, P. P. 1993 Pennsylvania Kosztarab, M. 1978 (1994) Virginia Kotrba, M. 1997 GERMANY Kruse, J. 2000 Alaska Kula, R. R. 2004 Kansas Labandeira, C. C. 1993 District of Co- lumbia Lago, P. K. 1984* Mississippi Lakin, K. R. 1993 North Carolina LaPolla, J. S. 2005 District of Columbia Landry, B. 2003 SWITZERLAND Lavigne, R. 1999 AUSTRALIA Lee, S. 1999 SOUTH KOREA Lewis, J. A. 1994 District of Columbia. Lewis, P. A. 1974 Ohio Lewis, R. E. 1958 Iowa Li, H. 2003 CHINA Lingafelter, S. W. 1997 District of Co- lumbia Lisowski, E. A. 1988 Washington Loechelt, H. K. 1988 Washington Loeffler, C. C. 1992 Pennsylvania Long, S. 2003 Virginia Maier, C. T. 1976 Connecticut Main, A. J., Jr. 1965 EGYPT Manley, D. G. 1984 South Carolina Manglitz, G. R. 1956 (1989) Nebraska Marsh, P. M. 1960 (1997) Kansas Marshall, S. 1982 Ontario Mason, H. C. 1949 (1973) Maryland Mathis, W. N. 1976* Virginia Mawdsley, J. 2001 District of Columbia MacKay, W. P. 2005 Texas McCreadie, J. W. 2003 Alabama McGovran, E. R. 1937 (1973) Maryland McKamey, S. H. 1989 District of Colum- bia VOLUME 108, NUMBER 2 McPherson, J. E. 1985 Illinois Mead, F. W. 1976 (1995) Florida Menke, A. S. 1969 Arizona Metlevski, J. 2003 Kansas Metzler, E. H. 1998 New Mexico Micheli, C. 2001 Maryland Michener, C. D. 1994 Kansas Miller, D. R. 1972 Maryland Miller, G. L. 1981 Maryland Miller, S. E. 1980* District of Columbia Mitchell, R. T. 1949 (1978) Maryland Moore, T. E. 1950 Michigan Morse, J. C. 1976 South Carolina Moulton, J. K. 1994 Tennessee Mudge, A. D. 2001 Oregon Muegge, M. A. 2003 Texas Munro, J. B. 2000 California Munson, S. C. 1938 (1975) District of Columbia Nakahara, S. 1968 Maryland Navarrete-Heredia, J. L. 2003 MEXICO Neff, S. E. 1969 Pennsylvania Nelson, C. H. 1969 Tennessee Neunzig, H. H. 1956 North Carolina Norrbom, A. L. 1989 District of Colum- bia Novelo-Guitérrez, R. 1999 MEXICO Nuhn, T. P. 1981 Virginia Ochoa, R. 1999 Maryland O’ Neill, K. M. 2000 Montana Opler, P. A. 2001 Colorado Orr, R. L. 1990 Maryland Oswald, J. D. 1987 Texas Ouellette, G. D. 2006 District of Colum- bia Pakaluk, J. 1992* District of Columbia Paras, F 1999 Maryland Parker, C. R. 1977 Tennessee Parrish, D. W. 1963 (1987) Maryland Pennington, W. 2001 Tennessee Perez-Gelabert, D. E. 2000 Maryland Peters, J. G. 1999 Florida Peterson, R. V. 1952 Utah Pinto, J. D. 1982 Oregon Plakidas, P. 2003 Pennsylvania Pogue, M. 1996 District of Columbia 485 Poiner, G. 1999 Oregon Polhemus, D. A. 1993 Hawaii Polhemus, J. T. 1964 Colorado Porter, C. H. 1984 Georgia Price, R. D. 1963 Arkansas Pulawski, W. J. 1975 California Rabaglia, R. J. 2002. Maryland Rainwater, C. F. 1954 (1975) Maryland Rainwater, H. I. 1964 (1983) Maryland Revol, L. M. 1999 FRANCE Richards, A. B. 1996 California Rider, D. A. 2003 North Dakota Richardson, H. H. 1939 (1976) New Jer- sey Robbins, R. G. 1979* Maryland Robbins, R. K. 1986 District of Columbia Robbins, T. O. 1989 Texas Robinson, H. E. 1963 District of Colum- bia Rong, B. 2005 CHINA Root, R. B. 1984 New York Rothschild, M. J. 1989 Maryland Roughley, R. E. 2001 Manitoba Rozen, J. G., Jr. 1956 New York Rueda, P. L. M. 2003 Maryland Ruiter, D. 1976 Colorado RUSSELL, L. M. 1930 (1988) Maryland Russo, R. A. 2004 California Saugstad, E. S. 1979 West Virginia Scarbrough, A. G. 1971 Arizona Schaefer, C. W. 1985 Connecticut Schauff, M. E. 1980 Maryland Schick, K. 1994 California Schiff, N. M. 1991* Mississippi Schmude, K. L. 1990 Wisconsin Schubert, R. D. 2001. Maryland Schultz, T. R. 2000 District of Columbia Schawaroch, V. 2003 New York Scudder, G. G. E. 1984 British Columbia Sedlacek, J. D. 1988 Kentucky Setliff, G. P. 2002 Maryland Seybold, S. J. 2004 California Shaffer, J. C. 1974 Virginia Shands, W. A. 1940 (1991) South Caro- lina Shaw, S. R. 1991 Wyoming Sheng, M.-L. 2005 CHINA 486 Shepard, W. D. 1992 California Shorthouse, J. D. 1986 Ontario Simmons, R. A. 2002 North Dakota Sites, R. W. 1989 Missouri Skelley, P. E. 1992 Florida Sloan, M. J. 1983 (1990) District of Co- lumbia Smiley, R. L. 1964 Maryland SMITH, D. R. 1965* (2005) District of Columbia Snelling, R. R. 1972 California Solis, M. A. 1985* District of Columbia SPANGLER, P. J. 1958* (2002) Missouri Spilman, R. E. 1950 (1977) Maryland Spinelli, G. R. 1983 ARGENTINA Staines, C. L. 1975 Maryland Starr, C. K. 1987 TRINIDAD & TO- BAGO Stebnicka, Z. T. 2004 POLAND Steck, G. J. 1988 Florida Steiner, W. E. 1979 Maryland Steinly, B. A. 1983 Ohio Stewart, R. D. 1985 Maryland Stibick, J. N. L. 1992* Maryland STOETZEL, M. B. 1971 (2002) Mary- land Strazanac, J. S. 2001 West Virginia Strom, D. 2003 Florida Sullivan, J.B. 2003 North Carolina Sutherland, C. M. 1974 New Mexico Sutherland, D. W. S. 1973* Maryland Tauber, C. A. 2003 California Taylor, M. C. 2004 Maryland Tennessen, K. J. 1982 Wisconsin Thompson, F C. 1968* District of Co- lumbia Thompson, J. V. 1953 (1985) New Jersey Togashi, I. 1983* JAPAN Torres-Miller, L. R. 1984 West Virginia Triplehorn, C. A. 1972 Ohio PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ulrich, H. 1978* GERMANY Valley, K. 1976 Pennsylvania Vandenberg, N. 1993 District of Colum- bia Vazquez, A. W. 1957 Virginia Voegtlin, D. 1981 Illinois Wahl, D. B. 1987 Florida Wallace, M. S. 2002 North Carolina Wallenmaier, T. E. 1979 Michigan Wan, X. 2003 CHINA Warren, A. D. 1999 Oregon Webb, D. W. 1981 Illinois Weber, N. A. 1941 (1981) Florida Wenzel, R. L. 1984 Illinois Wharton, R. A. 1981 Texas Wheeler, A. G., Jr. 1974 South Carolina Wheeler, T. A. 2001 Quebec White, G. B. 1977* Maryland Whitworth, T. 1999 Washington Wiker, J. R. 2003 Illinois Williams, G. L. 1984 Maryland Williams, H. B. 1977* District of Colum- bia Wilson, D. 1999 Mississippi Wilson, N. 1957 Iowa Wojtowicz, J. A. 2005 Tennessee Wood, D. M. 1987 Ontario Woodley, N. E. 1984* District of Colum- bia Woolley, J. B. 1986 Texas Wright, D. 2005 Ohio Yasunaga, T. 1998 JAPAN Young, D. K. 1981 Wisconsin Zack, R. 1982 Washington Zhang, L. 2004 CHINA Zhang, Y. 2004 CHINA Zuparko, R. L. 1993 California Zyla, J. D. 2000 Maryland PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 487—488 REPORTS OF OFFICERS EDITOR Volume 107 of the Proceedings included four issues with a total of 996 pages. Nine- ty-eight regular papers, 18 notes, 6 book re- views, minutes of Society meetings, reports of officers, instructions for authors, and the table of contents for volume 107 were pub- lished. About 122 manuscripts (regular pa- pers and notes) were submitted for consid- eration for publication from November 1, 2004 through October 31, 2005. Miscellaneous Publication “‘A phyloge- netic revision of the Napaeina (Lepidoptera: Riodinidae: Mesosemiini)”” by Jason P. W. Hall is in its final stages and should be pub- lished by the end of 2005. I extend thanks to members of the Pub- lications Committee—Ray Gagné, Tom Henry, and Wayne Mathis—for their ad- vice, encouragement, and support. Ray Gagné, as Book Review Editor, has ob- tained some excellent book reviews. I am especially grateful to the many reviewers for their time-consuming efforts and con- structive reviews of manuscripts. Their con- tributions are essential to help increase the quality of papers published in the Proceed- ings. Respectfully submitted, David R. Smith, Editor TREASURER SUMMARY FINANCIAL STATEMENT FOR 2005 Special General Fund Publication Fund Total Assets Assets: November 1, 2004 $45,804.81 $120,246.00 $190,315.75 Total Receipts for 2005 S75 17 S10 $ 11,614.74 $ 86,792.84 Total Disbursements for 2005 $76,324.34 $ 2,000.00 $)7/8.524:34 Assets: October 31, 2005 $44,658.57 $129,860.74 $174551931 Net Changes in Funds $ (1,146.24) $ 9,614.74 $ 8,468.50 Audited by the Auditing Committee, November 28, 2005, consisting of Norman E. Wood- ley, Chairman, John W. Brown, and Alex S. Konstantinov. Presented to the membership at the meeting of December 1, 2005. Respectfully submitted, Michael G. Pogue, Treasurer 488 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MEMBERSHIP SECRETARY In 2005, the Society received applications for membership from 12 people: Caroline A. A. Baker Xavier Preud’ homme Terry L. Carpenter Bao Rong Duane Flynn Mao-Ling Sheng Timothy Foard Catherine A. Tauber Ismael A. Hinojosa-Diaz John Anthony Wojtowicz John S. LaPolla Quig-He Zhang Each applicant was sent a letter of welcome to the Society and his/her name was read at a regular monthly meeting. The number of applications decreased 8% from 2004. Other letters from the Membership Secretary included six letters to guest speakers, four letters to special fund contributors, and four replies to requests for information. Respectfully submitted, Holly B. Williams, Membership Secretary PROC. ENTOMOL. SOC. WASH. 108(2), 2006, pp. 489-491 SOCIETY MEETINGS 1,093rd Meeting—June 2, 2005 The 1,093rd meeting of the Entomologi- cal Society of Washington consisted of the Annual Banquet, held at the National Mu- seum of Natural History, Smithsonian Insti- tution. An open bar starting at 6:00 p.m. was followed by the formal banquet at 7:00 p.m. Approximately 85 members and guests were in attendance. At 8:00 p.m., the herd was prodded and corralled into the Baird Audi- torium for a lively presentation by Dr. Mary Liz Jameson, of the University of Nebraska. She discussed some activities of ‘Team Scarab” and went on to describe the well- known and some lesser-known habits of the hyperdiverse Scarabaeidae, with plenty of anecdotes to please the crowd. Toward the end, she passed around an amusingly labeled box of coco-puff cereal to sample—at least, we hoped it was cereal. The meeting ad- journed at about 9:30 p.m. Respectfully submitted, Stuart McKamey, Recording secretary 1,094th Regular Meeting—October 6, 2005 The 1094th regular meeting of the En- tomological Society of Washington (ESW) was called to order in the Cathy Kerby room of the National Museum of Natural History, Washington, D.C., by President Ja- son Hall at 7:10 p.m. The meeting was at- tended by 22 members and 5 guests. The minutes were read and approved with mod- ification. There were 5 new applicants for mem- bership: Duane Flynn, John Wojtowicz, Is- mael Hinojosa-Diaz, Quig-He Zhang, and Mao-Ling Sheng. No new members were present; one visitor was introduced. In miscellaneous business, Dave Smith, a long-time and active member of ESW, was elected to an Honorary Membership, which became available with the passing of Karl Krombein. For exhibits, Warren Steiner brought a poster about the new beetle family, Merui- dae, that he just described with Dr. Span- gler. Chris Thompson had on hand the Spe- cies2000 Annual Checklist CD for 2005, and noted that the USDA Systematic En- tomology Laboratory was the largest con- tributor, providing more than 150,000 rec- ords. Dave Furth brought two new books: “Photographic Atlas of Entomology and Guide to Insect Identification,” by J. L. Castner, and “Scorpions of Southern Afri- ca” by J. Leeming. He also had BugFest T- shirts for sale at $10, and childrens size shirts with beetle illustrations. Dave Furth introduced the evening’s speaker, Dr. R. Toby Schuh of the American Museum of Natural History in New York. His presentation was entitled “Exploring the Southern Hemisphere: Plant Bug Plan- etary Biodiversity Inventory Field Work in Australia and South Africa.”’ Together with collaborators, Dr. Schuh won one of four Planetary Biodiversity Inventory (PBI) grants, the others going to projects on slime molds (1,000 species), the plant genus So- lanum (1,500 species) and catfish (2,000 species). By comparison, species richness of the mirid subfamilies Orthotylinae and Phylinae, on which Dr. Schuh’s project fo- cuses, 1s much greater than the others com- bined. These subfamilies are relatively spe- cies-poor in the Neotropics. The project fo- cused on other, likely undersampled re- gions: the Western Cape region of South Africa and eastern, central, and especially southwestern Australia—both foci have high floral species richness and endemism. The team collected by sweeping and beat- ing, with much effort to collect and identify hosts. The life history of Miridae facilitates host association because the adults are usu- ally monophagous and occur when the host is in flower. Their PBI collection increased available specimens by 20% (over 490 100,000), species richness by 20% (over 400), 100% host records, and 1000% host vouchers for Miridae. Dr. Schuh’s favorite spot was the plant-covered dunes of Kal- barri Park in Western Australia, which he described as the “‘Garden of Eden.”’ As their productivity attests, their field work was intelligently designed. The meeting was adjourned at 8:30 p.m. Refreshments were provided by the Society. Respectfully submitted, Stuart McKamey, Recording Secretary 1,095th Regular Meeting—November 3, 2005 The 1,095th regular meeting of the En- tomological Society of Washington (ESW) was called to order in the Cathy Kerby room of the National Museum of Natural History, Washington, D.C., by President Ja- son Hall at 7:04 p.m. The meeting was at- tended by 22 members and 21 guests. The minutes of the 1,094th meeting were ap- proved as read. There were no new applicants for mem- bership. Eight visitors were introduced. In miscellaneous business, Dave Furth and John Brown announced that all of this year’s officers, except president-elect and president, have accepted nomination to con- tinue next year. Voting will take place at an upcoming meeting. For exhibits, Edd Barrows described the journey of a mystery package and the rem- nants of its arachnoid contents after it was squished when finally opened. Jon Mawds- ley had the new book, “A Monograph of the Genus Monticora,”’ a genus of carabids, by J. Maes. Virginia Kirby displayed mis- cellaneous literature about insects including the useful “‘National Pest Control Associ- ation Field Guide to Structural Pests’ by FE. H. Smith & R. C. Whitman. Art Evans had the “‘Guide to the Grasshoppers of Wis- consin” by K. Kirk & C. R. Bomar, men- tioned that the new Virginia Society of Nat- uralists had its first meeting last month, and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON announced an upcoming BioBlitz of Poto- mac Gorge. Dolores Savingnano announced an upcoming biological survey of Arlington starting next year. Dave Furth brought two books: “A Dazzle of Dragonflies” by F L. Mitchell and J. L. Lasswell, “Evolution of Insects,” which includes a thorough pale- ontological perspective by D. Grimaldi and M. S. Engel. Art Evans showed summer slides from his upcoming book on insects of Virginia and the Carolinas. Lastly, War- ren Steiner had a slide show and book, “*Te- nebrionids: A Systematic Revision of the Genus Eucrytus and its Complex,” by K. Ando. Dave Furth introduced the evening’s speaker, Gary Hevel of the Entomology De- partment, Smithsonian Institution, who pro- vided a humble short title: “‘Backyard En- tomology.”’ For the past three years, Gary has been surveying his own suburban back- yard to put some teeth into the common claims by entomologists about how many species occur in a small area. The lot is only two acres, located in Silver Spring, Mary- land, yet he has collected more than 4,000 species. It’s noteworthy that his survey is of species, determined by specialists, rather than morphospecies. He uses a wide variety of collection methods and has netted some important finds. Among them, there are the first U.S. records for three species (a bu- prestid, a chalcidoid, and a moth). His sur- vey has been the subject of several news articles and the subject of a full-length Jap- anese documentary. The meeting was adjourned at 8:55 p.m. Refreshments were provided by the Society. Respectfully submitted, Stuart McKamey, Recording Secretary 1,096th Regular Meeting—December 1, 2005 The 1,096th meeting meeting of the En- tomological Society of Washington (ESW) was called to order in the Cathy Kerby room of the National Museum of Natural VOLUME 108, NUMBER 2 History, Washington, D.C., by President Ja- son Hall at 7:11 p.m. The meeting was at- tended by 17 members and 7 guests. The minutes of the 1,095th meeting were ap- proved as read. In miscellaneous business, President Hall read ESW Proceeding’s editor Dave Smith’s Annual Report. John Brown summarized Treasurer Mike Pogue’s Annual Report. Membership Secretary Holly Williams gave her Annual Report. Dave Furth presented the slate of officers for 2006, with Michael Gates as President-Elect. There were no new nominations from the floor and the slate was approved unanimously. There were no new applicants for mem- bership. Two visitors were introduced. For exhibits, Dave Furth announced that the Smithsonian Entomology library now has a nearly complete set of the journal “Mitteilungen des Entomologischen Ver- eins Stuttgart,’ published by the Staatlich- en Museum fiir Naturkunde—apparently the only copies of the journal in the Western Hemisphere. He had on hand the recent vol- ume 39, “‘Jilustrated Catalogue and Bibli- ography of World Anthribids” by J. Rhein- heimes. Dave Furth introduced the evening’s speaker, Dr. Desmond Foley, National Re- source Council Research Associate and sci- entist at the Smithsonian’s Walter Reed Biosystematics Unit. His presentation was entitled “‘The Curious Case of the Ant & the Mosquito and Other Tales of Culicid 49] Complexity.”” There are about 3,200 spe- cies of mosquitos, about 80 of which are major disease vectors but only a few of which bite humans. Through two examples, Dr. Foley demonstrated the biological di- versity of mosquitos. The first was the New Guinea genus Malaya with a highly modi- fied proboscis that is useless for blood feed- ing. Instead, the mosquitos hover over ant trails and pounce on unlucky individuals, which regurgitate their food onto the mos- quito’s proboscis. He even provided a vid- eo, donated by a Japanese colleague, of the stranger-than-fiction behavior. The second example was his study of the Australian Anopheles annulipes complex, which are Malaria vectors. Anomalous patterns of rabbit infection suggested the presence of more than one species, with differing biol- ogies. Genetic analysis by allozymes and cross-infertility studies suggest that the complex comprises 18 to 25 cryptic spe- cies. Molecular sequence and genetic dis- tance studies are congruent. The complexity of the Anopheles annulipes complex ex- emplifies the need for greater taxonomic resolution to medically important mosqui- tos. The meeting was adjourned at 8:15 p.m. Refreshments were provided by the Society and Ralph Eckerlin, who provided his tra- ditional December-meeting eggnog. Respectfully submitted, Stuart McKamey, Recording Secretary NEW PUBLICATION A Phylogenetic Revision of the Napaeina (Lepidoptera: Riodinidae: Mesosemiin1) by Jason P. W. Hall Published by the Entomological Society of Washington, December 2005. 235 pp., 162 figures including 8 color plates. Hardback. ISBN 1-891276-52-2. Butterflies of the subtribe Napaeima are revised; 6 genera and 44 species are treated. Chapters include phylogenetic analysis, key to the genera, habitats and ecology of adults, food plants and immature stages, biogeography and conservation, and generic and species accounts. See inside back cover for ordering. ENTOMOLOGICAL SOCIETY OF WASHINGTON http://entomology.si.edu/ES W/ES W Menus. lasso Information and Officers Bylaws History First 100 Years Past Presidents History of the ESW Seal Information for Authors Available Publications Meetings Members Database Membership Application Subscription 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 [Bie Srelngehed hi re Skye) olypd OO Meta SY AE De Tl NE CAND eR NT. A OME See Cee ne veal DT neAe Siy RTL A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. BockGrisseliiand wiGhaeliE? Schautiisimpp L997 see sano) ne hha hy Le eos Sa ee Revision of the Oriental Species of Aphthona Chevrolat (Coleoptera: Chrysomelidae), by Alexander S. Ronstantinayand Steyenww, eingatelter, S49pps 2002, ae A ee Np SLA oath e kh Revision of the Genus Anoplophora (Coleoptera: Cerambycidae), by Steven W. Lingafelter and E. | ules tne li Ko felorel ee 502221 54 0) oF 25 0) 0 PAR WENN ahaa cee ak POD iL aI Bf ie we ev ea bia te fa a A Phylogenetic Revision of the Napaeina (Lepidoptera: Riodinidae: Mesosemiini), by Jason P. W. Hall. DESIG) DMA UTS) As Ghee Bel ALE GAM Up RU SERRA A Ee a Raed MN, Jie 6 Fah te ae NL eer gee ear. US OO MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. No.1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 ___________ No.4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952... No.5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 No.6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi Sa ENS A 0 oat be 62 a Me Re BRO RAISE es Colpurint )droim SUmMAtra 08 Ny YNE © Seed) UR SCAT ee acs a Se 646 CARRILLO-RUIZ, HORTENSIA and MIGUEL ANGEL MORON—Study on the phylogenetic relationships of the hopliids (Coleoptera: Scarabaeoidea)...............0.-002 ee ee eee 619 CURLER, GREGORY R., J. PHASUK, J. CHANPAISAENG, and G. W. COURTNEY—A new species of Horaiella Tonnoir (Diptera: Psychodidae) from Thailand ................... 519 ESPINASA, LUIS and ADRIANA FISHER—A cavernicolous species of the genus Anelpistina (Zygentoma: Nicoletiidae) from San Sebastian Cave, Oaxaca, Mexico ................. 655 ETNIER, DAVID A.—A new Neophylax (Trichoptera: Uenoidae) from middle Tennessee ..... 684 FOOTTIT, ROBERT G., SUSAN E. HALBERT, GARY L. MILLER, ERIC MAW, and LOUISE M. RUSSELL—Adventive aphids (Hemiptera: Aphididae) of America north of Mexico... 583 GOK, ALI and EBRU GUL ASLAN—Species composition and abundance of fiea beetles (Coleoptera: Chrysomelidae) associated with moist habitats in Isparta and Burdur provinces, CRUE ECS (0 a papa tee Se aA Mae en a es ONS Sc eis tar tes eal hn eRe CREP NCP aA ee 543 LAGO, PAUL K. and STEVEN C. HARRIS—A new species of Hydropsyche (Trichoptera: Hydropsychidae) from Alabama, with notes on H. frisoni Ross and an unusual Hydropsyche Ciirehealed E83) airs be vad pin tt aA ARY DM Meal SR A Sag Te Sa it a Mi ER i naa SO CC 559 LEE, SEUNGHWAN and HYOJOONG KIM—A fern aphid, Neomacromyzus cyrtomicola Lee, new genus and new species (Hemiptera: Aphididae) on Cyrtomium falcatum (Dryopteridaceae) PAs ASALTOCK GAVESH ON Suit apste sere Vives one ay UDMeuet ADVE IMEI Ma ESS ee Ui ER ef kas fa 493 McKAMEY, STUART H.—Further new genus-group names in the Cicadellidae (Hemiptera)... 502 McKAMEY, STUART H.—Two new species of the Neotropical leafhopper genus Caldwelliola Youns (Hemiptera: Cicadellidae) with'a key toimales: 2). 20s oe ins a ee 534 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2006 STEVEN W. LINGAFELTER, President Jon A. Lewis, Custodian MICHAEL W. GATES, President-Elect MICHAEL G. POGUE, Treasurer STUART H. McKaAmMey, Recording Secretary JOHN W. BROWN AND DAvID G. FurTH, Program Chairs Ho..is B. WILLIAMS, Membership Secretary JASON P. W. HALL, Past President Davip R. Situ, Editor Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY WAYNE N. MATHIS Honorary Members WILLIAM E. BICKLEY Davipb R. SMITH RONALD W. HODGES PAUL J. SPANGLER MANYA B. STOETZEL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168. MEETINGS.— Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu- tion, on the first Thursday of each month from October to June, inclusive, at 7:00 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. An- nual 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 ad- dress changes to the Entomological Society of Washington, % Department of Entomology, Smithsonian Institu- tion, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomological 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. 747-748 OF THIS ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Wash- ington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 30 June 2006 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 493-501 A FERN APHID, NEOMACROMYZUS CYRTOMICOLA LEE, NEW GENUS AND NEW SPECIES (HEMIPTERA: APHIDIDAE) ON CYRTOMIUM FALCATUM (DRYOPTERIDACEAE) IN BASALT ROCK CAVES SEUNGHWAN LEE AND HYOJOONG KIM Laboratory of Insect Biosystematics, College of Agriculture and Life Sciences, Seoul National University, San 56-1, Sillim-dong, Gwanak-gu, Seoul, 151-742, Korea (South) (e-mail: seung@snu.ac.kr) Abstract.—A new cave-living fern aphid, Neomacromyzus cyrtomicola Lee, new genus and new species, is recognized on Cyrtomium falcatum (L.) Presl. (Dryopter- idaceae) in basalt rock caves located on Jeju Island, a volcanic island in the southernmost region of Korea. The new genus, Neomacromyzus, is established by the following characteristics: small body size (1.0—-1.5 mm) with sooty black coloring and pale legs; short dorsal and marginal hairs, which are strongly capitate or flabellate: thorax and abdomen strongly wrinkled dorsally; and post-siphuncular sclerites strongly spinulate. A key to the genera of fern-infesting aphids in the eastern Palaearctic and the Indian subregions is presented. Key Words: Asia Jeju Island is located in the southern- most region of the Korean Peninsula, its north latitude ranging from 33°19'10” to 33°25'35”", and its east longitude ranging momumpleGe27) 501. .t0 bal26°37155%19 he island originated from four main volca- nic eruptions in the Pleistocene period (1,200,000~30,000 years ago). There are numerous sizes of lava caves along with many basalt rock cliffs and volcanic cones emepresemtcdss by)! Mit.) alla (1,950 m), which is located at the center of the island. The climate of Jeju Island is subtropical in the seaside lower eleva- tions, typically temperate at higher ele- vations, and subarctic at the top of Mt. Halla. Therefore, the flora and fauna of Jeju Island is very rich, having more than 1,800 species of vascular plants and 79 known aphid species in the subfamily Aphidinae (Lee et al. 2002). fern aphids, new genus, Neomacromyzus, key to genera, Far East Cyrtomium falcatum (L.) Presi. (Dryopteridaceae) is a subtropical fern which typically grows in wooded, shady hollows or on shady basalt rock cliffs near the sea. With adequate sunlight, leaves of the Cyrtomium falcatum crown can grow to 40-80 cm long with up to 11 leaves. It is able to live in the humid, dark areas of lava caves that receive indirect sunlight, but, the plants remain small in caves, with few leaves larger than 10 cm in length. There are seven aphid genera (Macro- myzus Takahashi 1960; Macromyzella Ghosh et al. 1977; Shinjia Takahashi 1938; Micromyzodium David 1958; Mi- cromyzus van der Goot 1917; Taiwano- myzus Tao 1963; Amphorophora Buckton 1876) that are known to live on various ferns in the eastern Palaearctic and the Indian subregions (Ghosh 1974; Ghosh 494 et al. 1977; Miyazaki 1968, 1971; Mori- tsu) 1952. Takahashi 1921) 1938-5 1960: David and Narayana 1968; Tao 1963). Among them, Macromyzus woodwardiae (Takahashi 1921) is the only aphid recorded on Cyrtomium falcatum, a fern common on Jeju Island. Macromyzus woodwardiae often can be found in large colonies on the underside of the young leaves, developing before the fruiting of spores, wherever the host plants occur (Zee 2002): The tiny black aphid described below is found on the young plants of Cyrto- mium falcatum which grow on rock walls in small, dark lava caves. The new aphid is not congeneric with any previously known aphid. In this context, Neoma- cromyzus Lee, n. gen., is established for the species Neomacromyzus cyrtomicola Kees in. Sp: MATERIALS AND METHODS Samples of aphid colonies were pre- served in 80% ethyl alcohol before slide preparations. Three to five adult speci- mens were placed in a plastic centrifuge tube of 95% alcohol and boiled on a heat block for 2-3 minutes (not necessary after being preserved in ethyl alcohol for more than | month). Thereafter, the ventral side of the abdomen of each specimen was punctured by a minute insect pin and macerated in 10% potas- sium hydroxide (KOH) for about 5 min- utes. The macerated specimens were washed 5-6 times in distilled water for at least 5 minutes each time and dehy- drated in glacial acetic acid. After being totally dehydrated, each specimen was cleared in clove oil for 10—20 minutes and, finally, mounted in a drop of Canada balsam on a slide (see Blackman and Eastop 2000 for details). Illustrations for macerated specimens were taken by a digital camera (Baumer Optronic ARC HR33C, Germany), at- tached on an upright microscope (Leica DM 400B) at a resolution of 600 dpi. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Measurements for each specimens also were taken with the digital images by software (Image Lab version 2.2.4.0, MCM Design (Ltd.)). Abbreviations used for keys and de- scriptions are as follows: Ant.I, HI, VIb— antennal segment I, III, base of VI, respectively; apt.—apterous viviparous female, aptera; Ant.II]BD—basal diame- ter of antennal segment III; BL—length of body; A—length of entire antennae; 2HT— second segment of hind tarsus; PT— processus terminalis; SIPH—siphunculus; URS-—ultimate rostral segment. SYSTEMATICS Neomacromyzus Lee, new genus Apterous viviparous female.—Body oval and small, 1.01—1.51 mm in length; entirely sooty black, except shiny biack on abdominal terga 1—6 (Fig. 13). Ap- pendages pale including antennal flagel- lum and siphunculus (Fig. 13). Head strongly spinulate both dorsally and ventrally, with 4 pairs of dorsal hairs (Fig. 9). Dorsal hairs on head capitate, 1.52.0 times as long as basal width of antennal segment III. Antennal tubercles well-developed, subparallel and strongly granulated, with single capitate hair on each side. Median tubercle on frons weakly developed with two pairs of capitate hairs. Antenna pale smooth or weakly imbricated with short blunt hairs and 1-2 secondary rhinaria on Ant.III (Fig. 7-8); primary rhinaria on Ant.V ciliate. Rostrum attaining hind coxae; mandibular laminae with 3 long acumi- nate hairs on each side; ultimate rostral segment |.50—2.38 times as long as 2HT, 1.00—1.45 times as long as Ant.Vib, with two pairs of accessory hairs (Fig. 11). Thorax strongly wrinkled dorsally and reticulated marginally, with one pair of spinal hairs and 2 hairs on each marginal sclerite. Dorsal and marginal hairs on thorax strongly capitate or flabellate. Legs pale; first tarsal chaetotaxy 3:3:3. VOLUME 108, NUMBER 3 Abdomen strongly wrinkled dorsally and spinulated ventrally. Postsiphucular sclerites strongly spinulated as in Tox- optera Koch 1856. Dorsal and marginal hairs on abdomen flabellate or strongly capitate (Figs. 3-4); abdominal tergum III with 3 pairs of spinal hairs and 2-3 hairs on marginal sclerites; abdominal terga VI and VII each with 1 pair of spinal hairs. Siphunculus cylindrical, well-fringed, and weakly tapering distal- ly; darkly pigmented basally and pale distally (Fig. 5). Cauda short triangular, constricted in middle, with 4-5 long acuminate hairs (Fig. 12). Remarks.—This genus can be distin- guished from other aphid genera by the following characteristics: body small (1.0-1.5 mm), sooty black with pale appendages including legs, antennae, and siphunculus (Fig. 13); dorsal and marginal hairs short and strongly capi- tate or flabellate (Figs. 4, 13); thorax and abdomen strongly wrinkled dorsally (Figs. 1-2); and post-siphuncular scler- ites strongly spinulate. Neomacromyzus appears to be more closely related to the genus Macromyzus than the other fern infesting aphids, by the reticulated dorsum of abdomen, well- developed antennal tubercles, and in- festation on the same host plant, Cyrto- mium falcatum. The two may be distin- guished by the following combination of characters (those of Macromyzus in parenthesis): siphunculus pale, cylindri- cal, and not reticulated, at most less than one row of reticulation at distal apex (entirely dark, constricted and strongly reticulated at distal 1/6); dorsal setae on body short and strongly capitate or flabellate (relatively long, blunt or weak- ly capitate at apex), ultimate rostral segments with 4-6 hairs (with 10-13 hairs); antennal segment III with 1-3 secondary rhinaria on basal half (with- out secondary rhinarium). The new genus also can be distinguished from the other aphid genera of the tribe 495 Macrosiphini with well-developed fron- tal tubercles and capitate hairs (Capito- phorus, Pleotrichophorus, Chaetosiphon) by the strongly elevated scleroites at the base of the dorsal setae on the thorax and abdomen, body small and sooty black, and the host limitation on ferns. Etymology.—The name Neomacromy- zus, 1s masculine and derived from Latin prefix Neo- (new) and the other sympat- ric genus name, Macromyzus living on the same host plant. Type species.—Neomacromyzus cyr- tomicola Lee, n. sp. KEY TO THE GENERA OF FERN-INFESTING APHIDS IN THE EASTERN PALEARCTIC AND INDIAN SUBREGIONS 1. Siphunculus with polygonal reticulations on distal apex on both apterae and alate — Siphunculus not reticulated, at most one or two lines of wrinkles on distal apex (Fig. 5) 3 2. First tarsal chaetotaxy 4:4:4 or 4:4:3. Abdominal dorsum with darkly pig- mented, tuberclelike, elevated scleroites at base of dorsal hairs. On various ferns: Dryopteris, Rumobra, Woodwardia, Poly- stichum, Asplenium, Cyrtomium falcatum. In far eastern and southern Asia (India, MN Korea, Japan, China, Taiwan) (Type species: Myzus woodwardiae Takahashi PAD oe ono eebee Macromyzus Takahashi 1960 — First tarsal chaetotaxy 3:3:3. Abdominal dorsum evenly pale without elevated scler- oites at base of dorsal hairs. Found on Dryopteris arida (Don) Kuntze, Polysti- chum, Pteridium (Pteridaceae). In Korea (Jeju-do), Japan, China (Sichuan), Taiwan, Indonesia (Sumatra), India (Type species: Myzus polypodicola’ Takahashi OBA cae wer eran ore Macromyzella et al. 1977 3. All legs with extremely reduced tarsi without claws. First tarsal chaetotaxy 1:0:0. Found on Pteridium aquilinum var. latiusculum Underw. ex A.Heller (Hypole- pidaceae). In Korea, Japan, China (Zhejing, Sichuan), Nepal, India, Russia, Australia (Type species: Microtarsus pterydifoliae Shinji 1929 = Atarsos orientalis Mordvilko 1929) Shinjia Takahashi 1938 — Tarsi not reduced, with claws. First tarsal chaetotaxy 3:3:3, 4:4:4, or 5:5:5 4. Dorsal setae strongly capitate or fla- bellate (Figs. 3-4). Thorax and abdomi- nal dorsum with well-pigmented, elevated 496 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ae « WA). Figs. 1-12. Apterous viviparous female of Neomacromyzus cyrtomicola. 1, Entire body. 2, Dorsal surface of anterior dorsum of abdomen. 3, Marginal setae of thorax. 4, Dorsal and marginal setae of abdominal tergite 5—6. 5, Siphunculus. 6, Hind tibia and tarsus. 7, Antennal segment III-IV. 8, Antennal segment V-VI. 9, Head and antennal segment I. 10, Hind tarsus. 11, Ultimate rostral segment. 12, Cauda. VOLUME 108, NUMBER 3 Fig. 13. Apterous viviparous female of Neomacromyzus cyrtomicola on a young leaf of Cyrtomium falcatum. scleroites at base of dorsal setae (Figs. 2, 4). Tiny black aphid (Fig. 13). On Cyrtomium falcatum in Korea (Jeju-do) (Type species: Neomacromyzus cyrtomicola, TESST )iee 4S Ape gee Neomacromyzus, n. gen. Dorsal hairs acuminate or blunt, not capitate or flabellate. Base of dorsal setae on thorax and abdomen low, not elevated 5 Apterae without secondary rhinaria on antenna. Head spinulous, both dorsally and ventrally. Antennal tubercles diver- gent. On various ferns: Adiantum, Las- trea, Nephrolepides, Pitygramma, Polypo- dium, Pteris, Streptocarpus, Asplenium, Digitaria. In India, Japan (Type spe- cies: Micromyzodium filicium David L958) Pie 2 Meee Micromyzodium David 1958 Apterae with secondary rhinaria on anten- na. Head smooth or scabrous on dorsum, if scabrous, antennal tubercles strongly con- WGI EATIDS, shies ee ee Seat ah oleae RU utaaeS ie oA ie oot 6 Siphunculus cylindrical. Antenna and tibi- ae entirely black. On Osmunda, Athyrium. In Korea, Japan, China, Taiwan, India (Type species: M. niger van der Goot OM ies es Micromyzus van der Goot 1917 Siphunculus swollen. Antenna and tibiae paler at base thanjother pantssys eer 7 Body small, less than 2 mm in length. Head scabrous dorsally with short blunt hairs. Antennal tubercles strongly convergent. Abdomen with well-pigmented sclerites or scleroites on spinal or marginal region. Cauda normally with 4 hairs. On Athyrium, Dryopteris, Osmunda, Astilbe, Chrysosple- nium. In Japan, India, Taiwan (Type species: Myzus montanus Takahashi 1925) ieee Notes, acti Taiwanomyzus Tao 1963 Body usually larger than 3 mm in length. Head smooth with long acuminate hairs. Antennal tubercles divergent or parallel. Abdomen membranous. Cauda with more than 10 hairs. On various ferns (Athyrium, Lastrea, Pteris, Asplenium), Rubus, Geranium, Filipendula, Cryptotae- nia, Anemone. Widely distributed in Pa- 498 laearctic and Nearctic Regions (Type spe- cies: Amphorophora ampullata Buckton SHG) es SE Amphorophora Buckton 1876 Neomacromyzus cyrtomicola Lee, new species (Figs. 1-13) Apterous viviparous.—Co/or (in life): Body sooty black, with pale white capitate or flabellate hairs. Legs pale. Siphunculus pale, except black basal part. Cauda black. Antenna pale, except black Ant.I-I], becoming = gradually dusky towards distal apex (Fig. 13). Color (in macerated specimens): head dark brown, antenna dark brown on Ant.I-II, pale brown on Ant.III-VI, de- veloping dusky color at distal apex of each segment. Clypeus, mandibular lam- inae and anteclypeus concolorous with head; rostrum pale, except dark distal end of URS. Thorax pale brown, except dark brown marginal sclerite on each segment. Legs pale, except dark brown coxae and pale brown tarsi. Abdomen pale brown with pigmented scleroites on each elevated base of setae. Genital plate brown. Cauda dark brown. Siphunculus pale on distal 2/3, dark brown on basal WEN Morphology: Body oval and small, 1.01—1.51 mm long from antennal tuber- cle to end of cauda. Head ‘strongly spinulate both dorsally and ventrally with 4 pairs of dorsal hairs, 3 pairs of ventral hairs, and frequently 1—2 addi- tional irregularly located hairs. Dorsal hairs on head capitate and 1.5—2.0 times as long as Ant.IIIBD. Antennal tubercles well developed and strongly granulated with a single hair on each side; inner side of each tubercle parallel (Fig. 9). Median tubercle of frons weakly developed. Antenna 1.45—1.84 mm long; Ant.I—An- t.VI, 0.07-0.11, 0.06—0.41, 0.29-0.44, 0.21-0.29, 0.20—0.30, (0.09-0.13) + (0.41—0.57) (Ant. VIb + PT), respectively. Ant.I-II granulated with 7-11 and 4-6 short blunt hairs, respectively. Ant.III PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON smooth on basal half and weakly imbri- cated on distal half, with 1 secondary rhinarium at base along with 8—15 short blunt hairs. Longest hair on antenna distinctly shorter than Ant.IIIBD. Ant.I1V—V imbricated; primary rhinar- ium on Ant.V ciliate; longest diameter of primary rhinarium as long as or slightly shorter than middle width of segment; Ant.VI strongly imbricated with 2-3 hairs on Ant.VIb; PT 3.37—5.42 times as long as Ant.VIb. Rostrum attaining hind coxae; mandibular laminae with 2— 4 long acuminate hairs on each side; ultimate rostral segment long, 1.50—2.38 times as long as 2HT, 1.00—1.45 times as long as Ant.VIb with two pairs of accessory hairs (Fig. 11). Thorax strong- ly wrinkled dorsally or reticulated mar- ginally with one pair of spinal hairs and 2 marginal hairs on each side; hairs short, clavate or flabellate, and based on elevated tuberclelike scleroites. Hind coxae spinulate with 7-8 long acuminate hairs. Trochanters and femora mostly smooth with ca. 35 short hairs; longest hair on hind femur 1/2 of basal diameter of femur. Hind tibia smooth, with acuminate hairs; longest hair on tibiae shorter than diameter of the middle. First tarsus smooth; first tarsal chaeto- taxy 3:3:3; 2HT weakly imbricated with 4-5 short acuminate hairs. Abdomen strongly wrinkled dorsally, spinulate ventrally. Dorsal hairs on abdomen short and strongly flabellate on anterior segments and clavate on posterior seg- ment, based on pigmented, elevated scleroites (Figs. 3-4). Tergum III with 3 pairs of dorsal hairs and 2-3 hairs on marginal sclerites. Tergum VI with 2 hairs between siphunculi. Tergum VII with 2 spinal hairs and 2-3 marginal hairs on each side. Tergum VIII with 4 acuminate or weakly capitate hairs. Siphunculus cylindrical, imbricate, well- flanged at apex, weakly tapering distally, 1.92—2.30 times as long as cauda (Fig. 5). Post-siphuncular sclerite dark-pigmented VOLUME 108, NUMBER 3 Table 1. Biometric data of apterous viviparous females of Neomacromyzus cyrtomicola. 499 Length of (mm) No. of hairs on No. of Rhinaria on Ratio (times) Apterous Vivipara (7 = 17) Part Mean Range Body from antennal tubercle to caudal end (BL) 1.28 (1.01—1.51) Entire antennae (A) 1.63 (1.45—1.84) Antennal segment I (Ant.1) 0.09 (0.07—0.11) Antennal segment IT (Ant.IT) 0.08 (0.06—-0.41) Antennal segment III (Ant.III) 0.38 (0.29-0.44) Antennal segment IV (Ant.IV) 0.25 (0.21—0.29) Antennal segment V (Ant.V) 0.25 (0.20—0.30) Base of antennal segment VI (Ant.VIb) 0.11 (0.09-0.13) Processus terminalis of Ant.VI (PT) 0.48 (0.41—0.57) URS (Ultimate Rostral Segment) 0.13 (0.12—0.14) Hind femur 0.53 (0.44—0.63) Hind tibia 0.96 (0.81—1.09) Second segment of hind tarsus (2HT) 0.07 (0.06—0.08) Siphunculus (SIPH) 0.26 (0.21—0.31) Cauda 0.12 (O.11—0.14) Setae on Ant.III 0.01 (0.01—0.02) Setae on Tergite VI 0.02 (0.02—0.03) Mandibular lamina 3 (2-4) Ant. 9 (7-11) Ant.II 4) (4-6) Ant. III 12. (8-15) URS 6 (4-6) Tergite VI between SIPH 2 (2-2) Tergite VIII 4 (3-4) Median of Genital plate (GP) 2 QD) Posterior margin of GP 11 (7-14) Cauda 5 (4-6) Ant. III 2 (= 3)) Entire Antennae (A) / Body Length (BL) 1.30 (@RO5=1888)) PT / Ant.VIb 4.42 (3.37—5.42) PT / Ant.III e277 (1.07—1.93) URS / 2HT 1.78 (1.50—2.38) URS / Ant. VIb 1.19 (1.00—1.45) SIPH / Body 0.21 (0.17—0.26) SIPH / Ant.III 0.69 (0.55—1.00) SIPH / Hind femur 0.49 (0.41—0.55) SIPH / Cauda Daley, (1.92—2.30) Cauda Length/Basal width 1.66 (1.36-1.96) Setae on Ant.III / Ant. IJIBD 0.78 (0.39-1.09) Setae on Tergite VI / Ant. IIIBD 1.45 (0.77-2.19) and strongly spinulate as ventral stridu- latory apparatus of Toxoptera. Cauda short triangular, constricted in middle, strongly spinulate dorsally and ventral- ly, with 4-6 long acuminate hairs (Fig. 12). Etymology.—The species name is de- rived from the genus name of the host plant Cyrtomium and the Latin suffix — cola (inhabitant of, dwelling in, living among). Biology.—All type specimens were collected on young, small plants of Cyrtomium falcatum which were living on rock walls in basalt caves on Jeju Island, Korea. No specimens have been found on large, full-grown plants of C. falcatum, although the other fern-infest- 500 ing aphid, Macromyzus woodwardiae (Takahashi, 1921), is very common on C. falcatum plants which grow in rather bright areas of the woody/shady places on Jeju Island. The colonies of WN. cyrtomicola were found on both the upper and lower sides of the young leaves and on young shoots that had not spread leaves, whereas M. woodwar- diae is found only on the shady underside of newly grown leaves of the larger plants. It seems that N. cyrtomicola is adapted to young, small plants found growing in dark and humid volcanic rock caves. Winged adults have not been observed. Presumably, they are monoe- cious and holocyclic on C. falcatum. Type material.—Holotype: apterous viviparous female, CALS SNU collec- tion specimen no. 020415-SH31/apt.17, KOREA(South), Jeyu-do (Island), Nam- jeju, Seongsan, Hlchul-bong, in volcanic rock cave, 17.iv.2002, on Cyrtomium falcatum, \eg. Seunghwan Lee. Para- types: 16 apterous viviparous females and three nymphs in same collection as holotype; five apterous viviparous fe- males and one nymph, CALS SNU collection specimen no. 040527-HJ-001, KOREA(South), Jeyu-do (Island), Nam- jeju, Seongsan, Illchul-bong, in volcanic rock cave, 27.v.2004, on C. falcatum, leg. Hyojoong Kim. The type specimens, including the holotype, are deposited in the College of Agriculture and Life Sciences, Seoul National University (CALS SNU), Seoul, Korea. Five para- types (collection number 020415-SH31) are also located at the Insect Museum of the National Institute of Agricultural Sciences and Technology (NIAST, Su- won), Korea. ACKNOWLEDGMENTS We are grateful to Mr. Wonhoon Lee and Mrs. Youngbun Lee (College of Agriculture and Life Sciences, Seoul National University, Korea) for pre- paring the electronic illustrations and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON their technical assistance making slide preparations of aphids and measuring the morphological characters. This study was supported by the Brain Korea 21 program and Seoul National Univer- sity. LITERATURE CITED Blackman, R. L. and V. F. Eastop. 2000. Aphids on the World’s Crops. An Identification and Information Guide. 2nd edition. The Natural History Museum, London, United Kingdom. 466 pp. Buckton, G. B. 1876. Monograph of the Brit- ish Aphides, Vol. I. Ray Society, London. 190 pp. David, S. K. 1958. A new genus and three new species of aphids from India. Indian Journal of Entomology 20: 175-180. David, S. K. and K. Narayana. 1968. Three new species of aphids from South Western Hima- layas in India. Bulletin of Entomology 9: 99-103. Ghosh, A. K. 1974. Fern Infesting aphids (Insecta: Homoptera) in India. Indian Journal of Horticulture 31(1): 104—109. Ghosh, M. R., R. C. Basu, and D. N. Raychaud- huri. 1977. Studies on the aphids (Homoptera: Aphididae) from eastern India. XXXV. Three new genera and four new species from north- ern India. Oriental Insects 11(4): 579-586. Lee, S. 2002. New Record of Macromyzus wood- wardiae (Sternorrhyncha, Aphididae) on Cy- rtomium falcatum from Korea. The Korean Journal of Systematic Zoology 18(1): 127— (13)3}. Lee, S., J. Holman, and J. Havelka. 2002. Illustrated catalogue of Aphididae in the Korean peninsula. Part I. Subfamily Aphidi- nae. /n Park, K. T., ed. Insects of Korea, Vol. 9). 3228) joy oy. Miyazaki, M. 1968. A revision of the fern aphids of Japan with descriptions of three new species (Homoptera: Aphididae). Insecta Matsumur- ana 31(3): 13-24. ———. 1971. A revision of the tribe Macrosiphini of Japan (Homoptera: Aphididae, Aphidinae) Insecta Matsumurana 34(1): 1—247. Mordvilko, A. K. 1929. Aphids, pp. 163-204. In Philipjew, J. N., ed. Key for the Identification of Russian Insects. Moscow. Moritsu, M. 1952. The fern aphids in Japan. Bulletin of the Faculty of Agriculture, Yama- guchi University 2: 26-28. Shinji, O. 1929. Four new genera of Aphididae from Morioka, Japan. Lansania 1(3): 39-48. VOLUME 108, NUMBER 3 Takahashi, R. 1921. Aphididae of Formosa. Part I. Agricultural Experimental Station Govern- ment Formosa Report 20: 1—97. 1925. Aphididae of Formosa. Part IV. Department of Agriculture, Government Re- search Institute, Formosa, Report 16: 1—65. . 1938. List of the aphid genera proposed in recent years (Hemiptera). Tenthredo 2(1): 1-18. 501 . 1960. Four new genera of Aphididae from Japan. Kontyu 28(4): 223-229. Tao, C. C. 1963. Revision of Chinese Macrosiphi- nae (Aphidae, Homoptera). Plant Protection Bulletin (Taiwan) 5(3): 162-205. van der Goot, P. 1917. Notes on some Indian aphides. Records of the Indian Museum 13: 175-183. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 502-510 FURTHER NEW GENUS-GROUP NAMES IN THE CICADELLIDAE (HEMIPTERA) STUART H. MCKAMEyY Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 168, Washington, D.C. 20560-7012, U.S.A. (e- mail: smckamey@sel.barc.usda.gov) Abstract.—New replacement names are proposed for 11 genera or subgenera of leafhoppers in the subfamilies Coelidiinae, Ledrinae, Macropsinae, Scarinae, Typhlocybinae, and Ulopinae: Davduospina for Duospina Davies, Dialodia for Lodia Nielson, Doowella for Woodella Evans, Hobemanella for Bohemanella DeLong & Freytag (subgenus of Polana), Latycephala for Platycephala, Lesinda for Selinda Theron, Lonnia for Nollia Hamilton, Negoneura for Genoneura Dlabola, Olidiana for Lodiana Nielson, Ruana for Urana Coelho & Nessimian (subgenus of Gypona), and Varpulana for Parvulana DeLong & Freytag (subgenus of Polana). The new generic names circumscribe 116 valid species, 93 of which are here proposed in new combinations. Twenty-two of the remainder are in preoccupied scarine subgenera. Lodiana pectinata Yang & Zhang (1995) is preoccupied by L. pectinata Nielson (1982)—the new replacement name Olidiana yangi McKamey is proposed to replace the junior primary homonym. One of the new combinations involves a known vector of phytopathogens, the sandal leafhopper (now Olidiana indica [Walker]). The spellings of Humpatagallia Linnavuori & Viraktamath (Agalliinae) and Limassolla Dlabola (Typhlocybinae) are fixed as correct. Key Words: \eafhoppers, Homoptera, Auchenorrhyncha, new genus, sandal spike vector Numerous generic homonyms were discovered while preparing a database of world leafhoppers. Eleven new generic names were previously proposed for homonyms in the subfamilies Deltoce- phalinae and Selenocephalinae (McKa- mey 2003). This is the second in a series of new generic names and includes all remaining subfamilies except Evacanthi- nae and Idiocerinae, which will be dealt with separately. Two leafhopper genera were spelled multiple ways in their original descrip- tions: Humpatagallia Linnavuori & Vir- aktamath (1973a: 485; Agalliinae) and Limassolla Diabola (1965e: 663; Typhlo- cybinae). The principle of first revisor (ICZN; International Commission on Zoological Nomenclature 1999, Art. 24) is here applied to fix the above original spellings as correct, and the authorship is unchanged. The alternate spellings, Humptagallia and Limasolla, are incor- rect. The correct combination of the sandal leafhopper, an important vector of san- dal spike disease in Myanmar and India (Nielson 1968b), is now Olidiana indica VOLUME 108, NUMBER 3 (Walker). Out-dated combinations ap- pearing in literature include Coelidia indica or Lodiana indica. The taxa are arranged alphabetically within their réspective higher taxon. Letter designations for literature cita- tions through 1985 are consistent with the bibliographies by Metcalf (1964a) and Oman et al. (1990). SUBFAMILY COELIDIINAE Dialodia McKamey, new name Dialodia, nomen novum for Lodia Nielson 1932e: 222, preoccupied by Lodia Baird & Girard 1853. Type species by original designation: Lodia_ glabrosa Nielson. Dialodia alyssae, new combination Lodia alyssae Nielson 1996: 149 [n. sp.]. Dialodia glabrosa, new combination Lodia glabrosa Nielson 1982e: 224 [n. sp:|: Dialodia negans, new combination Lodia negans Nielson 1982e: 227 [n. sp.]. Dialodia parapectinata, new combination Lodia parapectinata Nielson 1988: 240 fosp- |: Dialodia pectinata, new combination Lodia pectinata Nielson 1982e: 225 [n. sp.]. Dialodia proxima, new combination Lodia proxima Nielson 1982e: 227 [n. sp.]. Lesinda McKamey, new name Lesinda, nomen novum for Selinda heron, 1986: 161, preoccupied, .by Selinda Moore 1883. Type species by original designation: Selinda bruggeni Theron. Lesinda bruggeni, new combination 503 Selinda bruggeni Theron 1986: 161 [n. sp.]. Olidiana McKamey, new name Olidiana, nomen novum for Lodiana Nielson 1982e: 86, preoccupied by Lodiana Ragonot 1888. Type species by original designation: Lodiana alata Nielson. Olidiana alata, new combination Lodiana alata Nielson 1982e: 115 [n. Sp:l: Olidiana alvea, new combination Lodiana alvea Nielson 1991: 461 [n. Sp. |: Olidiana aperta, new combination Lodiana aperta Nielson 1982e: 103 [n. sp.]. Olidiana bedardi, new combination Lodiana bedardi Nielson 1982e: 128 [n. sp.]. Olidiana bifurcata, new combination Lodiana bifurcata Nielson 1982e: 117 [fn SpE Olidiana bigemina, new combination Lodiana bigemina Zhang 1990: 105 [n. sp.]. Olidiana biungulata, new combination Lodiana biungulata Nielson 1982e: 113 [n. sp.]. Olidiana boninensis, new combination Coelidia boninensis Matsumura 1914b: o2 [me spsl: Lodiana boninensis: Nielson 1982e: 134 [n. comb.]. Olidiana brevis, new combination Tettigonia brevis Walker 1851b: 774 [n. Spel: Lodiana brevis: Nielson 1982e: 132 [n. comb.]. Olidiana brevisina, new combination 504 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Lodiana brevisina Zhang 1990: 104 [n. Spl: Olidiana brevissima, new combination Lodiana brevissima Zhang 1990: 106 invisp.|. Olidiana cladopenis, new combination Lodiana cladopenis Zhang 1990: 101 lin: seal: Olidiana corneola, new combination Lodiana corneola Nielson 1982e: 114 [ne-sp:i. Olidiana ctenostyla, new combination Lodiana ctenostyla Nielson 1991: 457 [me Spoe|: Olidiana cupraria, new combination Coelidia cupraria Walker 1857b: 173 paves) oe) Lodiana cupraria: Nielson 1982e: 108 [n. comb.]. Olidiana curta, new combination Lodiana curta Nielson 1982e: 91 [n. sp.]. Olidiana curvispinata, new combination Lodiana curvispinata Zhang 1994: 81 fine Spel: Olidiana fasciculata, new combination Lodiana fasciculata Nielson 1982e: 99 [me spel Olidiana fissa, new combination Lodiana fissa Nielson 1991: 460 [n. sp.]. Olidiana flavofascia, new combination Lodiana flavofascia Zhang 1994: 97 [n. sp.]. Olidiana fringa, new combination Lodiana fringa Zhang 1994: 104 [n. sp.]. Olidiana furcata, new combination Lodiana furcata Nielson 1991: 453 [n. sp-l: Olidiana genista, new combination Lodiana genista Nielson 1982e: 129 [n. sp.]. Olidiana gladia, new combination Lodiana gladia Nielson 1991: 458 [n. sp.]. Olidiana hamularis, new combination Lodiana hamularis Xu 2000a: 219 [n. sp.]. Olidiana huangi, new combination Lodiana huangi Zhang 1994: 80 [n. sp.]. Olidiana huoshanensis, new combination Lodiana huoshanensis Zhang 1994: 86 poser Olidiana indica, new combination Coelidia indica Walker 1851b: 855 [n. sp.]. Lodiana indica: Nielson 1982e: 137 [n. comb.]. Olidiana know!/toni, new combination Lodiana knowltoni Nielson 1982e: 101 roles) 6 Olidiana kodeti, new combination Lodiana kodeti Nielson 1982e: 124 [n. sp.]. Olidiana kuohi, new combination Lodiana kuohi Xu 2000a: 221 [n. sp.]. Olidiana lamina, new combination Lodiana lamina Nielson 1982e: 91 [n. sp.]. Olidiana laminapellucida, new combina- tion Lodiana laminapellucida Zhang 1994: 104 [n. sp.]. Olidiana laminispinosa, new combination Lodiana 1994: LOS [in -spl: Olidiana longilamina, new combination laminispinosa Zhang VOLUME 108, NUMBER 3 Lodiana longilamina Zhang 1994: 88 [n. sp.]. Olidiana marginifrons, new combination Coelidia marginifrons Walker 1870b: S10) asp. |: Lodiana marginifrons: Nielson 1982e: 96) |xemst:; n_ comb:|: Olidiana mecistenata, new combination Lodiana mecistenata Yang [in Yang & Aman, 1995: 412 m.. .sp-|: Olidiana munda, new combination Lodiana munda Nielson 1982e: 107 [n. sp.]. Olidiana mutabilis, new combination Lodiana mutabilis Nielson 1982e: 120 [n. sp.]. Olidiana neilsoni, new combination Lodiana neilsoni Zhang 1994: 76 [n. sp.]. : Olidiana nigridorsum, new combination Lodiana nigridorsum Cai & Shen 1998: 42 [n. sp.]. Olidiana nocturna, new combination Jassus nocturnus Distant 1908g: 332 [n. sp.]. Lodiana nocturna: Nielson 1982e: 137 [n. comb.]. Olidiana opulenta, new combination Jassus opulentus Distant 1908g: 336 [n. sp.]. Lodiana opulenta: Nielson 1982e: 134 [n. comb.]. Olidiana ornata, new combination Lodiana ornata Nielson 1982e: 127 [n. Spal Olidiana parapectinata, new combination Lodiana_ parapectinata Nielson 1991: 456 [n. sp.]. Olidiana pectinata, new combination Lodiana pectinata Nielson 1982e: 110 (aS: | 505 Olidiana pectiniformis, new combination Lodiana pectiniformis Zhang 1994: 87 [n. sp.]. Olidiana pectita, new combination Jassus pectitus Distant 1908g: 329 [n. sp.]. Lodiana pectita: Nielson 1982e: 99 [n. comb.]. Olidiana peniculata, new combination Lodiana peniculata Nielson 1982e: 111 [n-sSp-): Olidiana perbrevis, new combination Lodiana perbrevis Nielson 1991: 453 frat, Sy 54) b Olidiana perculta, new combination Jassus percultus Distant 1908g: 337 [n. sp.]. Lodiana perculta: Nielson 1982e: 131 [n. comb.]. Olidiana platyfasciata, new combination Lodiana platyfasciata Xu 2000a: 218 [ime sp=l- Olidiana polyspinata, new combination Lodiana polyspinata Zhang 1994: 107 [n. sp.]. Olidiana praetextus, new combination Jassus praetextus Distant 1908g: 338 SS: Lodiana_ praetextus: 122 [n.scomb#l. Olidiana recurvata, new combination Nielson 1982e: Lodiana recurvata Nielson 1998: 129 irasis | Olidiana reidi, new combination Lodiana reidi Nielson 1982e: 106 [n. sp.]. Olidiana ritcheri, new combination Lodiana ritcheri Nielson 1982e: 118 [n. sp.]. Olidiana ritcheriina, new combination 506 Lodiana ritcheriina Zhang 1990: 102 [n. sp.]. Olidiana scopae, new combination Lodiana scopae Nielson 1982e: 124 [n. sp.]. Olidiana scutopunctata, new combination Lodiana scutopunctata Zhang 1994: 83 lia: sips Olidiana serra, new combination Lodiana serra Nielson 1982e: 97 [n. Spe: Olidiana setacea, new combination Lodiana setacea Nielson 1982e: 120 [n. Spel: Olidiana signata, new combination Lodiana signata Zhang 1994: 91 [n. sp.]. Olidiana singularis, new combination Lodiana singularis Nielson 1991: 454 [n. sp.]. Olidiana spiculata, new combination Lodiana spiculata Nielson 1982e: 93 [n. Spel: Olidiana spina, new combination Lodiana spina Zhang 1990: 103 [n. sp.]. Olidiana spira, new combination Lodiana spira Nielson 1991: 458 [n. spel: Olidiana tantula, new combination Lodiana tantula Nielson 1991: 459 [n. Sa: Olidiana tongmaiensis, new combination Lodiana tongmaiensis Zhang 1994: 79 [ne -sp.]- Olidiana uenoi, new combination Lodiana uenoi Hayashi 1995: 197 [n. sp.]. Olidiana unica, new combination Lodiana unica Nielson 1982e: 95 [n. sp.]. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Olidiana ventrosola, new combination Lodiana ventrosola Nielson 1982e: 103 [n. sp.]. Olidiana viraktamathi, new combination Lodiana_ viraktamathi Nielson 1991: 455 [n. sp.]. Olidiana xanthopronotata, new combina- tion Lodiana xanthopronotata Zhang 1994: Tapes Olidiana zhengi, new combination Lodiana zhengi Zhang 1994: 106 [n. sp.]. Olidiana yangi McKamey, new name Lodiana_ pectinata Yang & Zhang 1995: 40: mn. sp. Preoccupied® by 7 pectinata Nielson (1982e). SUBFAMILY LEDRINAE Latycephala McKamey, new name Latycephala, nomen novum for Platyce- phala Ge 1992: 250, preoccupied by Platycephala Fallén 1820. Type species by original designation: Platycephala tortilla Ge. Latycephala decussata, new combination Platycephala decussata Ge 1992: 252 [n. sp.]. Latycephala graminea, new combination Platycephala graminea Ge 1992: 253 [me Spel: Latycephala laminata, new combination Platycephala laminata Ge 1992: 253 [n. sp.]. Latycephala tortilla, new combination Platycephala tortilla Ge 1992: 251 [n. sp.]. SUBFAMILY MACROPSINAE Lonnia McKamey, new name Lonnia, nomen novum for Nollia Hamil- ton 1983d: 19, preoccupied by Nollia VOLUME 108, NUMBER 3 Eisenack 1979. Type species by origi- nal designation: Neopsis pallidus Lin- navuori & DeLong 1977c. Lonnia pallida, new combination Neopsis pallidus Linnavuori & De- one 197 7ce> l/S*(ne sp:|: Nollia pallida: Hamilton 1983d: 19 [n. comb.]. Lonnia rustica (Hamilton), new combina- tion Nollia rustica Hamilton 1983d: 20 [n. comb.]. SUBFAMILY SCARINAE A historical, perhaps more familiar name for Scarinae is Gyponinae. Hobemanella McKamey, new subgeneric name Hobemanella, nomen novum for Polana (Bohemanella) DeLong & Freytag 1972f: 242, preoccupied by Bohema- nella Ramme 1951. Type species by original designation: Gypona bohemani Stal. Polana_ (Hobemanella) subgeneric combination bohemani, new Gypona bohemani Stal 1864a: 81 [n. sp.]. Polana (Bohemanella) bohemani: De- Monge &oeFreytag 1972f: 247 [n. subg. comb. | Polana (Hobemanella) alia, new subgene- ric combination Polana (Bohemanella) alia DeLong & Freytag 1972f: 247 [n. sp.]. Polana (Hobemanella) ardua, new sub- generic combination Polana (Bohemanella) ardua DeLong arreyiae W972i 253) (nm. sp: Polana (Hobemanella) assula, new sub- generic combination Polana (Bohemanella) assula DeLong & Freytag 1972f: 249 [n. sp.]. 507 Polana (Hobemanella) celsa, new subge- neric combination Polana (Bohemanella) celsa DeLong & Freytag 197212 250i |msp: |: Polana (Hobemanella) chelata, new sub- generic combination Polana (Bohemanella) chelata DeLong & Freytag 1972f: 244 [n. sp.]. Polana (Hobemanella) chifama, new sub- generic combination Polana_ (Bohemanella) chifama_ De- Long & Freytag 1972f: 250 [n. sp.]. Polana (Hobemanella) elabora, new sub- generic combination Polana (Bohemanella) elabora DeLong é& Freytac. (9721-3252) it. Spl: Polana (Hobemanella) macula, new sub- generic combination Polana (Bohemanella) macula DeLong & Preytagel9 726252) |nspul: Polana (Hobemanella) orbita, new sub- generic combination Polana (Bohemanella) orbita DeLong & Freytag 1972t 252m. sp.4. Polana (Hobemanella) resilara, new sub- generic combination Polana (Bohemanella) resilara DeLong & Freytag 1972f: 249 [n. sp.]. Polana (Hobemanella) resupina, new sub- generic combination Polana_ (Bohemanella) resupina De- Long & Freytag 1972f: 244 [n. sp.]. Polana (Hobemanella) retenta, new sub- generic combination Polana (Bohemanella) retenta DeLong & Freytag 1972f: 247 [n. sp.]. Polana (Hobemanella) scela, new subge- neric combination Polana (Bohemanella) scela DeLong & Freytag 1972f 253: [nesp*|: Polana (Hobemanella) belema, new sub- generic combination 508 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Polana (Bohemanella) belema DeLong 1984e: 45 [n. sp.]. Polana_ (Hobemanella) subgeneric combination luteonota, new Polana (Bohemanella) luteonota De- onpel979e7t54.[nr sp: Polana (Hobemanella) putara, new sub- generic combination Polana (Bohemanella) putara DeLong OVS ae 29S nes pale Ruana McKamey, new subgeneric name Ruana, nomen novum for Gypona (Urana) Coelho’ & Nessinuanm 19915 5575" pre- occupied by Urana Melichar 1901. Type species by original designation: Gypona (Urana) urata Coelho & Nes- simian. Gypona (Ruana) urata, new subgeneric combination Gypona (Urana) urata Coelho & Nes- sian 99-7557 ine Sp.t. Varpulana McKamey, new subgeneric name Varpulana, nomen novum for Polana (Parvulana) DeLong & Freytag 1972f: 289, preoccupied by Parvulana Fagot 1891. Type species by original desig- nation: Polana (Parvulana) alata De- Long & Freytag. Polana (Varpulana) alata, new subgeneric combination Polana (Parvulana) alata DeLong & Breytag, 19728, 292. [m., sp.). Polana (Varpulana) bidens, new subgene- ric combination Polana (Parvulana) bidens DeLong & Freytag 19721: 290) [nasp.|- Polana (Varpulana) pandara, new subge- neric combination Polana (Parvulana) pandara DeLong & Freytag 1972f 290:{meisp:): Polana (Varpulana) tulara, new subgene- ric combination Polana (Parvulana) tulara DeLong & Ereytas 1972 290 |n- spl SUBFAMILY TYPHLOCYBINAE Negoneura McKamey, new name Negoneura, nomen novum for Genoneura Dlabola 1994: 78, preoccupied by Genoneura Matthew 1889. Type spe- cies by original designation: Genoneura alboviridis Dliabola. Negoneura alboviridis, new combination Genoneura alboviridis Diabola 1994: 80 paleersy 05) 5 SUBFAMILY ULOPINAE Davduospina McKamey, new name Davduospina, nomen novum for Duospina Davies 1988: 58, preoccupied by Duos- pina Hodges 1966. Type species by original designation: Cephalelus capen- sis Evans. Davduospina capensis, new combination Cephalelus capensis Evans 1947c: 145 [ne spel: Duospina capensis: Davies 1988: 58 [n. comb.]. Davduospina sheilae, new combination Duospina sheilae Davies 1988: 62 [n. sp.]. Dayduospina twanella, new combination Duospina twanella Davies 1988: 60 [n. sp.]. Doowella McKamey, new name Doowella, nomen novum for Woodella Evans 1966a: 87, preoccupied by Woo- della Haque 1956. Type species by original designation: Woodella wanun- garae Evans. Doowella wanungarae, new combination Woodella wanungarae Evans 1966a: 87 fake fo) os VOLUME 108, NUMBER 3 ACKNOWLEDGMENTS J. Dorsey assisted in the search for generic homonyms and M. Nielson encouraged replacement of coelidiine homonyms. 1... Deitz, F.C. Thomp- son, and two anonymous reviewers pro- vided helpful comments on an earlier draft of the manuscript. LITERATURE CITED Baird, S. F. and C. Girard. 1853. Smithsonian Miscellaneous Collections 2(5): 116 [original reference not seen, citation of Reptilia, follow- ing Neave]. Cai, P. and Shen, X. 1998. New species of family Cicadellidae from Mt. Funiu in Henan (Homoptera: Cicadelloidea). Fauna and Tax- onomy of Insects in Henan 2: 37-52. Coelho, L. B. N. and J. L. Nessimian. 1991. Descrigao de duas novas espécies e de um novo subgenero de Gypona Germar (Homoptera, Cicadellidae, Gyponinae). Revista Brasileira Biologia 51(3): 555—559. Davies, D. M. 1988. Leafhoppers (Homoptera: Cicadellidae) associated with the Restoniaceae. 1. The tribe Cephalelini (Ulopinae). Journal of the Entomological Society of South Africa 51(1): 31-64. DeLong, D. M. 1979d. 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Erganzungen zur iranischen, israe- lischen und benachbarten Zikadenfaunen mit Bescheibungen 30 neuer Taxone (Homoptera, 509 Auchenorrhyncha). Sbornik Narodniho Mu- sea V. Praze Rada B Prirodni Vedy 49(1—4): 41-110. Eisenack, A. 1979. Nollia eifelensis (Acritarcha), ein Pyrocystis-ahnliches Mikrofossil aus dem Unter-Devon der Eifel. Senckenbergiana lethaea 60(1—3): 52. Evans, J. W. 1947c. Some new Ulopinae (Homo- ptera, Jassidae). Annals and Magazine of Natural History 14(11): 140-150. . 1966a. The leafhoppers and froghoppers of Australia and New Zealand (Homoptera: Cicadelloidea and Cercopoidea). Memoirs of the Australian Museum 12: 1—347. Fagot, P. 1891. Histoire malacologique des Pyr- énées Frangaises et Espagnoles. Liste, & c. Bulletin du Société de Ramond 26(3): 226. Fallén, K. F. 1820. Oscides Sveiciae 2: 1-10 [original reference not seen, citation of Co- leoptera, following Neave]. Ge, Z. 1992. Homoptera: Cicadelloidea, pp. 243— 316. In Chen, S., ed. Insects of the Hengduan Mountains Region, Vol. 1. Science Press, Being. 865 pp. Hamilton, K. G. A. 1983d. Revision of the Macropsini and Neopsini of the New-World (Rhynchota: Homoptera: Cicadellidae), with notes on intersex morphology. Memoirs of the Entomological Society of Canada 123: 1-223. Haque, A. F. M. 1956. The foraminifera of the Ranikot and the Laki of the Nammal Gorge, Salt Range. Memoirs of the Geological Survey of Pakistan. Palaeontologia Pakistanica 1: 1— 300. Hayashi, M. 1995. A new species and a new record of coelidiine leafhoppers (Homoptera, Cica- dellidae) from the Ryukyu Islands, Japan. Special Bulletin of the Japanese Society of Coleopterology 4: 197-202. Hodges, R. W. 1966. Review of New World species of Batrachedra, with description of three new genera (Lepidoptera: Gelechioidea). Transac- tions of the American Entomological Society 92: 585-651. International Commission on Zoological Nomen- clature. 1999. International Code of Zoolog- ical Nomenclature. 4th ed. International Trust for Zoological Nomenclature, London. xxix + 306 pp. Linnavuori, R. E. and D. M. Delong. 1977c. The leafhoppers (Homoptera: Cicadellidae) known from Chile. Brenesia 12&13: 163-267. Linnavuori, R. E. and C. A. Viraktamath. 1973a. A new cicadellid genus from Africa. Revue de Zoologie et de Botanique Africaines 87(3): 485-492. 510 Matthew, G. F. 1889. Transactions of the Royal Society of Canada 6 (1888, section 4): [49]-62. plate IV [original reference not seen, citation from L. L. Deitz, pers. comm.]. Matsumura, S. 1914b. Die Coelidinen Japans. Transactions of the Sapporo Natural History Society 5: 81—90. McKamey, S. H. 2003. Some new generic names in the Cicadellidae (Hemiptera: Deltocephalinae, Selenocephalinae). Proceedings of the Ento- mological Society of Washington 105(2): 447-451. Melichar, L. 1901. Monographie der Acanalonii- den und Flatiden (Homoptera). Annales des k. k. Naturhistorischen Hofmuseums 16: 178— 258. Metcalf, Z. P. 1964a. General Catalogue of the Homoptera. Fascicle VI. Cicadelloidea. Bibli- ography of the Cicadelloidea (Homoptera: Auchenorrhyncha). United States Department of Agriculture, Agriculture Research Service, 349 pp. Moore, F. A. 1883. A monograph of Limnaina and Euploeinae, two groups of diurnal Lepidoptera belonging to the sub-family Euploeinae, with descriptions of new genera and species. Pro- ceedings of the Zoological Society of London 1883: 201-324. Nielson, M. W. 1968b. The leafhopper vectors of phytopathogenic viruses (Homoptera, Cicadel- lidae). Taxonomy, biology and virus trans- mission. United States Department of Agri- culture Technical Bulletin 1382, 386 pp. 1982e. A_ revision of the subfamily Coelidiinae (Homoptera: Cicadellidae). IV. Tribe Coelidini. Pacific Insects Monograph 38: 1-318. 1988. New genera and new species of Neotropical Coelidiini (Homoptera: Cicadelli- dae: Coelidiinae). Great Basin Naturalist 48(2): 232-349. . 1991 [dated 1990]. New Oriental leafhop- pers in the tribe Coelidiini with a revised key to the species (Homoptera: Cicadellidae: Coeli- diinae). Entomologica Scandinavica 21(4): 445-475. 1996. Two new genera and some new species of Neotropical and Oriental leafhop- pers in the tribe Coelidiini (Homoptera: Cicadellidae). Transactions of the American Entomological Society 122(2—3): 147-160. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 1998. New species of coelidiine leafhoppers from southern China (Homoptera: Cicadelli- dae). Mitteilungen des Museum ftir Natur- kunde Berlin Deutsche Entomologische Zeits- chrift 45(2): 129-130. Oman, P. W., W. J. Knight, and M. W. Nielson. 1990. Leafhoppers (Cicadellidae): a Bibliogra- phy, Generic Check-list and Index to the World Literature 1956-1985. CAB Interna- tional Institute of Entomology, Wallingford, UK. [iv] + 368 pp. Ragonot, E. L. 1888. Nouveaux genres et espéces de Phycitidae et Galleriidae. Paris. 52 pp. Ramme, W. 1951. Zur Systematik, Faunistik und Biologie der Orthopteren von Stidost-Europa und Vorderasien. Mitteilungen aus dem Zool- ogischen Museum in Berlin 27: 1-431. Stal, C. 1864a. Hemiptera Mexicana enumeravit speciesque novas descripsit (cont.). Entomolo- gische Zeitung. Herausgegeben von dem En- tomologischen Vereine zu Stettin 25: 49-86. Theron, J. G. 1986. New genera and species of southern African Coelidiinae (Homoptera: Cicadellidae),with description of the new tribe Equeefini. Phytophylactica 18(4): 153-163. Walker, F. 185lb. List of the Specimens of Homopterous Insects in the Collection of the British Museum 3: 637-907. 1857b. Catalogue of the homopterous insects collected at Sarawak, Borneo, by Mr. A. R. Wallace, with descriptions of new species. Journal of the Proceedings of the Linnaean Society of London 1: 141-175. 1870b. Catalogue of the homopterous insects collected in the Indian Archipelago by Mr. A. R. Wallace, with descriptions of new species. Linnean Society of London Journal of Zoology 10: 276-330. Xu, G. 2000. Three new species of Lodiana (Homoptera: Cicadellidae) from China. En- tomologia Sinica 7(3): 218—222. Yang, J. and Zhang, L. 1995. Homoptera: Cica- delloidea, pp. 37-44. In Zhu, T., ed. Insects and Macrofungi of Guitanshan, Zhejiang. Zhejiang Science and Technology Publishing House. 318 pp. Zhang, Y. 1990. A Taxonomic Study of Chinese Cicadellidae (Homoptera). Tianze Eldonejo, Yangling 1990: 1-218. 1994. A Taxonomic Study of Chinese Coelidiinae (Homoptera: Cicadellidae). Henan Science and Technology Press, 151 pp. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 511-518 THE THIRD-STAGE LARVA AND PUPA OF DEMODEMA BREVITARSIS (BLANCHARD) (COLEOPTERA: SCARABAEIDAE: MELOLONTHINAE) FROM SOUTHERN BRAZIL MIGUEL ANGEL MORON AND JOSE ROBERTO SALVADORI (MAM) Departamento de Entomologia, Instituto de Ecologia, A.C. Apdo. Postal 63, Xalapa, Veracruz 91000, México (e-mail: moron_ma@ecologia.edu.mx):; (JRS) Embrapa- Centro Nacional de Pesquisa de Trigo, Caixa Postal 451.99001-970 Passo Fundo, Rio Grande do Sul, Brazil (e-mail: jrsalva@cnpt.embrapa.br) Abstract.—The third-instar larva and pupa of Demodema brevitarsis (Blanchard) from Tapejara, Rio Grande do Sul, Brazil, are described. This is the first description of immature stages in this genus. A key to the larvae of seven genera of New World Macrodactylini (sensu Jato), is given. Key Words: Demodema, larva, pupa, taxonomy, Macrodactylini, key, America The genus Demodema Blanchard of the Macrodactylini consists of eight species distributed from southern Brazil, Bolivia and Paraguay to Argentina (Evans 2003). Adults of Demodema species are commonly attracted to lights, but nothing is known about their habits and life cycle. From 1999 to 2001, the second author (JRS) collected white grubs feeding on soybean roots near Tapejara, Rio Grande do Sul, Brazil, and after rearing in the laboratory he obtained chafers identified as D. brevi- tarsis (Blanchard). In this paper we describe for the first time the third-instar larva and pupa of Demodema_ brevitarsis. Technical terms are those of Ritcher (1966) and Moron (1986). Voucher specimens are deposited in the Entomological Collection Instituto de Ecologia, Xalapa, Mexico (IEXA) and in the Insect Collection of Centro Nacional de Pesquisa de Trigo-Embrapa, Passo Fundo, RS, Brazil (CNPT). Of the 1,066 species and 57 genera of Macrodactylini listed from the New World (Evans 2003), the larvae of only 10 species in six genera are described at present. Consequently, it is difficult to establish a set of diagnostic characters that distinguish the larvae of Macrodac- tylini from larvae of other tribes of Melolonthinae. However, based on Ritcher (1966), Costa et al. (1988), and our own studies, we propose the follow- ing combination of characters as di- agnostic for larvae of Macrodactylini: head without ocelli (except in some Macrodactylus); mandibles without ven- tral stridulatory area; haptomerum usu- ally present, epipharynx with 2 to 6 heli; lacinia with a row of 3 unci usually longitudinal; maxillary stridulatory area without anterior process; last antennal segment with a single, dorsal sensory spot; respiratory plates of spiracles not constricted (except in some Macrodacty- lus); raster with longitudinal palidia or with palidia absent; anal slit usually angulate or Y-shaped; lower anal lip with sagittal cleft or groove; each tarsal claw bearing 2 setae; and claws of ale PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mesothoracic and metathoracic legs sometimes reduced in size. KEY TO THE KNOWN THIRD-INSTAR LARVAE OF THE GENERA OF MACRODACTYLINI (SENSU EVANS 2003) FROM THE AMERICAS (MODIFIED FROM RITCHER 1966 AND COSTA ET AL. 1988) 1. Tormae united. Raster lyre-shaped and WITROMGIDAlIGIAN 4 me Meta key an te oe ae 2 — Tormae not united (Fig. 3). Raster usually not lyre-shaped, with or without palidia. ... 3 2. Raster with dense triangular patch of very short, stout conical setae on each side. Hap- tomerum of epipharynx with a transverse row of 2to4heli. ...... Dichelonyx Harris — Raster with setae rather uniformly dis- tributed. Haptomerum often absent. Coenonycha Horn 3. Haptomerum of epipharynx with 4 heli. All legs with well developed claws. Abdominal spiracles with lobes of respiratory plate constricted, surrounding bulla. Raster with a pair of short, longitudinal, parallel palidia. Macrodactylus Latreille — Haptomerum of epipharynx with 5 or 6 heli (Fig. 3). Claws of mesothoracic and/or metathoracic legs reduced in size (Fig. 14). Abdominal spiracles with lobes of respira- tory plate curved but not constricted around bulla (Fig. 15). Raster with single palidium on each side of septula, usually not parallel (Fig. 16). 4. All abdominal spiracles similar in size. ... 5 — Abdominal spiracles 6 to 8 smaller in size than spiracles 1—5 (Fig. 1). 5. Raster with a pair of palidia parallel anteri- orly and widely divergent posteriorly. Palli- dia monostichous anteriorly and _ polysti- chous along lower anal lobes. Thoracic spiracles much larger than abdominal spira- cles, with lobes of respiratory plate directed POStchiOnlyA eee ee Plectris Serville — Raster with a pair of palidia moderately divergent posteriorly. Pallidia entirely monostichous. Thoracic spiracles with lobes of respiratory plate directed ventrally TIsonychus Mannerheim 6. Claws of prothoracic legs long and sharp, claws of mesothoracic legs — slightly shorter, claws of metathoracic legs min- ute. Raster with pair of longitudinal palidia that are not divergent posteri- orly. Anoplosiagum Blanchard — Claws of prothoracic and mesothoracic legs long and sharp, claws of metathoracic legs shorter (Figs. 12-14). Raster with a pair of monostichous palidia that are nearly par- allel anteriorly and widely divergent post- eriorly (Fig. 16). .... Demodema Blanchard LARVAE OF DEMODEMA BLANCHARD, 1850 The larval description of Demodema brevitarsis (Blanchard) 1s the first for the genus. Based on current knowledge of the larvae of Macrodactylini (sensu Jato), the larva of Demodema are most similar morphologically to those of Anoplosia- gum species. The known larva of Demo- dema have one pair of palidia widely divergent posteriorly, each palidium con- sisting of a row of 25-27 pali. The known larva of Anoplosiagum have one pair of longitudinal, feebly curved pali- dia, each palidium consisting of a row of 10-13 pali. Larvae of both genera have the abdominal spiracles 6 to 8 smaller than the spiracles 1-5, and the hapto- merum of epipharinx with 6 stout hell. Demodema brevitarsis (Blanchard, 1850) (Figs. 1-16) Third-instar larva.—This description is based on 32 third instar larvae collected in soybean fields. Locality data: Brazil: State of Rio Grande do Sul, Tapejara (28°03'14"S; 52°07'20”"W), =17-I-2001, 630 m altitude, J.R. Salvadori (12 larvae) (IEXA); same data (20 larvae) (CNPT). Body (Fig. 1): Approximate dorsal length: 46-53 mm. Color yellowish white. Moderate vestiture of yellowish setae. Head (Figs. 1, 2): Maximum width of head capsule 4.5-4.9 mm. Cranium yel- low, smooth to slightly rugose. Frons with 6-7 posterior frontal setae, 4 exterior frontal setae, 2—3 anterior angle setae, and 5 anterior frontal setae on each side. Dorsoepicranium with 4 setae on each side. Ocelli not defined. Clypeus subtra- pezoidal with 7-8 exterior clypeal setae on each side and 2 central setae. Preclypeus weakly sclerotized, without setae. Labrum nearly symmetrical with 12 posterior setae, one long lateral seta on each side, 2 long central setae and 2 long anterior VOLUME 108, NUMBER 3 without pternotorma. Laeotorma nar- row and elongated, with subquadrate Fig. 1. Demodema brevitarsis, third-instar larva. Scale line = 2 mm. setae. Epipharynx (Fig. 3): Corypha with 15-18 stout setae. Haptomerum wide, prominent, with 8-10 sensilla, behind process a _ transverse irregular row of 6 stout heli. Acanthoparia with 12 curved setae, progressively enlarged toward anterior border; penultimate seta on each side with apex narrowly cleft. Chaetoparia with 75-95 setae on each side. Proplegmatia absent. Plegmatia present, each plegmatium with 9—10 long plegmata. Dexiotorma wide and long, pternororma. Nesia with small sensorial cone and large sclerotized plate. Laeo- phoba abundant. Dexiophoba scarce. Crepis vague. Mandibles (Figs. 4-5) bi- colored, right mandible with 1 apical scissorial tooth and wide blade anterior to scissorial notch and | scissorial tooth posterior to notch. Stridulatory area absent. Molar area with 3 lobes. Calx wide. Brustia formed by 12-14 setae. 514 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-7. mandible, ventral view. 5, Left mandible, ventral view. 6, Left maxilla, internal view of distal part. 7, Left maxilla, ventral view of distal part. Scale lines = 1 mm, except figs. 3, 6-7 = 0.5 mm. Lateral edge with 2 basal setae. Left mandible (Fig. 5) with | apical scissorial tooth and wide blade anterior to scissor- ial notch and | scissorial tooth posterior to notch. Stridulatory area absent. Mo- lar area with 2 lobes. Acia absent. Demodema brevitarsis, third-instar larva. 2, Head, frontal view. 3, Epipharynx. 4, Right Brustia formed by 18—22 short setae. Lateral edge with 2 basal setae. Maxilla: Galea and lacinia fused (Fig. 8) forming mala. Mala with large uncus at apex and 3 subterminal large unci (Figs. 6—7). Stridulatory area (Fig. 9) with row of VOLUME 108, NUMBER 3 PCa JS a> NR ™ 14) r ay | 9 oS @ Figs. 8-16. Demodema brevitarsis, third-instar larva. 8, Hypopharynx and right maxilla, dorsal view. 9, Stridulatory area of maxilla. 10, Last antennal segment, dorsal view. 11, Last antennal segment, ventral view. 12, Tarsungulus of anterior leg. 13, Tarsungulus of middle leg. 14, Tarsungulus of posterior leg. 15, prothoracic spiracle. 16, Raster. Scale lines = 0.5 mm, except fig. 8 = 1 mm. 14-16 acute, small teeth, without distal, truncate process. Labium (Fig. 8): Dor- sal surface with short, wide truncate process. Hypopharyngeal sclerome with abundant phobae on both laterobasal sides. Glossa with 22—26 setae at middle, and | lateral row formed by 7—9 setae on each side. Antenna: Second segment 2 516 times longer than first segment, and nearly as long as third segment. Surface of last segment (Figs. 10-11) with 1 dorsal and 2—3 ventral sensory spots. Thorax (Fig. 1): Prothorax with small anterolateral areas weakly sclerotized. Thoracic spiracles (Fig. 15) with C- shaped respiratory plate 0.65 mm high and 0.45 mm wide; plate with 10—12 holes across diameter at middle; holes narrowly oval. Dorsoventral diameter of bulla 4 times longer than width of the narrowed respiratory plate. Distance between two lobes of respiratory plate widely open. Dorsal surface of each segment with scattered transverse rows of medium, size, erect Setae. Legs: Tar- sungulus of anterior and middle legs (Figs. 12-13) with long and acute apex. Tarsungulus of posterior legs (Fig. 14) with short and sharply pointed apex. All with | basal seta and | preapical seta. Abdomen (Fig. 1): Spiracles similar to those on thorax but variable in size. Spiracle of abdominal segment I: 0.55 mm high and 0.35 mm wide; seg- ment II: 0.60 mm high and 0.30 mm wide; segment III: 0.55 mm high and 0.28 mm wide; segment IV: 0.60 mm high and 0.35 mm wide; segment V: 0.45 mm high and 0.30 mm wide; seg- ments VI-VII: 0.40 mm_ high and O25 mm Mwider “and sseement— Will 0.30 mm high and 0.20 mm wide. Dorsal areas of segments I-VII with many scattered short, spinelike setae and irreg- ular, transverse rows of slender, long setae. Dorsal areas of segments VIII and IX only with irregular, transverse rows of slender, long setae. Dorsal area of segment X with mixture of long, short, and medium size setae. Segments IX and X not fused dorsally. Spiracular area and pleural lobes of abdominal segments I— IX with only a few setae. Raster (Fig. 16) with pair of palidia widely angled and divergent toward anal border, each con- sisting of row of 25-27 pali. Septula wide, open posteriorly, with finely rugose PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON surface. Tegilla composed of 25-28 hook-tipped, long setae. Lower anal lip with a shallow, rounded fovea before brief cleft on middle of border, and scattered mixture of short and long setae along anal border. Pupa.—This description is based on 18 pupae reared from third instar larvae collected on soybean roots. Locality data: Brazil: State of Rio Grande do Sul, Tapejara, 1-VII-1999, J. R. Salva- dori (1 male pupa, 8 female pupae) (IEXA); 3 male pupae, 6 female pupae (CNPT): Form: Body elongate, robust, exarate (Fig. 17). Body length: 19-21 mm. Red- dish yellow. With very fine velvety microtrichia, mainly on last abdominal segments. Head: Strongly reflexed down- ward. Antenna and mouth parts clearly separated. Ocular canthus and com- pound eyes well-differentiated. Clypeus convex. Labrum tumid. Surface of frons convex. Thorax: Pronotal disk slightly convex, with very shallow depression along midline. Meso- and metanota differentiated. Mesoscutellum widely an- gled, apex rounded. Metascutellum not defined. Pteroteca widened, with apex rounded, free, compressed around body; hind wing teca slightly longer than elytral teca. Meso-metasternal area with- out prominence. Protibia with 2 short process on external border. Meso- and metatibiae each with 2 rounded, short apical spurs. All tarsomeres vaguely defined. Abdomen: Tergites I-VII con- vex, without dioneiform organs. Tergites VIII-X fused, crearly narrowed toward end of body. Last tergite with paired, short, conical urogomphi, apically sharp- ly pointed (Figs. 17-19). Tergo-lateral tubercles absent. Spiracle I elongate, not prominent, covered by hind-wing teca. Spiracles II-IV tuberculiform, with open atrium and ringlike, sclerotized peritreme. Spiracles V—VI not promi- nent, opened, with ringlike narrow peri- treme. Spiracle VII tuberculiform, with VOLUME 108, NUMBER 3 SI Figs. 17-19. Demodema brevitarsis, pupa. 17, female, dorsal view. 18, apex of female abdomen, ventral view. 19, apex of male abdomen, ventral view. Scale lines = 1 mm, except fig. 17 = 2 mm. 518 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON open atrium and narrow _peritreme. Spiracle VIII closed, surrounded by fine rugae. Sternites II—-VII convex. Last sternite with genital ampulla widely rounded in female pupa (Fig. 18) and narrowly prominent in male pupa Gag. 19): Biological notes.—Larvae were found feeding on roots of Glycine max L. (Leguminosae) growing in common oxi- sol of the humid regime (hapludox). Depending on soil moisture, larvae and pupae were located at 18—25 cm deep. In the same locality, the larvae of this species have been found also on roots of corn (Zea mays L.), wheat (Triticum eastivum L.), ““aveia-preta” (Avena stri- gosa L.), and “‘azevem” (Lolium multi- florum Lam.) (Gramineae). Adults fly between December and February, and are attracted to electric lights. No host plants are known for the adults. The life cycle apparently is not regular. Some- times the adults fly each year or in other times fly every two years; this is under study by the second author (JRS). ACKNOWLEDGMENT This paper is a contribution to the project ““Coledpteros Lamelicornios de America Latina”’ supported by Departa- mento de Entomologia, Instituto de Ecologia, A.C. Xalapa, México (account 902-08-011). LITERATURE CITED Costa, C., S. A. Vanin, and S. A. Casari-Chen. 1988. Larvas de Coleoptera do Brasil. Museo de Zoologia, Universidade de Sao Paulo, Sao Paulo. 282 pp. 165 estampas. Evans, A. V. 2003. A checklist of the New World chafers (Coleoptera: Scarabaeidae: Melo- lonthinae). Zootaxa 211: 1-458. Moron, M. A. 1986. El género Phyllophaga en Mexico. Morfologia, distribucion y sistematica supraespecifica (Insecta Coleoptera). Instituto de Ecologia, A. C. México. 341 pp. Ritcher, P. O. 1966. White grubs and Their Allies. A Study of North American Scarabaeoid Larvae. Oregon State University Press, Cor- vallis. 219 pp. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 519-524 A NEW SPECIES OF HORAIELLA TONNOIR (DIPTERA: PSYCHODIDAE) FROM THAILAND GREGORY R. CuRLER, J. PHASUK, J. CHANPAISAENG, AND G. W. COURTNEY (GRC, GWC) Department of Entomology, 3222 Science II, lowa State University, Entomology, Faculty of Agriculture, Kasetsart University, Bangkok 10900, Thailand Abstract.—The moth-fly genus Horaiella Tonnoir is reported for the first time from Thailand. Numerous adult specimens were collected in Malaise-trap samples from Khao Yai National Park, central Thailand, including males and females of a new species, Horaiella iota Curler. Genitalic characters of the male confirm that the Khao Yai specimens represent a new species. A description and a brief discussion of bionomics and comparison with other known species of Horaiella are given. Key Words: Horaiella, Psychodidae, moth flies, Thailand Tonnoir (1933) described the genus Horaiella from two species collected in the Teesta Valley of northern India, H. prodigiosa and #H. consimilis. Subse- quently, Alexander (1953) described H. kuatunensis from Fukien, China. Except for these descriptions, and despite the unusual structure and habitat of the immature stages, the genus remains poorly known and rarely collected. Specimens recorded in this paper were collected during an inventory of stream- inhabiting Diptera of Khao Yai Nation- Al Park o(KCY NP) in+ central | Thailand: Established in 1962 and covering ap- proximately 2,170 km?, KYNP is the oldest and one of the largest parks in Thailand. The park includes parts of four provinces, Nakhon Ratchasima, Saraburi, Nakhon Nayok, and Prachin- buri, and is known for its biotic diversity, especially its vertebrates and vegetation. Much of KYNP consists of a large sandstone plateau dissected by numerous streams and covered by tropical and submontane broad-leaved evergreen for- ests (Gray et al. 1994). The rich biota reflects partly the park’s altitudinal di- versity (60—1,350 m). Samples from two streams in KYNP _ yielded numerous adult specimens of Horaiella iota, a new species described herein. Though imma- ture stages were not collected, adults of the present species exhibit sufficient differences in size, wing venation and terminalia to separate them from other described species of Horaiella. MATERIAL AND METHODS Specimens recorded in this paper were collected in Malaise traps set in July 2000 and checked every two weeks for one year (through June 2001). Traps were placed “either over the stream or im riparian vegetation within 5 meters of the stream. These streams were located in Nakhon Nayok Province, as follows: 1) Huai Patabak near km 29 (= dis- tance’ from south “entrance of 520 KYNB)s [4a19° No lOl?213E S50 5am: asl. This is a permanent stream, approximately 2—3.5m wide, with substrata comprising mostly cobble and coarse gravel. Huai Patabak is in a moderately dense forest, which keeps the stream shaded throughout the year. 2) Small creek 6.2 km up Khao Khieo Road, 14°22’'N, 101°24’E, 952 m asl. This is a temporary stream (4 months w/out surface flow) with a maximum width of approximately 2 m. Sub- strata are comprised mostly of cob- ble, boulders, and coarse gravels, and the riparian zone is well deve- loped, providing dense shade through- out the year. Specimens were collected and _ pre- served in 70% EtOH. Slide-mounted material was cleared in cedarwood oil and mounted in Canada balsam, follow- ing procedures’ described elsewhere (Courtney 1990). Some adult specimens were dried chemically using hexamethyl- disilazane, and mounted on pins. Speci- mens were examined using an Olympus SZX-12 dissecting microscope and a Ni- kon E-800 compound microscope, and drawings were rendered with the aid of a drawing tube on the Nikon system. Measurements are given in millimeters, as a mean followed by a range in parentheses where applicable. Values were recorded according to procedures outlined in Hogue (1973) and Courtney (2000) with the following exceptions: wing: length = point of greatest length, width = point of greatest width; head: length = posterior-most point of vertex to apex of mouthparts. Palpomere pro- Figs. 1—6: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON portions are from basal to apical seg- ments. Horaiella iota Curler, new species (Figs. 1-12) Diagnosis.—A small Horaiella. Male: Palpus 3-segmented; wing length less than 2 mm; radial fork arising basal to the tip of R>,3, medial fork arising apical to the tip of R>,3; branches of medial fork divergent in comparison to those of radial fork; hypopygium with gonostyli straight, 9th tergite with lateral lobes elongate, directed laterally. Female. Pal- pus and wing characters identical to male; subgenital plate with a pair of smoothly-rounded posterior lobes. Description.—Male: Measurements (N = 5): head length 0.25; wing length 1.48 (1.34-1.63); wing width 0.43 (0.39—0.49); leg segment lengths: forefemur 0.56 (0.51—0.62), tibia 0.45 (0.41-0.51), Ist tarsomere 0.20 (0.18—0.22), 2nd tar- somere 0.04, 34 tarsomere 0.025, 4th tarsomere 0.02, 5th tarsomere 0.05; mid- femur 0.66 (0.60—0.72), tibia 0.52 (0.47— 0.57), 1st tarsomere 0.24 (0.22—0.27), 2nd tarsomere 0.04, 3™4 tarsomere 0.025, 4th tarsomere 0.02, 5th tarsomere 0.05; hind femur 0.70 (0.64—0.77), tibia 0.53 (0.49— 0.60), 1st tarsomere 0.26 (0.25—0.28), 2nd tarsomere 0.04, 34 tarsomere 0.025, 4th tarsomere 0.02, 5th tarsomere 0.05 Head (Fig. 1): Eyes rounded, dichop- tic, widely separated; antenna 17-seg- mented; scape wide basally, constricted apically; pedicel globular; flagellomere one about 1.5,3, medial fork arising apical to the tip of R>,3; branches of medial VOLUME 108, NUMBER 3 fork divergent in comparison to those of radial fork; Sc ending in R,, R, with three branches, M with two branches, CuA with two branches, A, present. Foretibia with strigil (sensu Tonnoir) inserted posteroapically, with a row of four spines inserted distally on its medial surface. All legs with tarsomeres 2-4 remarkably short; tarsomere five with dorsomedial lip protruding apically. Tarsal claws of unequal length, medial tarsal claw elongate. Chaetotaxy: Scu- tum with four elongate, rigid setae inserted anteriorly and six inserted later- ally; scutellum with four elongate, rigid setae inserted posteriorly. Terminalia (Figs. 7-8, 11-12): Ab- dominal tergite IX with lateral lobes elongate, directed laterally. Cercus pres- ent as a pair of lobes extending poster- iorly beyond aedeagus, with several setae inserted apically. Gonocoxites stout ba- sally, with seven rigid, elongate spines inserted posteriorally, the medial-most spine longer and wider than _ those laterally. Gonostyli cylindrical, straight, subequal in length to gonocoxites, with eight rigid, elongate spines and numer- ous short spines inserted medially. Ae- deagus contained in tubular structure, bifurcate apically, with lateral parameral lobes basally; basiphallus with single Y- shaped sclerite; distiphallus with single sclerite consisting of a disclike portion basally with elongate filamentous projec- tions posterolaterally. Female: Measurements (N = 5): Head length 0.33; wing length 1.79 (1.69—1.90); wing width 0.57 (0.53—0.61); leg segment lengths: forefemur 0.60 (0.55—0.65), tibia 0.49 (0.45—0.53), 1st tarsomere 0.21 (0.20—0.22), 2nd tarsomere 0.04, 314 tar- “somere 0.025, 4th tarsomere 0.02, 5th tarsomere 0.05: midfemur 0.68 (0.63— 0.71), tibia 0.54 (0.51—0.57), 1st tar- somere 0.26 (0.24—0.27), 24 tarsomere 0.04, 3™4¢ tarsomere 0.025, 4th tarsomere 0.02, Sth tarsomere 0.05; hind femur 0.72 (0.71-0.75), tibia 0.56 (0.53-0.57), 1st 523 tarsomere ©.28 (0.27-0:29), 2nd tar- somere 0.04, 34 tarsomere 0.025, 4th tarsomere 0.02, 5th tarsomere 0.05 Head (Fig. 2): Identical to male except the mouthparts about 1.32 longer. Thorax and appendages: Wing identi- cal in shape, but slightly larger than in male. Legs identical in shape and pro- portion, but longer than in male. Chae- totaxy: Identical to that in male. Terminalia (Figs. 9-10): Cercus short, not longer than preceding abdominal segments, rounded apically; dorsal and ventral margins curled medially; lateral surface with numerous elongate setae, medial surface with dense setulae. Poste- rior margin of subgenital plate with a pair of rounded lobes, each lobe bearing a single posterolateral seta. Holotype.—Male. THAILAND. Na- khon Nayok Province: Khao Yai Nation- al Park, Huai Patabak, 14°19’N 101°21’E, 28.x—11.xi.2000, collected by Phasuk and Damrak, Malaise trap. Specimen mounted in Canada Balsam on slide, deposited in the National Museum of Natural History, Smithso- nian Institution, Washington, DC [USNM]. Allotype.—Female. same locality and date as holotype, mounted in Canada Balsam on slide, deposited USNM. Paratypes.— 4 6, 4 °. Same collection data as holotype and allotype, mounted in Canada Balsam on slides. Paratypes deposited in the National Insect Collec- tion, Department of Agriculture, Bang- kok, Thailand, and the Iowa State Insect Collection, Department of Entomology, Iowa State University, Ames, IA. Other material examined.—THAI- LAND. Nakhon Nayok Province. Khao Yai National Park, Huai Patabak, 14°19’N 101°21'E, collected by Phasuk and Damrak, Malaise trap, 2—16.1x.2000, 1 2; same location, 16—30.ix.2000, 2 2, 2 2: same location, 30.ix—16.x.2000, 13 6, 7 2: same location, 16—28.x.2000, 28 4, 31 2; same location, 1 1=25:x0:2000), 176, 524 1 2; same location, 25.xi—9.xii.2000, 5 2, 3 2; Khao Yai National Park, creek 6.2 km up Khao Khieo road, 14°22’N 101°24’E, collected by Phasuk and Damrak, Malaise trap, 16—28.x.2000, 1 2: same location, 11—25.x1-2000, 1 &. Other material deposited in the Depart- ment of Entomology, Kasetsart Univer- sity, Bangkok, Thailand, the Iowa State Insect Collection, Department of Ento- mology, Iowa State University, Ames, IA, and the Department of Entomology and Plant Pathology, University of Tennessee, Knoxville, TN. Etymology.—“‘Iota” is a Greek term that is used as both a number and a term to describe a very small amount of something. Horaiella iota is remarkably small in comparison to other members of the genus, hence the species name. Distribution.—This species is current- ly known only from two locations at Khao Yai National Park in central Thailand; however, it may occur in the surrounding areas as well. Bionomics.—Although Malaise traps were maintained throughout the year 2000, H. iota was captured only from September to December, with the great- est number of captures during mid- to late October. This suggests a univoltine life cycle and a peak activity period that coincides with the post-monsoon; how- ever, collection of the immature stages will provide a more exact assessment of phenology. Taxonomic notes.—Adults of H. iota are distinct among known species of Horaiella due to their small size and characters of the wing veins (e.g., radial fork arising basal to the medial fork). For the male, notable affinities are with H. consimilis and H. prodigiosa, the former being most similar to H. iota PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with respect to the structure of the 9th tergite, cerci and aedeagus. Horaiella iota females are similar to those of JH. prodigiosa, but differ in the shape of the subgenital plate (in H. iota, the posterior lobes are rounded and about as wide as long). ACKNOWLEDGMENTS We thank K. Damrak for field assis- tance and the National Research Council of Thailand, Royal Forestry Depart- ment, and Khao Yai National Park for granting permission to collect. Support for various aspects of this work were from the Thailand Research Fund (Roy- al Golden Jubilee Program), National Science Foundation (Grant DEB- 0103144), and National Geographic So- ciety (Grant 5347-94). LITERATURE CITED Alexander, C. P. 1953. Undescribed species of Nematocerous Diptera. Part II. Bulletin of the Brooklyn Entomological Society 48: 41— 42. Courtney, G. W. 1990. Cuticular morphology of larval mountain midges (Diptera: Deuteroph- lebiidae): Implications for the phylogenetic relationships of Nematocera. Canadian Jour- nal of Zoology 68: 556-578. . 2000. Revision of the net-winged midges of the genus Blepharicera Macquart (Diptera: Blephariceridae) of eastern North America. Memoirs of the Entomological Society of Washington, No. 23, 99 pp. Gray, D., C. Piprell, and M. Graham. 1994. National Parks of Thailand (Revised Edition). Industrial Finance Corporation of Thailand, Bangkok. 247 pp. Hogue, C. L. 1973. A taxonomic review of the genus Maruina (Diptera: Psychodidae). Bulle- tin of the Los Angeles County Natural History Museum 17: 1-69. Tonnoir, A. L. 1933. Description of remarkable Psychodidae and their early stages, with a theory of the evolution of the larvae. Records of the Indian Museum 35: 53-75. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 525-533 THE MINUTE PIRATE-BUG GENUS XYLOCORIS DUFOUR (HEMIPTERA: HETEROPTERA: ANTHOCORIDAE) FROM RICE MILLS IN THAILAND KAZUTAKA YAMADA, TOMOHIDE YASUNAGA, YUKINOBU NAKATANI, AND TosHIyvA HIROWATARI (KY, TH) Entomological Laboratory, Graduate School of Life and Environmen- tal Sciences, Osaka Prefecture University; Sakai, Osaka 599-8531, Japan (e-mail: kazu2000@plant.osakafu-u.ac.jp; hirowat_t@envi.osakafu-u.ac.jp); (TY) JICA-SV, c/o Natural History Museum, Tribhuvan Univ., Swayambhu, Kathmandu, Nepal (e- mail: kasumi@wlink.com.np); (YN) Laboratory of Insect Systematics, National Resources Inventory Center, National Institute for Agro-Environmental Sciences; Tsukuba, Ibaraki 305-8604, Japan (e-mail: nakatany@affre.go.jp) Abstract.—Three species of the anthocorid genus Xy/ocoris Dufour are reported from rice mills in southern Thailand. They are considered effective biological control agents against stored-product insect pests. Two of them, belonging to the subgenus Proxylocoris Carayon, are described as new: X. cerealis Yamada and Yasunaga, n. sp., and X. hyalinipennis Yamada and Yasunaga, n. sp. A key is provided to help distinguish these species. Key Words: Heteroptera, Anthocoridae, Xy/ocoris, new species, Thailand, biological control Xylocoris Dufour, 1831, is a large anthocorine flower bug genus, compris- ing about 50 species in the world, principally in the MHolarctic Region (jo Chu 1969, “Henry 1988; “Peéricart 1996). These bugs usually prey on various small arthropods, and some have been reported to be effective natural enemies against pests in stored-food facilities and grain mills (e.g., Péricart 1972, Lattin 2000). It is necessary to accurately identify species to establish modern taxonomic research for such natural enemies. There is no adequate taxonomic study on the genus Xy/ocoris in the Oriental Region. During continuing investigations of rice mills in Thailand, Nakatani collected three species of XYyv/ocoris. One of them is X. (Arrostelus) flavipes (Reuter, 1875), a well-known predator of stored-product insect “pests: The’ other two’ ‘represent undescribed species belonging to the subgenus Proxylocoris Carayon, 1972. Two species, Xylocoris (Proxylocoris) cerealis and X. (P) hyalinipennis, are described here as new, and the well- known _X. (A.) flavipes is diagnosed. In addition to diagnoses and descriptions of these three species, biological informa- tion on Xy/ocoris in rice mills is docu- mented, and key is provided to distin- guish the three Xy/ocoris species from rice mills in Thailand. MATERIALS AND METHODS Dried specimens of each species were used. For genitalic observations, the 526 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON entire abdomen was removed from the body, soaked in 5% hot KOH solution for 3—5 minutes, and placed into glycerin on a glass slide. Illustrations of the structure were drawn with the aid of a Nikon Stereoscopic Zoom Microscope SMZ1500. Scanning electron microscopy was used to observe the details of surface structures. Depositories of material are abbrevi- ated as follows: National Institute of Agro-Environmental Sciences, Tsukuba (NIAES); Entomological Laboratory, Osaka Prefecture University, Sakai (OPU). Abbreviation is used for the collector of the specimens examined: YN (Yukinobu Nakatani). All measure- ments are given in millimeters. Only selected references are cited for each taxon; see the catalogue of Péricart (1996) for further information. Termi- nology for descriptions follows Carayon C1972). TAXONOMY Genus Xylocoris Dufour Xylocoris Dufour 1831: 423. Type species by monotypy: Xylocoris rufipennis Dufour 1831. Diagnosis.—Recognized by the oval to elongate-oval body, dorsum shining and smooth, pronotum with stout, long, suberect corner setae, well-developed fossula spongiosa at apex of foretibia, midtibia bears smaller foussula spon- giosa, stout, long spines distributed along predominant part of the metatibia, sickle-shaped paramere curved toward the posterolateral end and ectosperma- lege usually opening at right side of abdominal terga III and VIII in female. Remarks.—This genus 1s divided into four subgenera, Xy/locoris s. str., Prox- yvlocoris Carayon, Arrostelus Kirkaldy, and Stictosynechia Reuter, based on the position of the tibial teeth in the male and the surface structure of the ostiolar peritreme, besides the peculiar copulato- ry direction. All species of Xylocoris are predaceous and often found in habitats such as under the bark of trees, among dead plant materials, and in litter layers of shaded forests. A few species are found in stored-food facilities, especially grains, and preferably prey on small arthropods. Some species have brachypterous, sub- brachypterous, and macropterous forms. Subgenus Proxylocoris Carayon Proxylocoris Carayon 1972: 594 (as a subgenus of Xylocoris). Type species by original designation: Piezostethus afer Reuter 1884. Remarks.—Proxylocoris is distingui- shed from other subgenera by a combi- nation of the following characters: male fore- and midlegs with tibial teeth always present, row of tibial teeth on midleg longer than that of foreleg; canaliculi of ostiole distinct and densely distributed (Figs. 8-9); ectospermalege placed be- tween abdominal terga II and III (Figs. 17,21): This subgenus has contained approx- imately 20 species; most of these are distributed in the Palearictic Region (Péricart 1996). A single member, X. clarus (Distant, 1910), is known from the Oriental Region (India and Myanmar). Xylocoris (Proxylocoris) cerealis Yamada and Yasunaga, new species (Figs. 1, 4, 8, 11—14) Xylocoris (Proxylocoris) sp. 1: Nakatani 2004: 38. Diagnosis.._Recognized by the shiny, oblong-oval body, semi-transparent he- melytron, infuscate clavus and corium that are darkened along the inner margin with a dark subtriangular spot at the mesial inner margin (Fig. 4), medially angulate paramere in posterior view VOLUME 108, NUMBER 3 Figs. 1-3. X. (Arrostelus) flavipes, °, (Fig. 13), and much smaller, weakly sclerotized ectospermalege (Fig. 14). Description.—Body (Fig. 1) oblong- oval; dorsal surface shiny, with pale yellow, short, reclining pubescence. Head (Fig. 1) blackish brown, smooth, head about 0.8 times as long as wide across eyes; apex of head tinged with pale yellow; three pairs of long, erect setae on each side of tylus, near anterior inner margin of eyes, and between eyes and ocelli; length of anteocular portion as long as eye in dorsal view; vertex 3.6 times as wide as eye in dorsal view; ocelli Figs. 4-7. nN i) — Xylocoris spp. from Thailand. 1, X. (Proxylocoris) cerealis, ¢. 2, X.(P.) hyalinipennis, ° . 3, subbrachypterous. Scale bars: 1.0 mm. reddish brown. Antenna yellowish brown; segment I dark brown, almost reaching to apex of head, with sparse, short setae; segment II tinged with dark brown on apical %3, thickened toward apex, about 0.78 times as long as width of head, with densely distributed, re- clining setae; segments III and IV each shorter than segment II, with sparsely distributed, long, suberect setae; segment III slightly shorter than IV; length of segments I-IV (4/2): 0.16/0.16, 0.38/ 0.40—-0.41, 0.31/0.31—0.34, and 0.34/0.34— 0.38. Rostrum yellowish brown, attain- Left forewings of Xylocoris spp. 4, X. (Proxylocoris) cerealis. 5, X. (P.) hyalinipennis. 6, 7, X. (Arrostelus) flavipes, macropterous and subbrachypterous. Scale bars: 0.5 mm. sy PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 8—10. ing base of mesocoxae; segment I and basal half of segment II tinged with dark brown; segment III more than twice as long as II. Pronotum (Fig. 1) varying from black to blackish brown, sparsely pubescent, with posterior half weakly depressed and transversely strigosed; collar incomplete; anterior margin of pronotum curved, slightly shorter than mesal length; lateral margin nearly straight; posterior margin slightly concave, twice as long as anterior margin. Scutellum black to blackish brown, apex pale yellow, anterior “% of lateral side bearing long, suberect setae. Hemelytron (Figs. 1, 4) semitransparent, tinged with yellowish brown, with sparsely distributed, short, reclining pu- bescence and minute punctures; clavus and corium darkened along inner mar- gin, with dark subtriangle spot at mesial inner margin; membrane transparent, with indistinct vein. Ostiolar peritreme (Fig. 8) brown; ostiole wide, curved anteriorly; apex of ostiole not reaching anterior margin of metapleuron; supra- coxal area with transverse rows of furrows. Legs yellowish brown, stout, with densely distributed, pale yellow pubescence; each femur dark brown, with apex slightly yellowish; each tibia Ostiolar peritremes of Xy/ocoris spp. in left lateroventral view. 8, XY. (Proxyvlocoris) cerealis. 9, X. (P.) hyalinipennis. 10, X. (Arrostelus) flavipes. Scale bars: 0.1 mm. Abbreviations: sca = supracoxal area; cn = canaliculi. bearing long, stout spines. Abdomen blackish brown; scissure on abdominal terga reaching segment III. Male genitalia (Figs. 11-13): Pygo- phore longitudinally narrow and basally constricted, posteroventrally with long setae; longest setae slightly shorter than half of paramere; paramere medially angulate, slightly bent at apex. Female genitalia (Fig. 14): Ectosper- malege much smaller than that of X. galactinus and X. hiurai and weakly sclerotized, extending posteriorly. Measurements: &/2: Body length 2.75/2.88—3.08; head length (excl. neck) 0.41/0.39-0.41; head width (incl. eyes) 0.48/0.51; vertex width 0.30/0.32—0.34; width between ocelli 0.23/0.26; rostral length 0.91/0.97—-1.03; anterior pronotal width 0.43/0.39—0.43; mesal pronotal length 0.47/0.47—0.48; basal pronotal width 0.98/1.06—1.09; length of embolial margin 0.90/0.94—1.00; length of lateral cuneal margin 0.50/0.48—0.50; maximum width across hemelytra 1.05/1.15—1.20. Holotype.— ¢, THAILAND: Songkhla Prov., Ranode, Rice mill factory, NO7- 48’ E100-16’, 11. xi. 2002, Y. Nakatani (NIAES). Paratypes—THAILAND: 1 ?, Na- khon Si Thammarat, NO8-15’ E100-01’, VOLUME 108, NUMBER 3 Figs. 11-21. (Proxylocoris) cerealis. 15-18, X. (P.) hyalinipennis. 19-21, X. (Arrostelus) flavipes. 11, 15, 19, Pygophores in dorsal view; 12, 16, 20, parameres in dorsal view; 13, 17, 21, parameres in posterior view; 14, 18, ecto- spermalege in dorsal view. Scale bars: 0.1 mm. 16. xi. 2002, YN (OPU); 1 2, Nakhon Si Thammarat, NO8-23’ E100-02’, 15.x1.2002, YN (OPU); 1 2, same data as holotype (NIAES). Distribution.—Southern Thailand. Etymology.—From the Latin “‘cerea- lis” (= of grain), referring to its habitat in cereal stocks in Thailand; an adjective. Remarks.—In_ general appearance, this new species resembles the cosmopol- itan X. galactinus (Fieber, 1836) and the temperate eastern Asian inhabitant ¥X. hiurai Kerzhner and Elov, 1976. Xy/lo- coris cerealis is distinguished from X. galactinus by the apex of ostiolar canal not extending to the anterior margin of the metapleuron (reaches the anterior 529 Male (11—13, 15-17, 19-21) and female (14, 18) genitalia of Xy/locoris spp. 11—14, X. margin of metapleuron in X. galactinus), medially angulate paramere (strongly curved in X. galactinus), and much smaller ectospermalege (rather larger ectospermalege in X. galactinus), and from X. hiurai by the weakly sclerotized and posteriorly extended ectospermalege (more darkened and obscure, subtrian- gular, and anteriorly broadened in X. hiurai). Xylocoris (Proxylocoris) hyalinipennis Yamada and Yasunaga, new species (Figsy 2; >, 91513) Xylocoris (Proxylocoris) sp. 2: Nakatani 2004: 38. 530 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Diagnosis.—Recognized by the much smaller body, mostly hyaline hemelytron that is tinged with ocherous, infuscate clavus and corium that are narrowly darkened along inner margins (Fig. 5), ostiolar peritreme accompanying narrow evaporative area and widened ostiole (Fig. 9), the strongly curved apical ' of the paramere that is furnished with a bifurcate process subapically (Fig. 17), and the anteriorly broadened ectosper- malege (Fig. 18). Description.—Body (Fig. 2) elongate oval; dorsal surface shiny, with bright yellow, short, reclining pubescence. Head (Fig. 2) blackish brown, smooth, about 0.75 times as long as width across eyes; apex of head broadly tinged with pale yellow; a pair of erect setae present at each side of tylus near anterior inner margin of eye, and between eye and ocellus; anteocular portion slightly lon- ger than eye in dorsal view; vertex 3.8 times as wide as eye in dorsal view; ocelli red. Antenna yellowish brown; segment I nearly extending to apex of head, with sparsely distributed, short setae; segment II sometimes tinged with dark brown at apex, gradually thickened to- ward apex, 3/4 as long as head width, with densely distributed, reclining setae; segments III and IV, each almost equal in length to II, with sparsely distributed, long, suberect setae; length of antennal segments I-IV (6/2): 0.13/0.13, 0.30/ 0.31, 0.30/0.30—0.31, 0.30/0.29—0.30. Ros- trum yellowish brown, extending to base of mesocoxae; segment III about twice as long as II. Pronotum (Fig. 2) black to blackish brown, sparsely pubescent, with weakly depressed posterior half; collar indis- tinct; anterior margin of pronotum curved, about as long as mesal length; anterior humeri rounded; lateral margin nearly straight; posterior margin slightly curved, slightly more than twice as long as anterior margin. Scutellum black to blackish brown; anterior “% of lateral side bearing long, suberect setae. Hem- elytron (Figs. 2, 5) nearly transparent, tinged with yellowish brown, with sparsely distributed, short, reclining pu- bescence and minute punctures; clavus and corium narrowly darkened along inner margin; membrane _ transparent, with indistinct veins. Ostiolar peritreme (Fig. 9) brown, with narrow evaporative area and rather broadened ostiole curved anteriorly; apex of ostiole almost reach- ing anterior margin of metapleuron; supracoxal area with transverse furrows. Legs yellowish brown, stout, densely clothed with pale yellow pubescence; each femur dark brown, tinged with yellowish brown at apex; each tibia bearing long stout spines. Abdomen blackish brown; scissure on abdominal terga extending to segment III. Male genitalia (Figs. 15-17): Pygo- phore basally narrow; paramere arising near posteroventral angle of pygophore, extending to near median projection on left margin of pygophore in dorsal view, strongly curved at apical 3 in posterior view, with subapical conspicuous, bi- furcate process. Female genitalia (Fig. 18): Ectosper- malege anteriorly broadened, with a com- plexly folded thin duct at right lateral margin. Measurements: &/°: Body length 2.25/ 2.30—2.38; head length (excl. neck) 0.30/ 0.31—0.32; head width (incl. eyes) 0.40/ 0.43; vertex width 0.28/0.27—0.28; width between ocelli 0.20/0.22—0.23; rostral length ?/0.97-1.03 (¢: apical two seg- ments lost); anterior pronotal width 0.34— 0.36/0.36—-0.38; mesal pronotal length 0.38—0.43/0.38—0.40; basal pronotal width 0.77—0.78/0.83—0.85; length of embolial margin 0.70/0.70—0.74; length of cuneal margin 0.35/0.38—0.40; maximum width across hemelytra 0.85/0.86—0.88. Holotype.—%, THAILAND: Nakhon Si Thammarat, Rice mill factory, N08- 23’ E100-02’, 15.xi.2002, Y. Nakatani (NIAES). . VOLUME 108, NUMBER 3 Paratypes.—THAILAND: 1 @&1 &, same data as for holotype (NIAES): 1 2, 1 2°, Nakhon Si Thammarat, NO8-15’ ETOO-017,01G6ix1 2002» YN (OPU);.2. 2, Songkhla Prov., Ranode, NO7-48’ E100- 1G. isa 2002.YIN: (OP). Distribution.—Southern Thailand. Etymology.—From the Latin “‘hyali- nus” (= transparent), combined with ““pennis’ (= wing), referring to the nearly transparent hemelytra; an adjective. Remarks.—This new species is very similar to X. clarus (Distant, 1910) in the coloration of the hemelytra and appear- ance of female genitalia from which it is distinguished by the apex of the para- mere extending to near median projec- tion on left margin of pygophore (apex of paramere extending well beyond median projection on left margin of pygophore in X. clarus) and the com- plexly folded thin duct at right lateral margin of ectospemalege (simple, vague, and conical ectospermalege in X. clarus). This new species is allied to former species, but may be distinguished by smaller body size, narrowly darkened along inner margin of clavus and corium, narrow evaporative area, broadened ostiole, and subapically curved paramere with a characteristic bifurcate process. Subgenus Arrostelus Kirkaldy Arrostus Reuter 1884: 35 (as subgenus of Piezostethus) (janior homonym of Ar- rostus Eahreus. 1872, Coleoptera): Type species by monotypy: Piezo- stethus flavipes Reuter 1884. Arrostelus Kirkaldy 1906: 119. New name for Arrostus Reuter. Remarks.—This subgenus may _ be separated from other subgenera by a combination of the following charac- ters: male tibial teeth always absent; supracoxal area of metapleuron distinct (Fig. 10); ectospermalege always absent; and copulation site and cicatrices present on anterodorsal area of abdomen. SS Four species, X. congoensis (Bergroth, 1905); X¢ ) flavipes “@Reuters 1875), Xi hirsutus Carayon, 1961, and X. queen- slandicus Gross, 1956, are known to occur in the tropical and subtropical zone around the world (Chu 1969, Péricart 1996). Xylocoris (Arrostelus) flavipes (Reuter) (Figs /33:6).7;, TO: 19=21) Piezostethus flavipes Reuter 1875: 65. Diagnosis.—Recognized by the small (1.8—2.2 mm), elongate-oval body (Fig. 3): dark brown head (Fig. 3) with pale yellow tylus; pale yellow antenna and rostrum; dark brown, unicolorous, high- ly polished, almost impunctate, and sparsely pubescent pronotum (Fig. 3); yellowish-brown hemelytron (Figs. 3, 6— 7), broadly darkened cuneus in macrop- terous form, corium and cuneus that are sometimes narrowly darkened in sub- brachypterous form; ostiolar peritreme (Fig. 10) with a smooth ostiole and supracoxal area; entirely pale yellow legs; pygophore (Fig. 19) with charac- teristic process basally; and moderately curved paramere (Figs. 20-21). Measurements: ¢/2: Body length 1.80—1.88/2.00—2.25; head length (excl. neck) 0.29—0.30/0.30—0.34; head width including compound eyes 0.35—0.36/ 0.38—0.41; vertex width 0.24—0.26/0.24— 0.26: width between ocelli 0.19—0.21/ 0.19—0.21; rostral length 0.81—0.84/0.81— 0.88: pronotal anterior margin width 0.31—0.32/0.34; mesal pronotal length 0.32—0.34/0.34—0.36; basal pronotal width 0.61—0.64/0.70—0.82; length of embolial margin 0.50—0.53/0.55—0.74 in subbra- chypterous, ?/0.65—0.75 in macropterous; length of cuneal margin 0.20—0.22/ 0.24—0.27 in subbrachypterous, ?/0.40— 0.45 in macropterous; maximum width across hemelytra 0.70—0.72/0.78—0.80 in subbrachypterous, ?/0.80—0.90 in mac- ropterous. 532 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Specimens examined: (Subbrachypterous form: subbrac.; macropterous form: mac.) TEALUAIND. 3165 il SaGubbracs)siNa= khon Nayok, N14-16° E101-04’, 2.x1.2002, YN (OPU); 1 & (subbrac.), Nakhon Si Thammarat, N0O8-24 100-00’, 15.x1i. 2002, YN (OPU); 2 (C*subbracei2 mac.), Nakhon Si Thammarat, NO8-23’ E100-02’, 15.xi.2002, YN (NIAES); 2 2 subbrac., 1 mac.), Phatthalung, N07-36° E100-07’, 12.xi.2002, YN (NIAES). Distribution.—Europe, northern and tropical Africa, Asia (Turkey, Saudi Arabia, Yemen, Pakistan, India, Indone- sia, Thailand, China), Australia, North America (U.S.A), South America. KEY TO XYLOCORIS SPECIES FOUND IN RICE MILLS IN THAILAND 1. Hemelytron yellowish brown, with dark- ened cuneus; femora pale yellow; ostiole smooth, lacking canalicu- Lit eee 3 AIT EES AMES Hemelytron mostly transparent or semi- transparent; femora dark brown; ostiole withecanaliculite es 5 tees auc ie yao asain ae D unicolorous X. (Arrostelus) flavipes tN Body larger (about 3.0 mm long); clavus and corium darkened along inner margins with a dark, subtrianguler spot at mesial inner margin; paramere angulate medial- | eters, 2. eee Body smaller (about 2.3 mm long); clavus X. (Proxylocoris) cerealis and corium narrowly darkened along inner margin; paramere curved subapically, with characteristic bifurcate process........ Spee COREY cs erica aka Gees Y. (P.) hyalinipennis BIOLOGY Visarathanonth and Sukprakarn (1988) reported 72 species of stored-product insect pests from Thailand. Present in- vestigations of rice mills, the following insect species were found from rice bran and polished rice: Rhyzopertha dominica (Fabricius) (Coleoptera, Bostrichidae), Lophocateres pusillus (Klug) (Col., Tro- gossitidae), Oryzaephilus | surinamensis (Linnaeus) (Col., Silvanidae), Cryptolestes pusillus (Schonherr) (Col., Cucuyidae), Tribolium castaneum (Herbst) (Col., Te- nebrionidae), Sitophilus spp. (Col., Cur- culionidae), Corcyra cephalonia (Stain- ton) (Lepidoptera, Pyralidae), and Sitotroga cerealella (Oliver) (Lep., Gele- chidae). They have been documented as major pests of paddy and milled rice. Xylocoris flavipes is known to prey on many developmental stages of various coleopteran and _ lepidopteran pests (Jay et al. 1968, Arbogast et al. 1971). In addition, X. sordidus is known to inhabit stored grain facilities in North and South America. The cosmopolitan species X. galactinus is a known predator of various insects found in the stored- grain habitats (Chu 1969, Péricart 1972). Thus some. stored-food insect pests can be regarded as prey of the anthocorid species reported here. Xy/o- coris flavipes is the only species known previously from Thailand (Visaratha- nonth et al. 1994). This species is widely distributed in temperate, subtropical, and tropical zones of the world. The native range of this species is difficult to determine because it has been trans- ported to so many parts of the world in commerce (Lattin 2000). Xylocoris flavipes is the most common in rice mill inhabitant in Thailand. In contrast, X. cerealis and X. hyalinipennis appear to be restricted in distribution and currently are found only in the southern Thailand, where they co-occur with X. flavipes in Phatthalung. ACKNOWLEDGMENTS Weare grateful to Dr. T. J. Henry (US. Department of Agriculture, Washington DG.) and Dry J.D: Latin (Department of Botany and Plant Pathology, Oregon State University) for reviewing the man- uscript with useful comments and sugges- tions. Yamada expresses his cordial thanks to Prof. M. Ishii and Mr. N. Hirai (Osaka Prefecture University) for their encouragement and advice on his study. Yasunaga also is appreciative of Dr. S. Miyamoto (Fukuoka City) for his con- tinuous interest and advice. This survey VOLUME 108, NUMBER 3 was partly supported by a research pro- ject, “Development of low-input technol- ogy for reducing post harvest losses of staples in Southeast Asia,” of the Japan International Research Center for Agri- cultural Sciences. LITERATURE CITED Arbogast, R. T., M. Carthon, and J. R. Roberts, Jr. 1971. Developmental stages of Xy/ocoris flavipes (Hemiptera: Anthocoridae), a predator of - stored-product insects. Annals of the Entomological Society of America 64: 1131— 1134. Carayon, J. 1972. Le genre Xylocoris: subdivision et espéces nouvelles (Hem. Anthocoridae). Annales de la Société Entomologique de France (n.s.) 8: 579-606. Chu, Y. I. 1969. On the bionomics of Lyctocoris beneficus (Hiura) and Xylocoris galactinus (Fieber) (Anthocoridae, Heteroptera). Journal of the Faculty of Agriculture of the Kyushu University 15: 1-136. Dufour, L. 1831. Description et figure du Xy/ocoris rufipennis, Hémiptére nouveau. Annales des Sciences Naturelles 22: 423-426. Henry, T. J. 1988. Family Anthocoridae, pp. 12-28. 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 Netherland. 958 pp. Jay. E., R. Davis, and S. Brown. 1968. Studies on the predaceous habits of Xylocoris flavipes (Reuter). Journal of the Georgia Entomolog- ical Society 3: 126—130. Kirkaldy, G. W. 1906. List of the genera of the pagiopodous Hemiptera-Heteroptera with their type species, from 1758 to 1904 (and also 533 of the aquatic and semi-aquatic Trochalo- poda). Transactions of the American Entomo- logical Society 32: 117-136. Lattin, J. D. 2000. Chapter 26. Minute pirate bugs (Anthocoridae), pp. 607-637. In Schaefer, C. W. and A. R. Panzini, eds. Heteroptera of economic importance. CRC Press, New York. 828 pp. Nakatani, Y. 2004. Heteroptera, pp. 31-39. In Hayashi, T., S. Nakamura, and P. Visaratha- nonth et al., eds. Stored rice insect pests and their natural enemies in Thailand. JIRCAS International Agricultural Series No. 13. Fun- ny Publishing, Bangkok. 79 pp. Péricart, J. 1972. Hémipteres Anthocoridae, Cimi- cidae et Microphyisidae de lOuest-paléarc- tique. Jn Faune de l’Europe et du_ bassin Méditerranéen 7: pp. 1-402. Masson, Paris. . 1996. Family Anthocoridae Fieber, 1836 - Flower bugs, minute pirate bugs, pp. 108—140. In Aukema, B. and C. Rieger, eds. Catalogue of the Heteroptera of the Palaearctic Region Vol. 2. Cimicomorpha I. The Netherlands En- tomological Society, Amsterdam. 359 pp. Reuter, O. M. 1875. Genera Cimicidarum Euro- pae. Bihang till Kongliga Svenska Vetenskap- sakademiens Handlingar 3: 1—66. . 1884. Monographia Anthocoridarum Or- bis Terrestris. Helsingforsiae, 204 pp. (also Acta Societatis Scientiarum Fennicae 16 (1885): 555-758). Visarathanonth, P., H. Nakakita, and P. Sitti- suang. 1994. Role of natural enemies in the regulation of stored-product insect popula- tions in rice storages in Thailand. JIRCAS Journal 1: 1-7. Visarathanonth, P. and S. Sukprakarn. 1988. Current problems of pests of stored products in Thailand. Proceedings of the symposium on pests of stored producs; Bogor, Indonesia. Biotrop Special Publication 33: 45—54. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 534-542 TWO NEW SPECIES OF THE NEOTROPICAL LEAFHOPPER GENUS CALDWELLIOLA YOUNG (HEMIPTERA: CICADELLIDAE) WITH A KEY TO MALES STUART H. MCKAMEY Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0168, U.S.A. (e-mail: smckamey@sel.barc.usda.gov) Abstract.—Two new species of Caldwelliola are described, C. sinuata from Honduras and C. trilineata from Ecuador, bringing the current total valid species in the genus to eleven. A key to males of all species is presented. The species C. bipunctata Nielson and Godoy, 1995, is placed in synonymy under C. /utea Signoret, 1855, n. syn. One species, C. caucana Young, has been implicated as a likely vector of the bacterium Xyle/la fastidiosa Wells, which causes the “‘crespera”’ disease, coffee leaf scorch, in Colombia and elsewhere in the Neotropical Region. Key Words: Cicadellinae, coffee leaf scorch, Xy/ella, Honduras, Ecuador, Hortensia, Chlorogonalia There is some truth in the generaliza- tion that “most leafhoppers are small and green,” as many are cryptically colored and blend into the green foliage upon which they feed, and most are less than a centimeter long. The subfamily Cicadellinae (sensu. Young 1968), or sharpshooters, is especially large and diverse. The New World, mostly tropical fauna alone comprises over 1,500 species in over 230 genera. Moreover, the tribe Cicadellini, with over 1,100 New World species, contains some of the largest leafhoppers and also some of the most common small green leafhoppers. Within this tribe, Hortensia Metcalf and Bruner, Chlorogonalia Young, and Caldwelliola Young are commonly en- countered, are externally similar to each other, and all have small, green species distributed from Mexico through Boli- via. Although Hortensia is monotypic, individuals vary surprisingly in size across its range, yet are easily identified by the coloration of the head (Young 1977: fig. 654). Chorogonalia and Cald- welliola, however, are less easily distin- guished, especially from each other because of superficial similarities in size and color, especially that of the head (Figs. 1-2). Diagnostic features to dis- tinguish undissected males and females of these two genera are provided in a review of Chlorogonalia (McKamey 2006). When originally described (Young 1977), Caldwelliola contained eight spe- cies, six of them new. Nielson and Godoy (1995) described the species C. bipunctata and McKamey (2006) added one additional species (Chlorogonalia tharma Young). In the present paper, two new species of Caldwelliola are described and one is placed in synonymy. A key to males of all species 1s presented so that C. caucana Young, a likely vector VOLUME 108, NUMBER 3 Figs. 1-2. Oblique aspect of anterior portion. 1, Chlorogonalia ultima. 2, Caldwelliola reservata. of coffee leaf scorch disease (crespera in Spanish) in the Neotropics (McKamey 2006), can be identified. Caldwelliola Young Caldwelliola| Young 1977:1039. Type species by original designation: Te/ti- gonia reservata Fowler. Diagnosis.—Male with subgenital plate broad at base, strongly narrowed from base to apex (Fig. 3), styles truncate or narrowly rounded, aedeagal paraphyses present, bifurcate, sometimes flattened, adominal segment XI length 0.3—0.7x length of entire pygofer (Fig. 6); sternum II apodemes usually exceeding posterior margin of segment III (Fig. 5); female with sternum VII produced posteroventrally, 585 acute (Fig. 4), its length almost equal to exposed pygofer length (covering most of its base). The form of the male subgenital plate and female sternum VII are visible in undissected specimens and sufficient to distinguish Caldwelliola from Chlorogo- nalia, which have the subgenital plates strongly narrowed abruptly near their apices and female sternum VII only about half the length of the exposed portion of the pygofer. Description (modified from Young 1977).—Length: Including forewings in repose, 5.2—9.0 mm. Head: Crown moderately produced; median length of crown varying from slightly less than 3/5 to more than 3/4 interocular width; anterior margin broadly rounded in dorsal view; margin not angular at transition from crown to face; ocelli located on or slightly behind a line between anterior eye angles; crown surface with a transverse concavity before ocelli and with a shal- low median concavity; lateral clypeal sutures extending onto crown; clypeus slightly flattened in profile; muscle 1m- pressions indistinct; clypellus with con- tours of its lower portion slightly more horizontal than profile of remainder of face; transclypeal suture obsolete medi- ally. Thorax: Pronotum narrower than transocular width of head; dorsopleural carinae complete; disk smooth, without pubescence; without transverse rugae on its posterior portion. Forewing coria- ceous with membrane including all of apical cells except basal portion of out- ermost and extending basad a_ short distance along costal margin (except indistinct in C. /utea (Signoret)); veins distinct, 3 closed anteapical cells, bases of inner and outer cells almost in trans- verse line with claval apex, base of cell more proximal; four apical cells, base of fourth usually located more proximally than base of third; without an anteapical plexus of veins; forewings of female in 536 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 3-4. Distinguishing external features of Caldwelliola species in undissected males and females, respectively. SGP, subgenital plate; StV/J, abdominal sternum VII. 3, C. reservata male. 4, C. reservata female. Figs. 5-7. Distinguishing internal features of Caldwelliola. 5, C. reservata, cleared male abdomen, showing the large sternum II apodemes (APO) often, but not always present in Caldwelliola, and the blunt genital style (ST Y). 6-7, Dorsal and lateral aspects of large abdominal segment XI in Ca/dwelliola. 6, Male of C. caucana. 7, Female of C. reservata. VOLUME 108, NUMBER 3 rest position exceeding apex of oviposi- tor. Hindleg with femoral setal formula 2:1:1; first tarsomere length slightly greater than combined length of last 2 tarsomeres and with 2 parallel rows of small setae on plantar surface. Male terminalia: Pygofer moderately produced posteriorly; posterior margin varying from narrowly to broadly con- vex; without processes. Subgenital plates separate throughout their length; extend- ing almost as far posteriorly as pygofer apex; each broad at base, abruptly narrowing at about mid length; triangu- lar and with macrosetae uniseriate. Style without preapical lobe; apex truncate, narrowly rounded, or acute. Connective broadly Y-shaped or triangular. Aedea- gus symmetrical, with elongate basal apodemes, without processes. Paraphy- ses present, bifurcate, appearing con- torted or linear, narrow throughout. Female terminalia: Abdominal ster- num VII with posterior margin produced and angular or subangulate. Genital chamber without sclerites. Ovipositor with second valvulae each broadened beyond basal curvature and_ bearing quadrate teeth. Pygofer with macrosetae on posterior half. Coloration: Most species dull green except head, anterior portion of prono- tum, mesonotum, and scutellum some- times tan to yellow (except C. lutea, which is entirely yellow); crown and frontoclypeus usually with brown or black markings visible in both dorsal and anterior views. Distribution.—Mexico, Central Amer- ica, Venezuela, and Colombia to Bolivia, west-central Brazil. Comments.—Most species of the ge- nus Caldwelliola have little intraspecific color variation. The three exceptions, revealed by their uniform genitalia (es- pecially the aedeagus) are C. andamira (Figs. 16, 25), which has two distinct patterns of crown coloration, and C. lutea. (Figs: 195. 26) andi «C. reservata 537 (Figs. 20-21), which each have two distinct patterns of pronotal coloration. The original description of C. /utea, based on a Guatemalan specimen, states that the prothorax has two dark spots along the median line (as in Fig. 19). Fowler (1900), who had specimens from Guatemala and Mexico, mentioned that the posterior spot is sometimes absent, presumably referring to the illustrated specimen, from Atoyac, Veracruz, which had only one spot (as in Fig. 26). Young (1977) saw specimens from Mexico, Guatemala, El Salvador, and Costa Rica, and illustrated a specimen from Veracruz that had only the anterior spot. Nielson and Godoy (1995) de- scribed C. bipunctata from Costa Rican specimens, distinguishing their new spe- cies from C. lutea “by the presence of 2 black spots on the pronotum, the paraphysis branched before the middle and by the shorter sternal abdominal apodemes.” While these features differ from the illustration provided by Young (1977)3 “the “spots ‘on? thes pronotum match Signoret’s description. Among specimens in the National Museum of Natural History (USNM), there are several series with the single, anterior pronotal spot; all are from Mexico, from Veracruz (Cordoba and Jalapa) and San Luis Potosi (Tamazunchale). The USNM specimens with two pronotal spots are from Guatemala, El Salvador, and Nicaragua. The Jalapa specimen illustrated by Young (1977) was reexa- mined and discovered to differ somewhat from Young’s illustration, having the paraphysis branched before the middle and shorter abdominal apodemes. In the absence of any known differences, in- cluding the number of spots on the pronotum, C. bipunctata is considered to be a junior synonym of C. lutea. It seems that northern populations, from Mexico, have one spot on the pronotum, while southern populations, from Gua- temala to Costa Rica, have two. Never- theless, Nielson and Godoy (1995) wrote that “C. lutea” (with the one-spotted pronotum) occurs in Costa Rica. More collecting may expose mixed popula- tions. Species LIstT C. andamira Young 1977: 1044. Vene- zuela. C. caucana Young 1977: 1043. Colom- bia. C. cunahua Young 1977: 1046. Peru. C. insularis Young 1977: 1041. Vene- zuela, Trinidad, Tobago. C. lutea (Signoret). Mexico, Guatemala, El Salvador, Nicaragua, Costa Rica. Tettigonia lutea Signoret 1855: 773. Caldwelliola bipunctata Nielson and Godoy 1995: 194, new synonymy. C. reservata (Fowler). Honduras, Belize, Costa Rica, Panama, Colombia. Tettigonia reservata Fowler 1900: 267. C. selvola Young 1977: 1046. Peru. C. sinuata, n. sp. Honduras. C. solimoeana Young 1977: 1048. Peru, west-central Brazil. C. tharma (Young). Ecuador, Peru, Brazil. Chlorogonalia tharma Young 1977: DiiGe C. trilineata, n. sp. Ecuador. Kry TO MALES OF CALDWELLIOLA 1. Crown of head with conspicuous, separate median black spot, without longitudinal or transverse black lines (Figs. 17—18, 20—22); aedeagus in lateral view broadest at mid length or, if not, pronotum with pale collar along anterior manetasya ult. tye ae 2 Crown of head without discal spot (Figs. 16, 19, 24); aedeagus broadest basal- ly or distally or, 1f at mid length (¢rilineata, n. sp., Fig. 35), then shaft apex emarginate 5 2. Sternal abdominal apodemes very slender, not capitate, extending beyond posterior ioeheeatnN (Oi Sinermayvhonl MIM (Varker S))) 9 sooo 6 46 o 3 538 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Sternal abdominal apodemes not extending posteriorly to margin of sternum III, Capitateionrmot) 4-0, 6 aie oe eee 4 Frontoclypeus black below dorsal, distal macula, with which it is contiguous (Fig. 15); more than 8.5 mm in length including forewings in repose...... tharma Frontoclypeus pale green except dorsal, distal black macula (Figs. 8-14); less than 6.5 mm in length including forewings in jor (Oa, WS) sscacoaccbance reservata Pronotum and mesonotum without black spots (Pig. 18) 2:5. 58 keen eecieeeee insularis Pronotum and mesonotum each with pair of blacksspots (Fis) 17) a ee caucana Sternal abdominal apodemes longer than WAGE) 2 ah, Pies eos oe yaa en 6 Sternal abdominal apodemes wider than longi): MSPS. - ORS See ee 8 Length less than 5.5 mm; without conspic- uous black spots on pronotum or mesono- (GUITARS IK) Gas coos ceo 6c andamira Length more than 6.0 mm; with or without black spots on pronotum or mesonotum 7 Entirely pale yellow except with 1 or 2 median black spots near mid length of pronotum and pair of black spots on mesonotum (Figs. 19526) -2=...-.2- lutea Mostly dark green, crown with median line and postfrontal sutures black (Fig. 24) . plghge tt ns oleae ogee cee nee trilineata, n. sp. Pysoter rounded distally == - = =--=- selvola Pygofer subacute distally (Figs. 36, 38) . . 9 Posteroventral pygofer margin concave; head with black spot at apex ..... cunahua Posteroventral pygofer margin linear or convex; head without black spot at apex. 10 .Head without black transverse line across crown; pygofer dorsal margin in lateral view uneven, almost linear and with its highest poimt atiapex nee solimoeana Head with solid, black, transversely sinu- ous line across crown; pygofer dorsal margin in lateral view weakly and evenly concave, and with its highest point before apex (aie. 38) ce cea sinuata, N. sp. Caldwelliola trilineata McKamey, new species (Figs. 15, 24, 34-36) Diagnosis.—Crown lacking median spot, with median line and postfrontal sutures black; pronotum and mesonotum lacking dark maculae. Description.—Length including fore- wings in repose: Male 7.2 mm. Head: VOLUME 108, NUMBER 3 5390 Figs. 8-15. Anterior views of Caldwelliola species. 8, C. andamira. 9, C. caucana. 10, C. insularis. 11, C. lutea. 12, C. selvola. 13, C. reservata (oblique view). 14, C. sinuata. 15, C. trilineata. Figs. 16-24. Detail dorsal views of Caldwelliola species. 16, C. andamira. 17, C. caucana. ites, (Es insularis. 19, C. lutea from Yepocapa, Guatemala. Figs. 25-26. Dorsal habitus views of two 20-21, C. reservata from Central America and — variably colored Caldwelliola species. 25, C. anda- Colombia, respectively. 22, C. tharma. 23, C. mira (compare with Fig. 16). 26, C. lutea from sinuata. 24, C. trilineata. Veracruz, Mexico. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 27-34. Median length of crown 0.6 interocular width and 0.3 transocular width; ocelli located behind a line between anterior eye angles, each slightly closer to adja- cent anterior eye angle than to median line of crown. Thorax: Pronotum lateral margins parallel. Apodemes of abdomi- nal sternum II slightly longer than wide; twisting to appear falcate in dorsal view and curved medially; apices separated from each other by approximately their width, not attaining posterior margin of tergum III. Male terminalia: Pygoter (Fig. 36) broadly produced dorsoposter- iorly, subacute, with large macrosetae scattered toward margins posteriorly and dorsally, lacking cluster of smaller macrosetae; subgenital plate abruptly constricted at mid length; connective broadly triangular; style short, not at- Lateral habitus views of Ca/dwelliola species. 27, C. andamira. 28, C. caucana. 29, C. insularis. 30, C. lutea. 31, C. reservata (Colombia). 32, C. selvola (forewing detached). 33, C. sinuata. 34, C. trilineata. taining constriction of subgenital plate, apex bluntly rounded. Aedeagus (Fig. 35) with ventral half of shaft narrow throughout in ventral view, in lateral view with distinct convexity in middle—in ventral view wider than ventral half of shaft; apex weakly emar- ginate, ventral margin with short con- cavity preapically; paraphyses present, bifurcate, long and gradually narrowing to apex, but not exceeding pygofer margins. Coloration: Mostly dark green; crown of head with median line and postfrontal sutures black (Fig. 24); fron- toclypeus (Fig. 15) with dorsal, distal maculae present but obscured by mostly dark brown color of remaining surface; anterior portion of pronotum concolor- ous with posterior portion. Female: Un- known. VOLUME 108, NUMBER 3 (37) (38 Figs. 35-38. Lateral views of aedeagus and male pygofer, respectively, of new Caldwelliola species. 35—36, C. trilineata. 37-38, C. sinuata. Material examined.—Holotype (Na- tional Museum of Natural History, Smithsonian Institution, Washington, D.C.) male with labels ““Archidona/ (Ecuad))/ RoiAiaensch S.,’ and “HOLO- TYPE/ Caldwelliola/ trilineata/: McKa- mey.” Paratype male, same locality and collection as holotype and with paratype label. Etymology.—The specific name is Latin for 3-lined, referring to the black longitudinal lines on the crown. Comments.—Although lacking a black spot on the crown as in C. reservata and C. insularis, C. trilineata has an aedeagus closely resembling those of both these species. The crown markings most close- ly resemble those of some C. andamira (Fig. 16). Caldwelliola sinuata McKamey, new species (Figs. 14, 23, 33, 37-38) Diagnosis.—Head with solid, black, transversely sinuous line across crown. Description.—Length including forew- ings in repose: Male 5.4mm, female 5.8 mm. Head: Median length of crown 0.7-0.8X interocular width and 0.4 transocular width; ocelli located behind a line between anterior eye angles, each slightly closer to adjacent anterior eye 541 angle than to median line of crown. Pronotum. Lateral margins parallel; apo- demes of abdominal sternum II slightly wider than long, not attaining posterior margin of tergum III. Male terminalia: Pygofer (Fig. 38) broadly produced pos- teriorly, subacute, dorsal margin weakly and evenly concave in lateral view, with large macrosetae scattered across poste- rior half and with cluster of smaller macrosetae near ventral margin at mid length; subgenital plate gradually con- stricted at mid length; connective broad- ly triangular; style short, not attaining constriction of subgenital plate, apex acute. Aedeagus in lateral view (Fig. 37) subparallel throughout length, though narrowest near mid _ length, curved ventrally then dorsally; apex produced posteriorly; paraphyses pres- ent, bifurcate, gradually narrowing, long, exceeding pygofer margins dorsoposter- iorly. Coloration: Male and female head yellow green throughout except with with solid, black, transversely sinuous line across crown (Figs 14, 23); prono- tum lacking dark maculae, anterior portion yellow along margin; mesono- tum and scutellum with midline black. Material examined.—Holotype (Na- tional Museum of Natural History, Smithsonian Institution, Washington, D.C.) male with labels “‘“Tegucigalpa/ Honduras/ MT30,[19)17;" “Ra. Dyer Colt/ Noe 22997/ 23296 sand *HORO- TYPE/ Caldwelliola/ sinuata/: McKa- mey. Paratypes (National Museum of Natural History): 2 males, same data as holotype, 1 female with labels “‘La Ceiba/ Honduras,/* “Pas Dyer ‘Coll and “11198to/ 11334/ vu-19-[19]16.” All paratypes labeled as such. Etymology.—The specific name refers to the aforementioned diagnostic color- ation of the crown. ACKNOWLEDGMENTS I thank M. Touchet for production of the photographs and M. A. Solis, D. R. 542 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Davis, and two anonymous reviewers for helpful comments on the manuscript. LITERATURE CITED Fowler, W. W. 1900. Order Rhynchota. Suborder Hemiptera-Homoptera (continued). Biologia Centrali Americana 2: 265-272. McKamey, S. H. 2006. Review of the Neotropical leafhopper genus Chlorogonalia (Hemiptera: Cicadellidae), with notes on the genus Cald- welliola. Proceedings of the Entomological Society of Washington 108: 611-618. Nielson, M. W. and C. Godoy. 1995. A new genus and new species of cicadelline leafhoppers from Costa Rica (Cicadellidae: Cicadellinae). Contributions on Entomology, International 1(2): 189-204. Signoret, V. 1855. Revue iconographique des Tettigonides. Annales de la Société Entomolo- gique de France 3(3): 765-836. Young, D. A., Jr. 1977. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New World Cicadellini and the genus Cicadella. North Carolina Agricultural Exper- iment Station Technical Bulletin 239. 1,135pp. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 543-549 SPECIES COMPOSITION AND ABUNDANCE OF FLEA BEETLES (COLEOPTERA: CHRYSOMELIDAE) ASSOCIATED WITH MOIST HABITATS IN ISPARTA AND BURDUR PROVINCES, TURKEY ALI GOK AND EBRU GUL ASLAN Stileyman Demirel University, Science and Art Faculty, Biology Department, 32260 Isparta, Turkey(e-mail: aligok@fef.sdu.edu.tr; egul@fef.sdu.edu.tr) Abstract.—An one-year survey was conducted on flea beetles associated with moist habitats in Isparta and Burdur provinces in the central part of the ‘Lake District’ in Turkey. Specimens were collected from various humid habitats divided into five major groups: moist meadows, marshes, moist pastures, lake sides, and stream banks. A total of 4,443 individuals belonging to 45 species were found, some of which are important pests of several wild plants. Habitat preference and relative abundance data are provided for each species. Among the genera collected in the study, Chaetocnema Stephens was the most abundant genus typically occurring in moist habitats. There was no significant difference in flea beetle species composition among the different habitat types. The highest species abundance and composition occurred in stream banks and moist meadows, whereas marshes have the lowest species abundance and composition. Key Words: Flea beetles, moist habitats, species composition, abundance, Turkey Alticinae is the largest subfamily of weeds belonging to Euphorbiaceae Chrysomelidae and can be distinguished from other Chrysomelidae by _ their greatly enlarged hind femora (Furth 1988). It as represented: by. 59. genera and more than 1,000 species in the Palearctic Region (Konstantinov and Vandenberg 1996). Alticines are highly specialized phy- tophagous insects and often follow their host plant distributions (Furth 1979, Jolivet 1988). Some of them are serious economic pests, causing direct damage and transmitting viruses; however, sev- eral, including species of Longitarsus Berthold and A/tica Muller have been used successfully in the biological control of weeds (Booth et al. 1990). Some species of Aphthona Chevrolat are also commonly used in biological control of (Konstantinov 1998, Jonsen et al. 2001, Konstantinov et al. 2001). Flea beetles are common in almost all types of habitats. The richest flea beetle communities occur in open areas near forests or scrublands often associated with rivers or lakes and in various kinds of meadows (Konstantinov and Vanden- berg 1996). Although most alticines are terrestrial, several of them, e.g., Altica, Chaetocnema Stephens and Longitarsus, are attracted by subaquatic plants along streams (Jolivet and Verma 2002). The Turkish Alticinae fauna consists of more than 250 species with recent contributions by Aslan et al. (1999, 2002) and Gok et al. (2002, 2003, 2004). Most of these works are faunistic studies. The Alticinae in Turkey are relatively poorly 544 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON studied from the ecological point of view including habitat use and host plant associations (Cilbiroglu and Gok 2004). Therefore, we regard this study as a pre- liminary step for future ecological studies on the Alticinae of Turkey. Burdur and Isparta are located in the central part of the ‘Lake District’, which is one of the most important wetland areas of Turkey. This region includes a series of lakes of different sizes and ecological conditions. Its topography 1s composed of narrow and long mountain ranges, and depression areas in between (Yildirim and Kebapg1 2004). This work is part of the authors’ studies on the Alticinae of Isparta and Burdur provinces, and the purpose is to assess flea beetle composition and abun- dance in habitats close to water. MATERIALS AND METHODS Site description.—The study was con- ducted in Isparta and Burdur provinces (between 29°33’ and 31°20'E and 37-03" and 38°05’'N), which are in the central part of the Lake District in Turkey, at the average altitude of 1,000 m. The area is located between central Anatolia and the Mediterrenean region of Turkey; therefore, both arid and temperate cli- matic conditions are present. The sam- pled moist habitats are categorized into five groups as follows: Moist meadows (MM): Includes nat- ural areas along or near a watercourse dominated by gramineous vegetation and scattered small shrubs. These areas are drier than marshes with many more plant species. Plants from Lamiaceae, Asteraceae, Rosaceae, and Salicaceae are common. Marshes’ (MA): Consists of open swampy areas that are closely associated with water, with rushes and sedges as the dominate plants. Plant species include Phragmites spp. (Poaceae), Typha spp. (Typhaceae), Bolboschoenus spp. (Cyper- aceae), and Carex spp. (Cyperaceae). There are also some other plants belong- ing to Asteraceae, Euphorbiaceae, and Brassicaceae. Moist pastures (MP): These are open areas located at moist, high elevations on cool mountain slopes at an average of 1,000-1,500 m. The areas are divided with small springs, and mainly include annual herbaceaus plant communities. Other parts away from the springs, are relatively drier. Dominant plants are species of Brassicaceae and rarely Ra- nunculaceae. Lake sides (LS): Includes humid areas adjacent to lakes, often surrounded by agricultural lands, especially wheat fields. Asteraceae and Poaceae are the dominant plant families. There are also orchards in surroundings. Stream banks (SB): These areas have the most diverse vegetation compared with other groups. There is a wide di- versity of plants, with many _ species coexisting in a small area. The areas are characterized by woodshrub vegetation and their undergrowth of herbaceous plants. The dominant woodshrub cover consists of the following families: Faga- ceae, Betulaceae, Salicaceae, and Rosa- ceae. The herbaceous cover is represent- ed by plants of the Lamiaceae, Fabaceae, Asteraceae, Convolvulaceae, Scrophu- lariaceae, and Boraginaceae. Sampling.—Fieldwork included sur- vey of two representative stations for each of the five habitats (Fig. 1). The ten stations were sampled biweekly from March through September 2003 when the adult flea beetles are active. The stations were sampled by collecting at each site for 60 minutes with a sweep net. Sampling was made randomly from all existing plants. A total of 4,443 beetles were collected and identified using the keys by Warchalowski (2003). The num- ber of adult individuals was counted for each species. The relative abundance of each species was determined by using the sample formula nj/Ny- < 100 (nj = VOLUME 108, NUMBER 3 5 TURKEY > — Cee.) Tefenni@ D Korkuteli @ MP1 @ Alnnyayla Fig. 1. Nn £ Nn @ Yalvag $.Karaagag K6prii River Aksu River HB @ Settlements A Stations The study area with stations: MM1, MM2, moist meadows; MA1, MA2, marshes; MP1, MP2, moist pastures; LS1, LS2, lake sides; SB1, SB2, stream banks. individuals of species 1; Nr = _ total individuals of all species). The specimens are deposited in the Department of Biology, Faculty of Art and Science, Suleyman Demirel University. RESULTS AND DISCUSSION In the study, forty-five species from ten genera of flea beetles were found associated with the five moist habitat types. Species associations with selected habitats are in Table 1. A list of flea beetle species collected is in Table 2, together with their number and relative abundance. Longitarsus and Chaetoc- nema were the largest genera with 12 and 10 species, respectively. Podagrica and Dibolia were the least species-rich genera with one species each. Although Longitarsus has the highest number of species, Chaetocnema was the most abundant genus, representing 36.3% of the total number of individuals captured during the study period. The number of species found in each habitat are as follows, in descending order: stream banks (SB) 32, moist meadows (MM) 28, moist pastures (MP) 21, lake sides (LS) 20, and marshes (MA) 19. The species richness did not vary significantly from habitat to habi- tat. More species were found in SB and MM, probably because these two habi- tats have the most diverse vegetation with many species of Lamiaceae, Aster- aceae, Salicaceae, and Graminae, which 546 Table 1. pastures; LS, lake sides; SB, stream banks. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Flea beetles associated with moist habitats. MM, moist meadows; MA, marshes: MP, moist Species MM MA MP LS SB Altica ancyrensis (Weise) Altica carduorum Guérin-Meéneville Altica deserticola (Weise) Altica lythri Aubé Altica oleracea (Linnaeus) Aphthona flaviceps Allard Aphthona nigriceps (Redtenbacher) Aphthona pygmaea Kutschera Chaetocnema concinna Marsham Chaetocnema conducta (Motschulsky) Chaetocnema coyei (Allard) Chaetocnema hortensis (Geoffroy) Chaetocnema major (Jacquelin du Val) Chaetocnema mannerheimi (Gyllenhal) Chaetocnema montenegrina Heikertinger Chaetocnema sahlbergi (Gyllenhal) Chaetocnema scheffleri (Kutschera) Chaetocnema tibialis (Mliger) Crepidodera aurata (Marsham) Crepidodera aurea (Geoffroy) Crepidodera lamina (Bedel) Dibolia occultans (Koch) Longitarsus bertii Leonardi Longitarsus kutschera Rye Longitarsus longipennis Kutschera Longitarsus luridus (Scopoli) Longitarsus lycopi (Foudras) Longitarsus melanocephalus (De Geer) Longitarsus parvulus (Paykull) Longitarsus pellucidus (Foudras) Longitarsus pratensis (Panzer) Longitarsus scutellaris (Rey) Longitarsus solaris Gruev Longitarsus succineus Foudras Neocrepidodera ferruginea (Scopoli) Neocrepidodera impressa obtusangula (J. Daniel) Phyllotreta aerea Allard Phyllotreta diademata (Foudras) Phyllotreta vittula (Redtenbacher) Podagrica menetriesi (Faldermann) Psylliodes anatolicus G6k and Cilbiroglu Psylliodes circumdatus (Redtenbacher) Psylliodes cupreus (Koch) Psylliodes magnificus Gruev Psylliodes reitteri Weise + ++t++4 + +t+t++¢+4++ + ++ +44 ~ +++ ¢t¢+¢ ¢4++4+ ++t+¢4t + + ++ +44 +++ +4 ++ +++ ++ 4+ 4+ 4+ 444 +++ 4+ +++ + +++ 44+ +++4+ + +++ + are characteristic food plant families of most Alticinae (Cilbiroglu and Gok 2004). According to Lawton (1983), changes of plant species composition and diversity influence the habitat com- plexity, hence the diversity and richness of phytophagous insect communities. The species numbers found in_ other three habitats (MP, LS, MA) were very similar (Fig. 2). As shown in Table 1, eight species are found in all habitats: Longitarsus long- VOLUME 108, NUMBER 3 Table 2. 547 Numbers and percent abundance of flea beetles collected. Species Number Collected Relative Abundance (%) Altica oleracea (Linnaeus) Chaetocnema coyei (Allard) Chaetocnema tibialis (Iliger) Altica deserticola (Weise) Chaetocnema concinna Marsham Longitarsus lycopi (Foudras) Longitarsus pratensis (Panzer) Neocrepidodera ferruginea (Scopoli) Crepidodera aurata (Marsham) Chaetocnema mannerheimi (Gyllenhal) Chaetocnema major (Jacquelin du Val) Longitarsus parvulus (Paykull) Chaetocnema conducta (Motschulsky) Longitarsus longipennis Kutschera Neocrepidodera impressa obtusangula (J. Daniel) Longitarsus pellucidus (Foudras) Longitarsus melanocephalus (De Geer) Dibolia occultans (Koch) Psylliodes cupreus (Koch) Aphthona pygmaea Kutschera Phyllotreta aerea Allard Phyllotreta vittula (Redtenbacher) Longitarsus succineus Foudras Longitarsus luridus (Scopoli) Phyllotreta diademata (Foudras) Podagrica menetriesi (Faldermann) Aphthona flaviceps Allard Psylliodes anatolicus G6k and Cilbiroglu Chaetocnema hortensis (Geoffroy) Psylliodes magnificus Gruev Longitarsus bertii Leonardi Crepidodera lamina (Bedel) Chaetocnema scheffleri (Kutschera) Chaetocnema sahlbergi (Gyllenhal) Longitarsus scutellaris (Rey) Psylliodes circumdatus (Redtenbacher) Crepidodera aurea (Geoffroy) Altica carduorum Guérin-Meéneville Altica ancyrensis (Weise) Longitarsus kutschera Rye Aphthona nigriceps (Redtenbacher) Chaetocnema montenegrina Heikertinger Longitarsus solaris Gruev Altica lythri Aubé Psylliodes reitteri Weise 585 SSF 542 1222. 379 8.53 Dif 6.12 214 4.82 180 4.05 WS 3.94 174 592 163 3.67 151 3.4 141 3) I17/ 122 2.74 104 2.34 102 Z29 87 1.96 82 1.85 81 1.82 81 1.82 76 el AZ 1.62 Tz 1.62 62 139 51 WES 48 1.08 48 1.08 32 Oz 3] 0.7 30 0.67 28 0.63 Dy 0.61 Dil 0.61 24 0.54 24 0.54 2p) 0.5 22 0.5 21 0.47 18 0.41 7 0.38 13 0.29 12 O27 1] 0.25 8 0.18 6 0.14 5) 0.11 ] 0.02 ipennis, Longitarsus luridus, Altica oler- acea, Chaetocnema concinna, Chaetoc- nema conducta, Chaetocnema_ coyel, Chaetocnema tibialis, and Psylliodes cu- preus. Four species were found in only one habitat type; of which two were in the genus Phyllotreta, and two in Psy- lliodes: Phyllotreta diademata, Phyllo- treta vittula, Psylliodes circumdatus and Psylliodes reitteri. Phyllotreta mainly prefer plants from Brassicaceae and Resedaceae which grow 548 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MM MA Fig. 2. pastures; LS, lake sides; SB, stream banks. in cultivated areas, roadsides, orchards, and shrubs (Mohr 1966; Furth 1979; Matsuda 1988, Nielsen 1988). The low density of Brassicaceae in moist habitats may be the reason for limited habitat occurrence of Phyllotreta. Many species of Psylliodes are also Brassicaceae feed- ers (Furth 1983). The restriction of each Psylliodes species to one habitat may be temporary or accidental. The most abundant species in number of individuals were Altica Chaetocnema_ coyei, and Chaetocnema ribialis*with t379o, 12:29; and 8.53%, respectively. Psylliodes reitteri was the rarest species, represented by a single individual. oleracea, Density and species composition of phytophagous beetles are affected by many factors, such as vegetation, humid- ity, temperature and host plants (Gillott 1995). Our studies revealed that moist habitats are preferred by a large number of moisture-loving flea beetles. MP Species percentage of the five habitats studied. MM, moist meadows; MA, marshes; MP, moist dale LS 3B LITERATURE CITED Aslan, I., B. Gruev, and H. Ozbek. 1999. A preliminary review of the subfamily Alticinae (Coleoptera, Chrysomelidae) in Turkey. Turk- ish Journal of Zoology 23: 373-414. Aslan, I., R. Beenen, and H. Ozbek. 2002. New records of leaf beetles from Turkey (Coleop- tera, Chrysomelidae). Entomologische Blatter 98: 231-235. Booth, R. G., M. L. Cox, and R. B. Madge. 1990. Gui- des to Insects of Importance to Man 3. Coleop- tera. University Press, Cambridge, U.K. 384 pp. Cilbiroglu, E. G. and A. Gok. 2004. Flea beetles (Coleoptera: Chrysomelidae) of Isparta, Tur- key, with habitat use and host plant associa- tions. Proceedings of the Entomological Soci- ety of Washington 106: 858-864. Furth, D. G. 1979. Zoogeography and host plant ecology of the Alticinae of Israel, especially Phyllotreta; with descriptions of three new species (Coleoptera: Chrysomelidae). Journal of Zoology 28(1): 1—37. 1983. Alticinae of Israel: Psy/liodes (Co- Journal of Israel leoptera: Chrysomelidae). Israel Entomology 17: 37-58. 1988. The jumping beetles (Alticinae)—The metafemoral spring. pp. 285-297. In Jolivet, P., apparatus of flea E. Petitpierre, and VOLUME 108, NUMBER 3 T. H. Hsiao, eds. Biology of Chrysomelidae. Kluwer Academic Publishers, Dordrecht, The Netherlands. 615 pp. Gillott, C. 1995. Entomology. Plenum Press, New York. 798 pp. Gok, A., E. G. GCilbiroglu, and Y. Ayvaz. 2003. New contribution to the Turkish flea beetle fauna (Coleoptera: Chrysomelidae: Alticinae). Israel Journal of Zoology 49: 319-320. . 2004. New records of flea beetles (Co- leoptera: Chrysomelidae) from Turkey. Phyto- parasitica 32: 360-362. Gok, A., E. G. Cilbiroglu, Y. Ayvaz, and M. Z. Yildirrm. 2002. Two new records for the Turkish flea beetle fauna: Phyllotreta reitteri Heik., 1911 and £&pitrix dieckmanni Mohr, 1968 (Coleoptera, Chrysomelidae, Alticinae). Israel Journal of Zoology 48: 254-255. Jolivet, P. 1988. Food habits and food selection of Chrysomelidae. Bionomic and evolutionary perspectives, pp. 1-24. Jn Jolivet, P., E. Petitpierre, and T. H. Hsiao, eds. Biology of Chrysomelidae. Kluwer Academic Publishers, Dordrecht, The Netherlands. 615 pp. Jolivet, P. and K. K. Verma. 2002. Biology of Leaf Beetles. Intercept Publishers, UK. 332 pp. Jonsen, I. D., R. S. Bourchier, and J. Roland. 2001. The influence of matrix habitat on Aphthona flea beetle immigration to leafy spurge patches. Oecologia 127: 287-294. Konstantinov, A. S. 1998. Revision of the Palearc- tic Species of Aphthona Chevrolat and Cladis- tic Classification of the Aphthonini (Coleop- tera: Chrysomelidae: Alticinae). Associated Publishers, Gainesville, Florida. 429 pp. 549 Konstantinov, A. S. and N. J. Vandenberg. 1996. Handbook of Palearctic Flea Beetles (Coleop- tera: Chrysomelidae: Alticinae). Contributions on Entomology, International 1(3): 237-439. Konstantinov, A. S., M. G. Volkovitsh, and M. Cristofaro. 2001. New data on Palearctic Aphthona (Coleoptera: Chrysomelidae) with description of a new species: Taxonomic and faunistic results of biological control explora- tion. Entomological News 112: 31-41. Lawton, J. H. 1983. Plant architecture and the diversity of phytophagous insects. Annual Review of Entomology 28: 23-39. Matsuda, K. 1988. Feeding stimulants of leaf beetles, pp. 41—56. Jn Jolivet, P., E. Petitpierre, and T. H. Hsiao, eds. Biology of Chrysome- lidae. Kluwer Academic Publishers, Dor- drecht, The Netherlands. 615 pp. Mohr, K. H. 1966. Chrysomelidae, pp. 95-299. In Freude, H., K. Harde, and G. A. Lohse, eds. Die Kafer Mitteleuropas, Krefeld. 299 pp. Nielsen, J. K. 1988. Crucifer-feeding Chrysome- lidae: Mechanism of host plant finding and acceptance, pp. 25-40. Jn Jolivet, P., E. Petitpierre, and T. H. Hsiao, eds. Biology of Chrysomelidae. Kluwer Academic Publishers, Dordrecht, The Netherlands. 615 pp. Warchalowski, A. 2003. Chrysomelidae: The Leaf Beetles of Europe and the Mediterranean Area. Natura optima dux Foundation, Wars- zawa. 600 pp. Yildirim, M. Z. and U. Kebapgi. 2004. Slugs (Gastropoda: Pulmonata) of the Lakes Region (GOoller Bélgesi) in Turkey. Turkish Journal of Zoology 28: 155-160. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 550-558 A NEW SPECIES OF EMERALD ASH BORER PARASITOID FROM CHINA BELONGING TO THE GENUS TETRASTICHUS HALIDAY (HYMENOPTERA: EULOPHIDAE) ZHONG-q!l YANG, JOHN S. STRAZANAC, YAN-XIA YAO, AND XIAO-yl WANG (ZY5 Ye XW) The Key laboratory. of Forest Protection, State roenesiry Administration of China, Research Institute of Forest Ecology, Environment and Protection, Chinese Academy of Forestry, Beijing 100091, China (e-mail: yangzq@ forestry.ac.cn); (JSS) Plant and Soil Sciences, West Virginia University, Morgan- town, WV 26506, U.S.A. Abstract.— Tetrastichus planipennisi Yang, new species, is described and illustrated based on individuals reared from mature larvae of the emerald ash borer, Agrilus planipennis Fairmaire (Buprestidae), collected from their galleries in the trunks of Fraxinus mandshurica Rupr. in northeastern China. Tetrastichus planipennisi 1s a gregarious endoparasite, with 56—92 individuals developing in a single host larva and observed parasitism rates from 32 to 65%. Based on reared field-collected material, the female to male ratio is 2.5:1. It 1s a koinobiont parasitoid and overwinters as a mature larva. The new parasitoid is a potential biocontrol candidate for A. planipennis. Key Words: Hymenoptera, Eulophidae, TJetrastichus planipennisi, new species, emerald ash borer, Agrilus planipennis Fairmaire, parasitoid, biology, biocontrol The emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Bu- prestidae), is native to eastern Asia, including Japan, Korea, China, Inner Mongolia and the Russian Far East. EAB larval feeding damages ash trees (Oleaceae, Fraxinus spp.) by forming zigzag galleries in the phloem and outer sapwood. During heavy infestations, in- dividual galleries coalesce, girdling and killing branches and/or entire trees. EAB mainly attacks F. mandshurica Rupr. and F. rhychophylla Hance. in northeastern China (Liaoning, Jilin, and Heilongjiang provinces) and F. velutina Torr. in northern China (Tianjin Municipality, Shandong, and Hebei provinces). His- torically, EAB in China has been con- sidered a minor pest, but our early investigations found in some locations, such as the seacoast forest-belt of Tian- jin, nearly 80% of the ash trees were infested with mortality 35%. In 2002, EAB was discovered attacking ash trees in southeastern Michigan, and has since spread to adjacent areas of Windsor, Canada, northwestern Ohio (Lucas and DeFiance counties), and northeastern Indiana. In those areas, EAB is devas- tating both ornamental and forest trees (McCullough 2002, Plant Health Divi- sion 2005). A joint cooperative program surveying for natural enemies of EAB in China was initiated in 2003 to screen potential biocontrol agents for use in North VOLUME 108, NUMBER 3 America and China. Several parasitoids have been found, the first reported and laboratory maintained is Spathius agrili Yang (Hymenoptera: Braconidae) (Yang et al. 2005). The second candidate for an EAB biocontrol program is another species of parasitic wasp in the genus Tetrastichus Haliday (1844) (Hymenop- tera: Eulophidae). This species is an abundant natural enemy of EAB in northeastern China with parasitism rates ranging from 32 to 65%. It is a koino- biont, allowing parasitized EAB larvae to continue development. Typical of other koinobionts, it is an endoparasite, and like many other eulophids, numer- ous larvae feed gregariously on a single host individual. The Tetrastichinae can be distin- guished from other Eulophidae by the following characters: postmarginal vein generally absent or much reduced; scu- tellum with two pairs of longitudinal grooves and 2 pairs of strong setae; female antenna with funicle 3-segmented, male antenna with funicle 4-segmented. Tetrastichus Haliday can be _ distin- guished from other Tetrastichinae genera mainly by the combination of the fol- lowing two characters: forewing with submarginal vein having usually | (rare 2) dorsal setae; propodeum with an inverted Y-shaped paraspiracular carina. Other characters to help define this genus are given by Graham (1991) and LaSalle (1994). The genus Tetrastichus 1s cosmopoli- tan with 482 species (Noyes 2003). Graham (1991) revised the 45 European species and LaSalle (1994) listed 21 North American species. The literature on Asian species is less complete. Kost- jukov (1995) produced a key for 155 species from the Russian Far East, describing many new species. In China 26 species have been reported and de- scribed by various authors (Liao et al. 1987; Sheng 1995; Sheng and Wang 1992, 1995; Sheng and Shen 1996; Yang Sail 1996; Zhu and Huang 2001, 2002; He 2004). The following description of a new Tetrastichus species includes biological observations related to its EAB bio- control potential. MATERIALS AND METHODS During 2003 and 2004, surveys were conducted in Liaoning, Jilin, and Hei- longjiang provinces for natural enemies of EAB. Stressed F. mandshurica with dead branches were located, most with trunk diameters of 5-15 cm. Portions of their trunk bark were peeled back to locate and trace the EAB larval galleries. When mature parasitoid larvae were located, they usually occupied the terminal end of their host gallery and laid adjacent to their mummified host. These wasp larvae and additional EAB larvae were collected and brought into the laboratory. Five wasp larvae and a piece of filter paper moistened with distilled water were placed into each vial (12 mm diameter < 74 mm length). In- dividual field sampled EAB larvae were placed in separate vials with a section of the ash twig (10 mm diameter and 60 mm length) with a trough carved between the bark and sapwood for the larva to feed and be observed. All vials were tightly plugged with cotton and maintained at ambient temperature (about 23°C) with the filter paper moist- ened every three days. During the EAB larval rearing, the bark flap over the twig trough was pulled back to observe the development of the EAB larva and determine if it was parasitized. When parasitoids emerged they were killed and point mounted for taxonomic study. Specimens were examined with an Olym- pus SZH10 Stereo Microscope. Micro- graphs were taken with a ZOEL 550LV scanning electron microscope. Morpho- logical terms follow Boucek (1988) and Gibson (1997). Authorship of the new species is attributed solely to Yang Zhong-qi. 552 Tetrastichus planipennisi Yang, new species (Figs. 1—26) Female (Figs. 1—26).—Length 2.7— 4.1mm. Body dark green with bronze green tint. Antenna fuscous, except scape fulvous with apical 1/4 brownish and pedicel slightly reddish. Legs with coxae and femora same color as body except trochanters fuscous; tips of femora, tibiae and tarsi yellow, and last tarsal segment brownish. Wings hyaline, vena- tion testaceous. Head: Wide as mesosoma, in dorsal View swidth 2:3>< jlength, POL, loux OOL, OOL equals OD. Temple short, 0.22 length of eyes. Head (Fig. 1) ante- rior view slightly wider than tall (25:22). Scrobes moderately deep, nearly smooth. Parascrobal areas with numerous pilifer- ous punctures near eyes. Eye sparsely pubescent, separated by 1.3—1.4x their length, length 1.9 width. Malar space 0.8 length of eye, malar sulcus nearly straight. Face medially convex. Mouth width 0.9-1.2* malar space. Clypeus with anterior margin deeply incised medially. Antennal (Fig. 1) toruli lower margin located on ventral line of eyes. Antenna (Fig. 2) with scape as long as eyes; scape length 4.8 its width, ex- tending beyond median ocellus, reaching vertex; pedicel plus flagellum 1.2—1.28x width of head; pedicel 2.2 as long as wide, distinctly shorter than F, (5.5:9); funicle (Fig. 3) much thicker than pedi- cel; F; as broad as F> and F3, but shorter than these segments (8:9:9); F, 2.8 and F, 3 3.0 as long as respective widths; clava (Fig. 4) slightly wider than flagel- lum (18:16), length 2.8 its width, distinctly shorter than F>, plus F; (12.5:17), and with a weak constriction between C, and C>; tip moderately acute; lengths of © 14.-and Gy 0:8 their respective widths, C3 trochiform as long as wide; terminal spine 0.3 length of C,. Each funicle segment with 3 rows of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON longitudinal sensilla, and mushroom-like sensilla scattered on apical half (Fig. 5); clava with 2 rows of sensilla on C,_5, and | on C;, and the mushroom-like sensilla (Fig. 5) restricted to the apical margin of C,. Mesosoma: Moderately convex, length 1.4 width (Fig. 6). Midlobe of meso- scutum slightly shorter than wide (22:24), median line complete and posterior two- thirds distinct, 5—7 adnotaular setae present on each side. Mesoscutum and scutellum finely reticulate, areoles elon- gate Scutellum length 1.1—1.2x width, with submedian grooves separated from each other by 1.3 the distance either submedian groove from its nearest sub- lateral (= axillular) groove, area bor- dered by anterior and posterior margins of scutellum and submedian grooves with length 2.4 width; two pair of setae lateral to submedian grooves, first pair placed two-thirds along length of sub- median groove from anterior margin, second pair near posterior margin, both pairs subequal in length to distance between submedian grooves. Propodeum (Fig. 7) length twice that of dorsellum, width 6.5 length, median area reticu- late; median carina broad, adpetiolar strip well developed; spiracles ovate, distance from metanotum equal its length; callus with 6 setae anterior to spiracle and 2 posterior to spiracle. Prepectus moderately reticulate. Meso- pleuron, mesepisternum and_= meta- pleuron reticulate. Legs slender; hind femur (Fig. 8) length 4.2 width, three rows of setae along length: dorsal and ventral margins, and anterior surface; hind coxa with posterior margin coarsely rugose (Fig. 9). Forewing (Fig. 10) cos- tal cell 0.8 length of marginal vein and 9.4 width, bare above, with a row of hairs below; submarginal vein with 1-2 setae; marginal vein length 4.0 length of stigma, and 12.0 length of postmar- ginal vein; basal cell open, bare above, below with 2-4 hairs; basal setal line with VOLUME 108, NUMBER 3 | i ai 4 j « i Figs. 1-11. Tetrastichus planipennisi, female. 1, Frontal view of head. 2, Antenna. 3, Funicle. 4, Clava. 5, Funicle sensilla. 6, Mesosoma, posterior half of scutellum and dorsellum. 7, Propodeum. 8, Hind leg. 9, Hind coxa. 10, Forewing. 11, Hindwing. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 12-16. Tetrastichus planipennisi, female. 12, Metasoma dorsal view. 13, Metasoma lateral view. 14, Metasoma lateral view of proximal half of metasoma. 15, Apex of last tergite. 16, Apex of last tergite four cercal setae. about 9 setae; speculum small, extending as a narrow strip along two-thirds of marginal vein length, cubital setal line borders posterior margin. Hindwing as rials! ent. 1s Metasoma: Petiole ringlike, gaster (Figs. 12—13) in air dried specimens very long, strongly acuminate, depressed with dorsum sunken, 1.9-2.0* as long as head plus mesosoma, 5.0 as long as broad, and narrower than mesoscutum (27:32). All tergites delicately reticulate, and setae present except on dorsum of Ist and 2nd tergite. Tip of hypopygium (Fig. 14) at 0.12 length of gaster. Last tergite extremely long, 4.4—5.8 as long as basal broad; cercal setae (Figs. 15—16) with longest one about twice length of other three. Ovipositor projecting some- what, 0.1 length of last tergite (Fig. 15). Male (Figs. 17—25).—Differs from fe- follows: Length 1.6—2.2 mm; dark blue; scape black, wing venation fuscous. Frons with parascrobal male as body areas having piliferous punctures much sparser (Fig. 17). Antenna (Figs. 18) with 4 funicular segments; scape moderately flattened, 2.5 1.6 as long as rom VOLUME 108, NUMBER 3 Figs. 17-25. Tetrastichus planipennisi, male. 17, Frontal view of head. 18, Antenna. 19, Funicle 1 and 2. 20, Clava. 21, Mesosoma in lateral view. 22, Forewing. 23, Hindwing. 24, Metasoma in dorsal view. 25, Metasoma in lateral view. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 26. planipennis 4th instar larva. broad. Mesosoma (Fig. 21) 1.7 as long as broad. Forewing (Fig. 22) with costal cell slightly shorter than marginal vein, latter 3.1 length of stigma; submargin- al vein usually with 1 seta, occasionally PeeGastem (ees, 24. 925). im) air dred specimens strongly compressed, slightly shorter than head plus (66:70). Etymology.—The specific epithet is derived from the specific name of its host, Agrilus planipennis Fairmaire. Diagnosis.—The female is unique among other having an mesosoma Tetrastichus extremely long gaster. Superficially sim- ilar to Baryscapus agrilorum (Ratze- burg), a parasitoid of Buprestidae (A gri- lus viridis L.) in Germany (Graham 1991), the new species differs from B. agrilorum which has a curved malar sulcus; submarginal vein with 3—4 dorsal setae; midlobe of mesoscutum with 9—12 adnotaular and) last™ tereite cof, gaster shorter, 3.3—-4.1 * as long as wide. The new species is also close to Tetra- stichus telon (Graham) (Graham 1961) from Europe, but the syntergum of this species is much shorter, 1.9—2.5 as long as broad and the submedian lines of scutellum are equidistant from each other and the lateral lines. Four other Tetrastichus species have been reported to parasitize Agrilus and other buprestid beetles including 7. u/mi Erdos, 1954; T. Graham, 1991; T. agrili Crawford, 1914; and 7. jinzhouicus Liao, 1987. setae, agrilocidus Tetrastichus planipennisi females Mature larvae of Tetrastichus planipennisi visible through the integument of an Agrilus can be separated from these species by the long metasoma (1.9-2.0 as long as head and mesosoma combined) and the extremely long last tergite (4.45.8 basal width) of the gaster. Type material.—Holotype *, Weihe, Heilongjiang Province, Yang Zhong-qi and Wang Xiao-yi , 22 April 2004, ex: Its mature larva collected in the gallery of A. planipennis (Note: It was reared from a field collected overwintering mature larva found adjacent to a mummified larvae of A. planipennis which had attacked and formed a gallery in the trunk of Fraxinus mandshurica on 22 March 2004, and the adult wasp emerged 22 April 2004). Paratypes: 3 ¥, data is the same as holotype; 5 2%, 1 4, Jingyuetan, Jilin Province, larvae collect- ed 19 March 2004, adults emerged 23 April 2004, other data as holotype; 8 2, 31 3, Benxi, Liaoning Province, larvae collected 18 March 2004, adults emerged 17-19 April 2004, other data as holotype; following all collected in’ Jingyuetan Forest Park, Changchun, Jilin Province by Yang Zhong-qi and Wang Xiao-yi, 6 *. pupae collected from single EAB gallery 18 Sep. 2004, adults emerged 27 Sep. 2004; 8 2, 2 4, larvae collected from single EAB gallery 18 Sep. 2004, pupated 29 Sep. 2004, adults emerged 13 Oct. 2004; 20 2, 10 ¢@, larvae collected from single EAB gallery 18 Sep. 2004, pupated 30 Sep. 2004, adults emerged 15 Oct. (153.32) 2.13 964 Warvaer collected from single EAB gallery 18 Sep. 2004, VOLUME 108, NUMBER 3 pupated 29 Sep. 2004, adults emerged 15 Oct. 2004; 66 2, 26 4, reared from a single EAB mature larva, 12 Oct. 2004 (Note: the EAB larva collected in its gallery on tree of FL manschurica 18 Sep. 2004, the parasitoid larvae inside the EAB larva broke through the remaining translucent EAB larva body skin 25 Sep. 2004, pupated 28 Sep. 2004, adults emegecd 2 Oct: 2004)2 48 2a 30'S, same date as the previous series except parasitoid larvae broke through the EAB larva 28 Sep. 2004, pupated 5 Oct. 2004, adults emerged 22 Oct. 2004. The holotype and most paratypes of the new species are deposited in the Insect Museum of the Chinese Academy of Forestry, Beijing, China, with the exception the following paratypes: 6 &, 3 ¢, National Museum of Natural History, Smithsonian Institution,Wa- shington, DC, USA; 6 °, 3 6 Canadian National Collection of Insects (CNCI), Agriculture and Agri-Food Canada, @tiawa, @anada. and 12" o> West Virginia University Arthropod Collec- tion, Morgantown, WV, USA. Biological observations.—Tetrastichus planipennisi is a gregarious endoparasi- toid of Agrilus planipennis larvae, pro- ducing 56-92 individuals from a single host. Feeding by the parasitoid larvae inside the EAB larva did not kill their host until it became a late mature larval stage (4th instar), eventually completely consuming the host body contents until only the host larval translucent integu- ment remained, in which the mature maggot-like larvae (3-4 mm) are clearly visible (Fig. 26). Wasp larvae then exited their host and soon pupated adjacent to its intact membranous integument, which is typically near the end of the gallery. It took approximately 15 days for adult wasps to eclose. The newly emerged wasp chewed a hole through the bark to exit the host gallery, with other emerging wasps in the brood often using the same hole. We estimate that at least S// four generations are produced a year in northeastern China, with the last gener- ation overwintering as mature larvae inside their host’s gallery. The sex ratio of the emerging adults was skewed towards females 2.5:1. The longevity at 25°C of the adult parasitoids was about 15 days for females and 13 days for males. ACKNOWLEDGMENTS We thank Michael Gates and Michael Schauff (Systematic Entomology Labo- ratory, USDA); and John Huber and Gary Gibson (Canadian National Col- lection of Insects, Agriculture and Agri- Food Canada) for their early reviews of the manuscript. Thanks are also due to Richard Reardon and Wu Yun, Forest Health Technology Enterprise Team, Morgantown, WV, for their coordina- tion during the study, Juli Gould, Otis Pest Survey, Detection, and Exclusion Laboratory, Otis ANGB, MA, and Cynthia J. Fritzler, Morgantown, WV, for suggestions to improve the manu- script. We are very grateful to our Chinese colleagues Gao Chang-qi, Sun Shou-hui, Song Li-wen, Zhang Xiao-jun, Jilin Forestry Academy and Liu Xiu- ying and Zhang Kai-peng, Jingyuetan Forest Park, Changchun, Jilin Province, for their assistance during our field studies. The research is part of an on- going Chinese Academy of Forestry, Forest. (Service-pahEt Sand lea and Animal and Plant Health Inspection Service, USDA cooperative program. LITERATURE CITED Bouéek, Z. 1988. Australasian Chalcidoidea (Hy- menoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. C.A.B_ International, Wallingford, U.K. 832 pp. Crawford, J. C. 1914. Some new Chalcidoidea. Insecutor Inscitiae Menstruus 2: 180-182. Erdés, J. 1954. Eulophidae hungaricae indescrip- tae. Annales Historico-Naturales Musei Na- tionalis Hungarici (series nova) 5: 323—366. Gibson, A. P. 1997. Morphology and terminology, pp. 16-28. In Gibson, A. P., J. T. Huber, and J. B. Woolley, eds. Annotated keys to the genera of Nearctic Chalcidoidea (Hymenoptera). NRC Research Press, Ottawa, 791 pp. Graham, M. W. R. de V. 1961. New species of Aprosto- cetus Westwood (Hym.: Eulophidae) from Britain and Sweden. Opscula Entomologica 26: 4-37. . 1991. A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae): Revision of the remaining genera. Memoirs of the American Entomological Institute, Number 49, 320 pp. He, J. 2004. Hymenopteran Insect Fauna of Zhe- jiang. Science Press, Beijing, 1373 pp, 43 pls. Kostjukov, V. V. 1995. Eulophidae, pp. 291—505. In Ler, P. A., ed. Key to the insects of Russian Far East, Vol. IV. Neuropteroidea, Mecop- tera, Hymenoptera. Pt 2. Hymenoptera. Dal’- nauka, Vladivostok, 600 pp. (in Russian). LaSalle, J. 1994. North American genera of Tetrastichinae (Hymenoptera: Eulophidae). Journal of Natural History 28: 109-236. [eto De Xe pene ei xen banoaandeal ea @he»ne 1987. Hymenoptera: Chalcidoidea (1). Eco- nomic Insect Fauna of China Fasc. 34. Science Press, Beijing, 241 pp., 24 pls. (in Chinese). McCullough, D. G. 2002. Emerald ash borer. Pest Alert, USDA, Forest Service. NA-PR-07-02 (Rev. Dec. 2002). U.S. Department of Agri- culture, Forest Service, 2 pp. Noyes, J. S. 2003. Universal Chalcidoidea Database. The Natural History Museum, London. (http:// www.nhm.ac.uk/entomology/chalcidoids). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Plant Health Division. 2005. Emerald Ash Borer. Canada Food Inspection Agency. (http://www. inspection.gce.ca/english/plaveg/protect/pestrava/ ashfre/agrplae.shtm]). Sheng, J. K. 1995. Two new species of Tetra- stichinae from China. Acta Agriculture Uni- versitatis Jiangxiensis 17(1): 21—24. Sheng, J. and G. Wang. 1992. Descriptions of four and some other Chinese record species in the genus of Tetrastichus (Hymenoptera: Eulophi- dae). Plant Protection of Jiangxi 15: 35—36 (in Chinese). Sheng, J. and B. Shen. 1996. A new Chinese record species in the genus Tetrastichus (Hymenop- tera: Eulophidae, Tetrastichinae). Acta Agri- culturae Universitatis Jingxiensis 18: 419-420. (in Chinese). Yang, Z. 1996. Parasitic wasps of bark beetles in China. Science Press, Beijing, 363 pp., 11 pls. (in Chinese with English summary). Yang, Z., J. S: Strazanac; P. Me Marsh. (© van Achterberg, and W.-Y. Choi. 2005. The first recorded parasitoid from China of the emerald ash borer: a new species of Spathius (Hyme- noptera: Braconidae: Doryctinae). Annals of the Entomological Society of America 98(5): 636-642. Zhu, C. and D. Huang. 2001. A taxonomic study of Eulophidae (Hym.: Chale.) in Zhejiang. Acta Zootaxonica Sinica 26(4): 533—547. Zhu, C. 2002. A taxonomic study on Eulophidae from Guangxi, China (Hymenoptera: Chalci- doidea). Acta Zootaxonica Sinica 27(3): 583-607. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 559-564 A NEW SPECIES OF HYDROPSYCHE (TRICHOPTERA: HYDROPSYCHIDAE) FROM ALABAMA, WITH NOTES ON #H. FRISONI ROSS AND AN UNUSUAL HYDROPSYCHE FROM FLORIDA PAUL K. LAGO AND STEVEN C. HARRIS (PKL) Department of Biology, University of Mississippi, University, MS 38677, U.S.A. (e-mail: plago@olemiss.edu); (SCH) Department of Biology, Clarion University, Clarion, PA 16214, U.S.A. Abstract.—Hydropsyche fenestra, n. sp., belonging to the scalaris group and collected in central Alabama, is described and illustrated. Variation in Hydropsyche frisoni Ross is discussed with illustrations provided to facilitate identifications. In addition, figures of a deformed Hydropsyche from northern Florida, which appears to represent an undescribed species, are presented. Key Words: WHydropsychidae, Hydropsyche scalaris group, Alabama, Florida The genus Hydropsyche is well repre- sented in the southeastern United States, with 24 species reported, or about half of all known North American members of the genus. Our ongoing caddisfly studies in the region have uncovered a distinctive new species from Alabama, where 23 species have been previously recorded (Harris et al. 1991). The new species was collected in the west-central part of the state near the juncture of the Cumber- land Plateau and East Gulf Coastal Plain physiographic provinces. A second Hydropsyche encountered in our Alabama collections, Hydropsyche frisoni Ross, appeared to show distinct differences from the Illinois specimens illustrated by Ross (1944). After exam- ining over a thousand specimens from Alabama, we decided that either the Alabama material represented a new, but closely related, species or that H. frisoni exhibited considerable variation. Since Alabama represents the southern- most range of this species, either of the above scenarios seemed reasonable. Af- ter examining the holotype and numer- ous paratypes of H. frisoni, we concluded that we were dealing with one variable species. By providing additional illustra- tions, we hope to facilitate the identifi- cation of H. frisoni in the southeastern United States. Finally, in processing a number of caddisfly collections taken by Andy Rasmussen in the panhandle region of Florida, we encountered two specimens of a deformed Hydropsyche that appear to represent a new species. The male specimens were unusual in that one side of the genitalic structures were entire; the other side appeared contorted and mal- formed. With no complete individuals, we are unwilling to assign a species name at this time. We have, however, provided illustrations and a brief description hop- ing this might assist recognition as additional material becomes available. Morphological terminology follows that of Schmid (1980). The holotype of the new species will be deposited in the National Museum of Natural History, 560 Hydropsyche fenestra lanes, le PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hydropsyche fenestra, male terminalia. a, Lateral view. b, Tergum IX & X, dorsal view. c, Inferior appendage, caudal view. d, Apex of phallic apparatus, ventral view. f, Apex of phallic apparatus, dorsal view. Washington, DC. The specimens of the Florida Hydropsyche are deposited in the personal Gcollectionvol S» ©€x Haris at Clarion University, PA. Type material of Hydropsyche frisoni are deposited at the Illinois Natural History Survey, Cham- paign, IL. Additional material is de- posited in the collections of the authors. Hydropsyche fenestra Lago and Harris, new species (Eies 1) Holotype.—Male. Alabama: Tusca- loosa County, Big Sandy Creek off Co. Hwy 59, on Gulf States Power property, 6 km S>Coaling; 16 ‘August 1991, ,3 an M, nearly devoid of brownish coloration, except distally, thus producing a clear window. Eyes, in dorsal view, half as wide as interocular width. Abdominal segment IX with pro- nounced median dorsal crest. Tergum X somewhat difficult to describe because of deformity, but left side in best developed specimen with dorsal apical corner rounded in lateral view giving the im- pression that a median cleft exists (similar to the condition seen in various members of the group, such as H. scalaris Hagen and H. frisoni); preanal appendage appears as a rounded setifer- ous wart near posteroventral corner. Inferior appendage, in caudal view, with basal segment moderately long and straight, slightly enlarged apically, apical segment rounded apically and one-third 564 length of base; in lateral view, basal segment tapering from narrow base to broader apex, apical segment slightly constricted in basal half, apical half evenly convex on posterior margin and diagonally truncate on anterior margin, tapering to distinct point. Phallic appa- ratus tubular, nearly straight in ventral view, phallobase slightly inflated in apical two-thirds and distinctly con- stricted before apex; in lateral view, sinuate, swollen ventrally before apex; apex approximately as wide as phallo- base; lateral lobes tapered apically, slightly convex on dorsal — surface, straight ventrally, without mesal dome, mesal cavity very deep, approximately 85% open ventrally. Discussion.— Although obviously be- longing to the scalaris group, these specimens do not resemble to any extent previously described species, and we believe they represent a new species. There is a temptation to simply dismiss the specimens as deformed individuals of H. incommoda, the species most com- monly collected in the study area, based on the shape of tergum X and the inferior appendages. The length propor- tions of the inferior appendage segments are, however, quite different, being 4: 1 in H. incommoda and 3:1 in the de- formed individuals. Differences 1n phallic structure are also apparent, with the phallobase being somewhat more sinu- ous in the deformed specimens (compare Pics sowith Pic lin Plintyetvalei979): The mesal cavity in H. incommoda is shallow, but is very deep in the speci- mens in question; however, the ventral opening does not differ significantly in either shape or extent to which it is open. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hopefully, specimens without deformi- ties will be obtained through continued collecting in the area and the identity of these unusual individuals can be ascer- tained. Material examined.—FL: Liberty Co. Little Sweetwater Creek, Apalachicola Bluffs and Ravine Preserve, 5km N. Bristol, N30.2821-W84.5908, 26 Oct 1995, M. Pescador, A. Rasmussen, | 2; same locality, but label includes “‘site 6,” 19 May 1994, M. Pescador, A. Rasmus- sen, SiC. Hanssy lc. ACKNOWLEDGMENTS Colin Favret and Kathy Methven Zeiders of the [linois Natural History Survey, Champaign, were very helpful in providing specimens of Hydropsyche frisoni for this study. We thank Andy Rasmussen, Florida A&M _ University, Tallahassee, for bringing the Florida specimens to our attention. LITERATURE CITED Flint, O. S., Jr., J. R. Voshell, Jr., and C. R. Parker. 1979. The Hydropsyche scalaris group in Virginia, with the description of two new species (Trichoptera: Hydropsychidae). Pro- ceedings of the Biological Society of Washing- ton 92: 837-862. Haris. S) ©. PEs © Neilvand) Bakes acousSSir Caddisflies of Alabama. Geological Survey of Alabama, Bulletin 142: 1-442. Ross, H. H. 1938. Descriptions of Nearctic caddis flies (Trichoptera) with special reference to the Illinois species. Illinois Natural History Sur- vey, Bulletin 21: 101—183. . 1944. The caddis flies, or Trichoptera, of Illinois. Illinois Natural History Survey, Bul- letin 23: 1-326. Schmid, F. 1980. Genera des Trichopteres du Canada et Etats adjacents, part 7. In Les insectes et arachnides du Canada. Agriculture Canada, Ottawa, 296 pp. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 565-574 TAXONOMIC STUDY ON THE LEAFHOPPER GENUS BHATIA (HEMIPTERA: CICADELLIDAE) FROM CHINA SUQIN SHANG, LIN SHEN, YALIN ZHANG, AND HOUHUN LI (SS, LS, YZ) Key Laboratory of Plant Protection Resources and Pest Manage- ment of Ministry of Education, Entomological Museum, Northwest A&F University, Yangling, Shaanxi 712100, China (e-mail: yalinzh@cnipm.com, yalinzh@ yahoo.com.cn); (SS, HL) Department of Biology, Nankai University, Tianjin 300071, China. Corresponding author: Yalin Zhang Abstract.—Five new species of the leafhopper genus Bhatia Distant, B. hastata Shang and Shen, B. digitata Shang and Shen, B. flabellata Shang and Shen, B. quadrispinosa Shang and Zhang, and B. unicornis Shang and Li, are described from China. Figures are provided for these species, Bhatia koreana (Kwon and Lee), a new record for China, and B. biconjugara Zhang and Zhang. A key to separate these species and B. satsumensis Matsumura from China is given. Key Words: species, China The paraboloponine leafhopper genus Bhatia Distant (Hemiptera, Cicadellidae, Selenocephalinae), described from a mis- identified type species (Webb 1994), was established by Distant (1908) and revised by Zhang and Webb (1996). Formerly, the genus contained nine species from Asia and the Pacific, of which B. satsumensis Matsumura and B. biconju- gara Zhang and Zhang were known from China. The record of B. olivacea (Me- lichar) from China (Kuoh 1966, Zhang and Zhang 1998) cannot be confirmed as the identification was based on a female specimen. In addition, it should be noted that the figures 142 and 143 in Kuoh’s (1966) paper are switched, fig. 143 shows the general appearance of Bhatia olivacea (Melichar) and fig.142 is for Phlogotettix cyclops (Mulsant and Rey). The present work describes and figures five new Bhatia species from China and B. koreana Kwon and Lee is a newly recorded species for China. A feature of the genus, Hemiptera, Cicadellidae, Selenocephalinae, Bhatia, new record, new the paraphyses of the aedeagus, is not present in one of the included species and also the degree of attachment to the aedeagus of the paraphyses varies consid- erably between species. Both situations also occur 1n some Cicadellinae genera, which, based on other features in these genera, 1s irrespective of any close relation- ship between the species (Young 1968: 13). Type specimens of the new species are kept in the collection of the Entomologi- cal Museum, Northwest A&F University (NWAFU), Institute of Zoology, Chi- nese Academy of Science (IZCAS), and China Agricultural University (CAU). Bhatia Distant 1908 Bhatia Distant 1908:357; Zhang and Webb 1996:12; Viraktamath 1998:170. Type species: Eutettix? olivacea Meli- char. By original designation. Melichariella Matsumura 1914:236—237; Ishihara 1954:243-245; Linnavuori 1960: 566 36; Linnavouri and Al-Ne’amy 1983: 23. Synonymy revived by Zhang and Webb 1996. Type species: Melichar- iella satsumensis matsumura. By orig- inal designation. Koreanopsis Kwon and Lee _ 1979:50 Synonymised by Zhang and Webb 1996. Type species: Koreanopsis kor- eana Kwon and Lee. By monotypy. Diagnosis.—Head slightly wider than pronotum; vertex short and wide, more than twice as broad as long, slightly longer medially than next to eyes, anterior margin rounded to face, with a transverse depression subapically; ocel- li on foremargin; antennae long. Prono- tum finely transversely striate. Forewing elongate with 4 apical cells and 3 sub- apical cells. Foretibia rounded dorsally, setal formula 1+4. Hind femur with apical setal formula 2+2+1. Male genita- lia with pygofer lobe internal ledge (if present) extended diagonally across lobe to its ventroposterior corner; with several macrosetae posteriorly. Subgenital plate triangular with inner side nearly straight. Connective Y- or H-shaped. Aedeagus with one or more basal processes or paraphyses (except flabellata). Distribution.—Widely distributed throughout the Oriental Region and Pacific areas. Remarks.—The genus is similar to Athysanopsis Matsumura but differs from the latter as follows: pygofer lobe internal ledge (if present) extended diagonally across lobe to its ventroposterior corner rather than along posterior margin; sub- genital plate with inner side nearly straight rather than with inner apical half constricted. Bhatia is separated from other genera of Paraboloponini in the key to genera by Zhang and Webb (1996). KEY TO SPECIES (MALES) OF CHINA 1. Aedeagus without basal processes or pa- FApMySesi (hig 55)) ener B. flabellate, n. sp. Aedeagus with at least one basal process... 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON N Aedeagus with a (Figs. 49, 50) — Aedeagus with one or two pairs of basal processes 3. Aedeagus with two pairs of short apical processes (Figs. 10—12) single basal process B. unicornis, n. sp. B. satsumensis — Aedeagus without apical process ....... 4 Aedeagus with a single pair of basal pro- GESSES.2 Agha ht oo. coe = Be SAR eee 5 — Aedeagus with two pairs of basal processes. . . 7 5. Aedeagal shaft short and robust, connected to paraphyses by membrane (Fig. 31) B. digitata, n. sp. connected to — Aedeagal shaft elongate, paraphyses directly 6. Apical process of style short and hooklike (Fig. 16) — Apical process of style long and straight (Fig. 20) B. hastate, n. sp. 7. Apical process of style with apical exten- VKopm (WEE SM) 5 os bid oo boo g' — Apical process of style without apical extension (Fig. 44)... .B. quadrispinosa, n. sp. B. koreana B. biconjugara Bhatia biconjugara Zhang and Zhang (Figs. 1—6) Bhatia biconjugara Zhang and Zhang 1998: 178, figs 1A—E. Distribution.— China (Sichuan, Guangxi). Bhatia satsumensis (Matsumura) (Figs. 7—12) Melichariella satsumensis Matsumura 1914: 237-238; Ishihara 1954: 242-243. Bhatia satsumensis: Zhang and Webb 1996:12, figs 128-133; Zhang and Zhang 1998: 179. Distribution.—Japan, China (Guang- dong). Bhatia koreana (Kwon and Lee) (Figs. 13-17) Koreanopsis koreana Kwon and Lee 1979:50, figs. 1—5. Bhatia koreana: Zhang and Webb 1996: 12, figs 139-144, 490. Specimens examined.—1 ¢ (IZCAS), Huoditang, Ningshan County, Shaanxi VOLUME 108, NUMBER 3 567 iS J i) la s) 6 Figs. 1-6. Bhatia biconjugara (from Zhang and Zhang 1998). 1, Pygofer in lateral view. 2, Subgenital plate, valve, style and connective in dorsal view. 3, Style in dorsal view. 4, Aedeagus in lateral view. 5, Connective in ventral view. 6, Aedeagus in posterior view. province, 1,580 m, 18.vii. 1998, coll. dark maculation and yellow spots; fore- Yuan Decheng. New record for China. wing brownish hyaline with dark patch Distribution. South Korea, China _ at tip of clavus; legs yellowish. (Shaanxi). . Head with ocelli visible dorsally, ap- proximately 3 times own diameter from corresponding eye. Face with laterofron- : tal suture extended to lateral margin of new species _ 4 ocellus; antennal ledge strong. Pronotum (Figs. 18-23) with lateral margins moderately long. Male.—Length (incl. tegmen): 8.0 mm. Forewing with appendix narrow. Yellowish brown, head and _ thorax Male genitalia with pygofer side long, marked dorsally with brown as in_ triangular; lobe without internal ledge. Fig. 18; face deep yellow, pronotum with Valve pentagonal. Subgenital plate elon- a yellowish band anteriorly, with dense gate, triangular shaped. Style apical Bhatia hastata Shang and Shen, 568 Figs. 7-12. Connective in ventral view. 9, Style in ventral view. 10, Aedeagus in lateral view. 11, Aedeagus in posterior view. 12, Aedeagus in ventral view. process very long and straight, slightly tapered to truncate apex, without lateral lobe. Connective H-shaped. Aedeagus with shaft narrow, apex slightly expand- ed caplike in dorsal/ventral view; pre- atrium long with a pair of elongate basal processes lateroposteriorly, strongly curved dorsally and exceeding apex of shaft. Holotype (IZCAS), Mt. Sheng- tang, Jinxiu, Guangxi Province, 900- 19,00 m, 28.v1.2000, coll. Yao Jian. Etymology.—This species is named for its spearlike aedeagal shaft in lateral view. Notes.—This species is similar to B. koreana but can be distinguished by: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Bhatia satsumensis (from Zhang and Webb 1996). 7, Genital segment in lateral view. 8, 1) pygofer long and triangular; 2) style with apical process very long, straight and slightly tapered apically, without a lateral lobe; 3) valve pentagonal; and 4) aedeagal shaft slightly expanded api- cally in dorsal/ventral view. Bhatia digitata Shang and Shen, new species (Figs. 24-31) Male.—Length (incl. tegmen): 6.8 ~ 7.5mm. Yellowish brown; head and thorax dorsally marked with brown as in Fig. 24; face brown with base of frontoclypus yellow; forewing — light brownish hyaline, with spot at tip of clavus; legs brown. VOLUME 108, NUMBER 3 569 Figs. 13-23. 13-17, Bhatia koreana. 13, Pygofer in lateral view. 14, Valve and subgenital plate in 15, Aedeagus and connective in lateral view. 16, Connective and style in ventral view. 17, 18-23. B. hastata. 18, Head and thorax in dorsal view. 19, Pygofer in lateral view. 20, Valve, subgenital plate and style in ventral view. 21, Aedeagus and connective in lateral view. 22, ventral view. Aedeagus in dorsal view. Style in ventral view. 23, Aedeagus in dorsal view. 570 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON GETS rer Carr, SSO SSS Figs. 24-38. Valve and subgenital plate in ventral view. 27, Style in ventral view. 28, Connective in ventral view. on oy. Fatiba ree ‘to LS = iS me (San. 24-31, Bhatia digitata. 24, Head and thorax in dorsal view. 25, Pygofer in lateral view. 26, 29 Aedeagus in dorsal view. 30, Apex of aedeagus posteriorly. 31, Aedeagus and connective in lateral view. 32-38, B. flabellate. 32, Head and thorax in dorsal view. 33, Pygofer in lateral view. 34 Valve, subgenital plate and style in ventral view. 35, Aedeagus and connective in lateral view. 36, Apex of aedeagus posteriorly. 37, Connective in ventral view. 38, Female segment in ventral view. Head with ocelli visible dorsally, ap- proximately twice own diameter from corresponding eye. Face with laterofron- tal suture extended to corresponding ocellus; antennal ledge strong. Pronotum with lateral margin very short. Forewing with appendix narrow. Male genitalia with pygofer side elon- gate, upturned apically; lobe without internal ledge. Valve trapezoid. Subgeni- tal plate triangularly shaped, distal half digitate. Style apical process very short, tapered to rostriform apex, lateral lobe slight. Connective H-shaped. Aedeagus VOLUME 108, NUMBER 3 with shaft very short and robust, globu- lar at apex in dorsal/ventral view with many fine spines; a pair of very long and narrow processes basoposteriorly —be- tween aedeagus and connective, linked to former by an elongate membranous band. Types.—Holotype: ¢ (IZCAS), Mt. Shengtang, Jinxiu, Guangxi Province, 900 m, 18.v.1999, coll. Yao Jian, Paratype: 12 (NWAFU), Baotianman, Neixiang, Henan Province, 1,300 m,_ 11.vu.1998, coll. Hu Jian. Etymology.—This species is named for its fingerlike aedeagal shaft. Notes.—This species is similar to B. koreana but differs by: 1) pygofer more elongate; 2) apical process of style shorter; and 3) aedeagal shaft short and robust, with many dense fine spines and basal processes linked to aedeagus by a mem- brane. Bhatia flabellata Shang and Shen, new species (Figs. 32—38) Male——CenethvGnel) tegmen): 75° — 8.0 mm. Female.—Length (incl. tegmen): 8.2 ~ 8.8 mm. Yellowish brown; head and thorax dorsally marked with brown as in Fig. 32; frontoclypus with a row of short transverse brown bands along each side; pronotum with dense yellow spots; fore- wing light brownish hyaline. Head with ocelli visible dorsally, ap- proximately twice own diameter from corresponding eye. Face with antennal ledge strong; laterofrontal suture extend- ed to corresponding ocellus. Pronotum with sides moderately long. Forewing with appendix broad. Male genitalia with pygofer side broad basally, dorsal margin sloping caudally to narrow, bluntly rounded apex; lobe without internal ledge. Valve pentagonal. Subgenital plate triangularly shaped with 571 digitate apex; fine setae along lateral margin. Style with apical process short, rostriform, lateral lobe prominent. Con- nective H-shaped. Aedeagal shaft elon- gate, strongly curved dorsally, expanded flabellate at apex, without processes. Types.—Holotype: ¢ (IZCAS), Linhai Villa, Jinxiu, Guangxi Province, 1,000 m, 2.vii.2000, coll. Li Wenzhu; Paratypes: 1 ¢ | 2 (IZCAS), Mt. Shengtang, Jinxiu, Guangxi Province, 900—-1,900 m, 28.vi1.2000, coll. Yao Jian; 1 2 (IZCAS), coll. Chen Jun, other data same as holotype. Etymology.—This species is named for the expanded flabellate apex of the aedeagus. Notes.—This species is allied to B. hastata but can be distinguished by: 1) pygofer bluntly rounded apically; 2) aedeagal shaft strongly curved dorsal- ly, with expanded flabellate apex; and 3) style with apical process rostriform. Bhatia quadrispinosa Shang and Zhang, new species (Figs. 39-45) Male.—Length (incl. tegmen): 6.5 mm. Yellow; head and _ thorax dorsally marked with brown as in Fig. 39; fore- wing light brownish hyaline with small brown spot at tip of clavus; legs yellow. Head with ocelli not visible dorsally, approximately twice own diameter from corresponding eye. Face with laterofron- tal suture not reaching corresponding ocellus; antennal ledge weak. Pronotum with lateral margin very short. Forewing with appendix narrow. Female pregenital segment with hind margin concave me- dially, midlength longer than that of preceding segment; ovipositor extending beyond apex of abdomen. Male genitalia with pygofer side broad at base, dorsal margin abruptly sloping subapically to triangular-shaped apex; lobe without internal ledge. Valve pen- tagonal. Subgenital plate triangularly shaped with digitate apex, with many Figs. 39—S0. fine setae on lateral margin. Style with apical process rostriform. Connective Y- shaped, stem nearly as long as arms. Aedeagal shaft short and robust, strong- ly curved dorsally, slightly expanded apically in lateral view, with a pair of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 39-45, Bhatia quadrispinosa. 39, Head and thorax in dorsal view. 40, Pygofer in lateral view. 41, Subgenital plate in ventral view. 42, Valve in ventral view. 43, Aedeagus in lateral view. 44, Style and connective in ventral view. 45, Aedeagus in dorsal view. 46—50, B. unicornis. 46, Head and thorax in dorsal view. 47, Pygofer in lateral view. 48, Valve, subgenital plate, connective and style in ventral view. 49, Aedeagus and connective in lateral view. 50, Aedeagus in ventral view. short basal processes and a pair of long lateroposterior processes, former not exceeding apex of shaft and latter longer than shaft. Types. gyinge, Holotype: nd a(CAW); Mt.Emei, Sichuan Qin- Province, VOLUME 108, NUMBER 3 20.vili. 1961, coll. Jin Ruihua. Paratype: | 2 (CAU), same data as holotype. Etymology.—This species is named for its four aedeagal processes. Notes.—This species is similar to B. biconjugara but differs in: 1) pygofer side broad basally with dorsal margin abrupt- ly sloping subapically to a triangular- shaped apex; 2) valve pentagonal; and 3) style with apical process hook-shaped. Bhatia unicornis Shang and Li, new species (Figs. 46—S0) Male.—Length (incl. tegmen): 6.8 ~ 7.5 mm. Sordid yellow; head and thorax dorsally marked with brown as_ in Fig. 46; forewing hyaline with a spot at tip of clavus. Head with ocelli visible dorsally, ap- proximately twice own diameter from corresponding eye. Face with laterofron- tal suture extending to corresponding ocellus; antennal ledge weak. Pronotum with sides very short. Forewing with appendix narrow. Male genitalia with pygofer side broad basally, dorsal margin abruptly sloping subapically to broadly rounded apex; lobe without internal ledge. Valve pen- tagonal. Subgenital plate triangularly shaped, tapered to digitate apex, with fine setae on lateral margin. Style with apical process moderately long and truncate apically. Connective Y-shaped, stem longer than arms. Aedeagus with shaft moderately long and robust, curved dorsally, a laterally serrate triangularly shaped flange subapically on each side; a long, stout medial process baso- posteriorly between aedeagus and con- nective, linked to former by membrane. Types.—Holotype ¢ (NWAFU), Mt. Dinghu, Guangdong Province, 18.vii.1985, coll. Zhang Yalin. Paratypes: 1 4, 1 & (NWAFU), same data as holotype. Etymology.—This species is named for its digitate subgenital plate. Notes.—This species is allied to B. flabellata but can be distinguished by: 1) style with apical process moderately long with apical margin truncate; 2) con- nective Y-shaped with stem longer; and 3) aedeagus with shaft moderately long and robust with a laterally serrate tri- angular-shaped flange subapically on each side and with a single process baso-poster- iorly linked to aedeagus by membrane. ACKNOWLEDGMENTS This study is supported by the National Science Foundation of China (Grant No. 30370180), the Ministry of Educa- tion, China (Grant No. 20030712015), and the China Postdoctoral Science Foundation (Grant No. 2004035509). We give our sincere thanks to M. D. Webb (The Natural History Museum, London) for revising the manuscript and also to Yang Chikun, Li Fasheng, and Cai Wanzhi (The China Agricultural University, Being) and Yang Xingke (The Institute of Zoology, Chinese Acad- emy of Sciences, Beijing) for arranging the loan of specimens. LITERATURE CITED Distant, W. L. 1908. Rhynchota-Homoptera. The Fauna of British India Including Ceylon and Burma, IV. Published Under the Authority of the Secretary of State for India in Council. Edited by Lt. Col. C. T. Bingham, 501, 282 figs. Ishihara, T. 1954. Revision of two Japanese genera of the Deltocephalinae (Insecta: Hemiptera). Zoological Magazine, Tokyo 63: 243-245. figs. 1-2. Kuoh, C. L. 1966. Economic Insect Fauna of China (Homoptera: Cicadellidae). Science Press, Beijing. 170 pp. Kwon, Y. J. and C. E. Lee. 1979. Some new genera and species of Cicadellidae of Korea (Homo- ptera: Auchenorrhyncha) nature and life in Southeast Asia. Kyungpook Journal of Bi- ological Sciences 9: 49-61. Linnavuori, R. 1960. Cicadellidae (Homoptera, Auchenorrhyncha) of Fiji. Acta Entomologica Fennica 15: 1-71. Linnavuori, R. E. and K. T. Al-Ne’amy. 1983. Revision of the African Cicadellidae (subfami- ly Selenocephalinae) (Homoptera, Auchenor- rhyncha). Acta Zoologica Fennica 168: 1—105. 574 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Matsumura, S. 1914. Die jassinen und einige neue Acocephalinen japans. Journal of the College of Agriculture, Imperial University of Tokyo 5: 165-240. figs 1-12. Viraktamath, C. A. 1998. Revision of the leafhop- per tribe Paraboloponini (Hemiptera: Cicadel- lidae: Selenocephalinae) in the Indian sub- continent. Bulletin of The Natural History Museum (Entomology) 67(2): 1—207. Webb, M. D. 1994. Bhatia Distant, 1908 (Insecta, Homoptera): proposed confirmation of Eutettix? olivaceus Melichar, 1903 as the type species. Bul- letin of Zoological Nomenclature 51: 116—117. Young, D. A. 1968. Taxonomic Study of the Cicadellinae (Homoptera: Cicadellidae). Part 1: 13. Smithsonian Institution Press, Washing- ton, D.C. 287 pp. Zhang, Y. L. and M. D. Webb. 1996. A revised classification of the Asian and Pacific Seleno- cephalinae Leafhoppers (Homoptera: Cicadel- lidae). Bulletin of The Natural History Muse- um (Entomology) 65(1): 1—103. Zhang, W. Z. and Y. L. Zhang. 1998. On Chinese species of the genus Bhatia Distant (Homo- ptera: Cicadellidae). Entomotaxonomia 20(3): 177-181. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 575-582 THE BIOLOGY OF CHIONODES HIBISCELLA (BUSCK) (LEPIDOPTERA: GELECHIIDAE), WITH DESCRIPTIONS OF THE IMMATURE STAGES JENNIFER L. APPLE AND DAviID ADAMSKI (JLA) Department of Biology, Willamette University, 900 State Street, Salem, OR 97301, U.S.A. (e-mail: japple@willamette.edu); (DA) Department of Entomology, Natural Museum of Natural History, P.O. Box 37012, Smithsonian Institution, Washington, D.C. 20013-7012 (dadamski@sel.barc.usda.gov) Abstract.—Observations are presented on the life history of Chionodes hibiscella (Busck) (Lepidoptera: Gelechiidae) on Hibiscus moscheutos L. (Malvaceae) in nontidal freshwater wetlands and intertidal habitats along the Chesapeake Bay in Maryland. In the summer, early instar larvae of this bivoltine species are leaf-tiers in young terminal leaves; later instars construct leaf rolls from mature leaves. In the fall, larvae from the second brood feed on the seeds of mature fruits and overwinter in the dried fruit capsule. The morphology of the larva and pupa is illustrated with images from digital day-light photography, scanning electron microscopy, and _ line drawings. Chaetotaxy of the larva, mouthparts and associated sensilla, claws of the thoracic legs, prolegs of the abdomen, and the anal fork are described. Key Words: Gelechiidae, Hibiscus, larval behavior, Malvaceae, morphology, pupa Species of Chionodes Hubner (Gele- chiidae) are found throughout the Hol- arctic and Neotropical regions. Howev- er, most of the nearly 200 species are known from North America north of Mexico. Host plant data are available for 79 species, but little is known about the complete life histories and feeding habits of most species (Hodges 1999). We describe the immature stages of Chio- nodes hibiscella (Busck) and add to the existing knowledge of its natural history and life cycle. Chionodes hibiscella is related to 65 other species recognized as the /ugubrella group (Hodges 1999). Males of the group share a hindwing with slender sex scales on the dorsal, basomedial surface; a vin- culum that is narrow to broadly U- shaped and a lobe arising from the posterolateral surface; and sternum 8 with a medial rectangular extension, the surface of which has a sclerotized, often obliquely lined zone anteromesally. Fe- males share an antrum lacking a poster- iorly directed extension of the ventral wall; apophysis anterioris arising dorsal- ly; and usually with a sclerotized rim on tergum 8 with an extremely short free part. Larvae in the /ugubre/la group have been reared from several plant families including Asteraceae, Betulaceae, Clado- ceraceae, Fabaceae, Malvaceae, Rham- naceae, Rosaceae, Selaginellaceae, and Sterculiaceae. Within the /ugubrella group, Hodges (1999) recognized 11 species complexes. The hibiscella complex contains six species, one of which 1s Chionodes hibiscella. Members of this complex are characterized by the following: valva highly sclerotized and less than half the 576 length of the tegumen; saccular lobe reduced and/or represented by a patch of setae; tegumen with anterior margin deeply emarginate; vinculum short; un- cus with posterior margin smooth; ster- num 8 with anteromesial margin tuning- fork shaped; an aedeagus lacking cor- nuti; ductus bursae short; and corpus bursae with two zones of sclerotization. All Nearctic species of the hibiscella com- plex are suspected or confirmed herbi- vores of Malvaceae. Of these species, C. hibiscella has received the most study and is known to feed on Hibiscus L. and Kosteletzkya Presl. (Hodges 1999). MATERIALS AND METHODS Observations on the natural history of Chinodes hibiscella (Fig. 1) were made as part of a broader study of herbivory and seed predation of Hibiscus moscheutos L. (Malvaceae) (Figs. 2—3) at the Smithso- nian Environmental Research Center (SERC), located along the Rhode River subestuary of the Chesapeake Bay (38°S3"N, 76°33'W) near Edgewater, Maryland, USA. Plants of H. moscheu- tos were monitored in 14—22 sites across three different habitat types: nontidal freshwater wetlands, intertidal habitats with frequent flooding and seasonally varying salinity (Fig. 2), and high marsh intertidal habitats with higher salinity and less frequent flooding. Marked plants were examined for herbivore damage in June of 2000— 2002. Larval specimens of Chionodes hibiscella were collected from leaves of Hibiscus moscheutos during June 2002. All larvae were either boiled in H5O and preserved in 70% EtOH, retained until pupation for preservation and study of the pupa, or maintained in the laborato- ry for adult emergence. During the period from August through October, fruits of Hibiscus moscheutos were collected as they ma- tured and dehisced on marked shoots. Fruits were stored individually in Zi- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ploc® bags, glassine envelopes, or small plastic cups and later dissected and examined for evidence of seed predators. Live or dead larvae found in fruits were preserved in 70% EtOH. For SEM study, larvae and pupae were cleaned in 10% alcohol with a camel’s hair brush and subsequently dehydrated in increasing concentrations of alcohol to absolute alcohol. After dehydration, specimens were critical point dried using a Tousimis critical point dryer, mounted on SEM stubs, and coated with gold-palladium (40/60%), using a Cressington sputter coater. The ultrastructure of the larva and pupa was studied with an Amray 1810 scanning electron microscope at an accelerating voltage of 10 kV. Gross morphological observations and measurements of the larva and pupa were made using a dissecting microscope (reflected light) with a calibrated mi- crometer. The Methuen Handbook of Colour (Kornerup and Wanscher 1978) was used as a color standard. All voucher specimens from this study are deposited in National Museum of Nat- ural History, Smithsonian Institution, Washington, DC. BIOLOGY The feeding behavior of Chionodes hibiscella is easily distinguished from that of other Hibiscus herbivores. Larvae or evidence of their presence were found in all three Hibiscus habitat types at SERC. Observations of C. hibiscella activity on Hibiscus moscheutos shoots in mid- to late-June were usually of leaves rolled from one or both edges that housed the larva as it fed (Figs. 6— 7). Our observations generally agree with those of Busck (1903), Heinrich (1921), Weiss and Dickerson (1919), and Ca- hoon and Stevenson (1986): the larva cuts a small notch in the petiole (Fig. 5) that causes the leaf to hang, but not to drop. Weiss and Dickerson (1919) sug- gested that partially severing the petiole VOLUME 108, NUMBER 3 Si Figs. 1-7. Adult, habitat, host, larva, and larval behavior of Chionodes hibiscella. 1, Adult. 2, Wetland habitat. 3, Flower of Hibiscus moscheutos. 4, Petiole of Hibiscus leaf, showing notch cut by larva. 5, Leaf margin rolled by larva. 6, Leaf roll constructed by larva, dangling from partially severed petiole. 7, Larva repairing a previously disturbed leaf roll. in this manner allows the larva to roll the wither and either remain attached to the leaf more easily as it wilts slowly and plant (Fig. 4) or drop to the ground. becomes more flexible. The larva even- June observations also included early tually pupates in these leaf rolls, which instar larvae, which typically were found 578 in the cluster of young leaves at the shoot tip (Fig. 8). Sometimes larvae pulled the upper surfaces of two small young leaves (~2 cm2 leaf area) together with silk and fed in the region between them. Larger young leaves occasionally were folded lengthwise along the midrib. Larvae also occurred in silken shelters constructed between a mature leaf and a dry young leaf that had been partially eaten at the base by younger instars. Observations of larvae at SERC are consistent with claims that Chionodes hibiscella is bivoltine. Busck (1903) col- lected adults in May and August in the District of Columbia, and Weiss and Dickerson (1919) reported adults flying in early June and late July and into August in New Jersey, indicating two flight periods a year for C. hibiscella. At SERC, mature larvae collected from Hibiscus leaves in early June 2002 pupated several weeks later. In Septem- ber and October of 2000 and 2001, late instar larvae were found in silken tubes constructed inside dry Hibiscus fruits. Signs of Chionodes’ presence included large holes in the fruit wall near the base, frass inside the fruit, and damaged seeds. Larvae also were encountered in silken shelters between the calyx and the fruit, with no signs of fruit damage. Cahoon and Stevenson (1986) reported similar patterns in herbivory in HAlibiscus moschuetos in a marsh along the Chop- tank River on Maryland’s eastern shore, with damage to leaves occurring from early June to late July and infestation of fruits noted in October. Larvae feeding on fruits in the fall at SERC are likely offspring of the gener- ation observed feeding on leaves in June. Weiss and Dickerson (1919) reported that the eggs laid by adults emerging from the early summer brood hatch by the end of August and feed at the shoot tip and around the inside of the fruit. They observed in New Jersey that larvae are mature by late October, and over- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON winter in this stage. At SERC, mature larvae could still be found in dried fruit capsules on standing dead shoots in mid- November 2002. Often the peduncle of the fruit was hollowed out by the larva and it becomes a common site for pupation (Fig. 10). Though little exami- nation of Hibiscus was conducted in winter and early spring at SERC, we have encountered pupae of Chionodes hibiscella in Hibiscus in mid-April. DESCRIPTIONS OF IMMATURES Larva (Figs. 7, 9, 11—21).—Length 10.0-14.1 mm (n = 10). Body smooth, pale gray or off white with paired dorsal, subdorsal, and lateral, irregularly shaped, longitudinal, grayish-red stripes from T3—A10 (Figs. 7, 9, 18-19, 21); T1— T2 grayish red (grayish-red pigmentation fades rapidly in specimens preserved in EtOH); sclerites of thoracic legs darkly pigmented, claw juxtaposed to a flat- tened, downcurved seta (Fig. 15); all pinacula brown centrally, gradually be- coming paler to outer margin; anal shield pale brownish yellow with small, darker pigmented spots on surface; Tl with shield dark brown along posterolateral margin, gradually becoming paler to anterior margin; head capsule dark brown except for 2 large, dorsolateral, amber spots on epicranium from outer margin of adfrontal sclerite to postero- lateral area of the epicranial notch, gradually darkening to the genal area. Head (Figs. 11-12): Hypognathous; ad- frontal sclerites narrow, gradually wid- ening from lateral margins of clypeus to epicranial notch; paired AF2 near apex of frons, but not beyond, about equal in length with AF1l; AFl and AF2 slightly shorter than Fl; C2 and Cl about equal in length; Fl equidistant to C2 as C2 is to Cl; labrum with 6 setae, 2 mesial, 2 near middle of lateral lobe, and 2 near outer margin of lateral lobe; outer labral setae about equal in length, about twice length of mesial setae; other setae of VOLUME 108, NUMBER 3 Dy Figs. 8—10. head capsule are illustrated in Figs. 11— 12. Mandibles (Fig. 20) slightly asym- metrical, quadridentate, and with pair of subequal setae on outer surface; maxilla and antenna with sensilla types and arrangement as illustrated (Figs. 13—14); six stemmata in a C-shaped arrange- ment; stemmata III and IV slightly approximate. Prothorax (Fig. 18): SD1 and D2 about equal in length, longest setae on shield, XD1 slightly shorter; SD1 in straight line with XD2 and XD1 along anterior margin; SDI about 2.5 times longer than XD2; XD1 slightly shorter than D2; SD2, D2, and D1 in straight line; distance between XD2 and XD1 about twice distance between SD1 and XD2; SD2 and XD2 about equal in length, SD2 equidistant to SDI and XD2; D2 about 2.5—3.0 times longer than XD2, D2 equidistant to XD2 and Larva and pupa of Chionodes hibiscella. 8, Early instar among terminal leaves of host (see arrow). 9, Mature larva. 10, Pupation site within fruit. XD1; L-group pinaculum trisetose, L1 closer to: 12. than: to L3; and about 2 times longer than L2 or L3; L2 and L3 about equal in length; L1 slightly longer than SV1, SV1 about twice as long as SV2; coxae about '% distance apart as distance between coxae on T2—T3 (not illustrated); V1’s about 1/3 distance apart as distance between V1’s on T2—T3 (not illustrated). Meso- and metathorax (Fig. 18): D2 and D1 on separate pina- cula, D2 about 2.5—3.0 times longer than D1; SDI and SD2 on same pinaculum, SD1 about 2.5—3.0 times longer than SD2; D1, SD1, SD2, and D2 in straight; L2 about twice length of L1, L1 slightly longer than L3; SV1 slightly posteriorad to L3; SV1 slightly shorter than L2. Abdomen (Figs. 16-17, 19, 21): Al—A2 (Fig. 19), with D2 about 2 times longer than D1; SD1 and D2 about equal in 580 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 11-17. palpus. Scale = 10 um. 14, Antenna. Scale = 10 um. 15, Claw of thoracic leg, with adjacent flattened and downcurved seta. Scale = 10 um. 16, Abdominal proleg. Scale = 100 um. 17, Anal shield and anal fork. Scale = 100 um. size; SD1 in vertical line with D1; SD1 slightly posteriorad spiracle; L1 about 3 times longer than L2, slightly poster- iorad spiracle; L2 in line with or slightly anterior spiracle; L3 longer than L2, and in line with or slightly posteriorad to D2; SV-group bisetose on Al, SV1 about 2 times longer than SV2; SV-group trise- tose on A2, SV1 about twice lengths of SV3 and SV2. A3—A6 with SD1 slightly anteriorad spiracle; one tonofibrillary Larva of Chionodes hibiscella. 11-12, Chaetotaxy of head. Scale = 100 um. 13, Maxillary platelet dorsoposterior to SD1 and near ventral margin of subdorsal longitudinal stripe; SV-group trisetose on_ proleg; crochets biordinal except, uniordinal and weak laterally (Fig. 16); V1’s slightly farther apart than on Al—A2 and A7— A8. A7 (Fig. 21), with SV-group bise- tose, SV1 slightly longer than SV2. A8 (Fig. 21), with SV-group unisetose, SV1 slightly longer than L3; L3_ slightly posteriorad to L1, and in line with SV1. VOLUME 108, NUMBER 3 581 Figs. 18-21. Mandible. 21, Abdominal segments 6—10. A9 (Fig. 21), with D2 about twice length of D1, D1 slightly anteriorad to D2; SD1 hairlike, and in line with D1; SD1 about equal in length to D1; L1 about twice as long as L2, and in line with L3 and SV1. A10 with anal plate (Figs. 17, 21), with SD2, D1, and D2 about equal in lengths, SD1 slightly longer; D1 slightly ante- riorad to SD1, and in line with SD2; D2’s decumbent; anal fork with 2 pairs of sickle-shaped furcae, mesial pair about twice as long as lateral pair. Pupa (Figs. 10, 22-24).—Length: 4.0— 5.4 mm (n = 10). Body smooth; femur of foreleg exposed as a_ short narrow Chaetotaxy of larva of Chionodes hibiscella. 18, Thorax. 19, Abdominal segments 1-2. 20, sclerite; tip of hindleg exposed between shghtly divergent apical portion of an- tenna; terminal six abdominal segments pivoting as a unit. Remarks.—Larvae for two species of Chionodes, C. arenella (Forbes) and C. trichostola (Meyrick), have been illus- trated previously by Hodges (1999). Both Chionodes hibiscella and C. trichos- tola possess a similar anal comb and SD1 on AQ is hairlike, whereas in C. arenella, the anal comb is absent and SDI on A9Y is_ setiform. Chionodes hibiscella appears to differ from C. trichostola by having D1, SD2, and XD2 equal in 582 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 22-24. Pupa of Chionodes hibiscella. 22, Ventral view. 23, Dorsal view. 24, Lateral view. Line scale = | mm. lengths on the prothoracic shield, and L3 is near equal in length with L2 on the prothorax. ACKNOWLEDGMENTS We thank C. Feller, J. O'Neill, and D. Whigham of the Smithsonian Environ- mental Research Center, Edgewater, MD, for support and advice. Research by JLA was supported in part by a Smithsonian Institution Postdoctoral Fellowship. LITERATURE CITED Busck, A. 1903. A revision of the American moths of the family Gelechiidae, with descriptions of new species. Proceedings of the United States National Museum 25: 767-938. Cahoon, D. R. and J. C. Stevenson. 1986. Pro- duction, predation, and decomposition in a low-salinity Hibiscus marsh. Ecology 67: 1341-1350. Heinrich, C. 1921. Some Lepidoptera likely to be confused with the pink bollworm. Journal of Agricultural Research 20: 807-836. Hodges, R. W. 1999. Gelechioidea, Gelechiidae, Chionodes. In Dominick, R. B. et al., eds. The Moths of America North of Mexico, fasc. 7.6., 339 pp. Huemer, P. and K. Sattler. 1995. A taxonomic revision of Palearctic Chionodes (Lepidoptera: Gelechiidae). Beitrage zur Entomologie 45: 3-108. Kornerup, A. and J. H. Wanschner. 1978. Methuen Handbook of Colour. 2.4 ed. Methuen and Co., Ltd., London, 243 pp. Weiss, H. B. and E. L. Dickerson. 1919. Insects of the swamp rose-mallow, Hibiscus moscheutos L., in New Jersey. Journal of the New York Entomological Society 27: 39-68. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 583-610 ADVENTIVE APHIDS (HEMIPTERA: APHIDIDAE) OF AMERICA NORTH OF MEXICO RoBERT G. Foottirt, SUSAN E. HALBERT, GARY L. MILLER, ErR1Ic Maw, AND LoulseE M. RUSSELL! (RGF, EM) Agriculture and Agri-Food Canada, Canadian National Collection of Insects, Ottawa, ON, KIA O0C6 Canada (e-mail: foottitrg@agr.gc.ca; mawe@agr.gc.ca); (SEH) Florida Department of Agriculture and Consumer Services, Division of Plant Industry, Gainesville, FL 32614, U.S.A. (e-mail: halbers@doacs.state.fl.us); (GLM, LMR) Systematic Entomology Laboratory, Plant Science Institute, Agricultural Research Service, U.S. Department of Agriculture, Beltsville, MD 20705, U.S.A. (e-mail: gmiller@sel.barc.usda.gov) Abstract.—We provide a compilation of 262 species of aphids that are considered as adventive to North America north of Mexico. Included for each species, where applicable, is reference to: the location and date of introduction of the first North American record; pest status in North America; principal economic hosts; and biogeographical origin. Information is also provided for species whose presence in North America is considered erroneous or questionable and for those species that are considered Holarctic or Beringian. Key Words: Aphidoidea, plant lice, plant pathogens, invasive species, biological control, quarantine, agriculture, forestry, horticulture, ornamentals, fruit trees In an era of a global economy and_ reintroduced into the United States commerce, the increase of adventive species represents both domestic and international concerns. If non-indige- nous species become pests, the economic consequences include loss of production, diminished quality, production cost in- creases, flexibility decreases in produc- tion/management decisions, increased risk of human disease, and damaging environmental and aesthetic effects (Hu- ber et al. 2002). Various imsects of agricultural concern have been identified as potentially damaging if introduced or 1 Tt is noteworthy that Louise Russell, a pioneer woman entomologist and specialist in the Sternorrhyncha, continues to publish at the age of 101. (Huber et al. 2002). Many aphid species have been added to the North American fauna since colonization by Europeans, and some of these have become serious pests. In addition to direct feeding damage, aphids constitute one of the largest groups of known plant-virus vectors (Chan et al. 1991). Aphids are primarily controlled through pesticide applica- tions, adding to crop production costs, and having environmental implications associated with increased pesticide use. The effects of some non-indigenous aphids in North America have been staggering. For example, by the early 1990’s, the Russian wheat aphid, Diur- 584 aphis noxia (Kurdjumov), was responsi- ble for $500—900 million cumulative losses in the western U.S. (Bernal et al. 1993, Morrison and Peairs 1998). The appearance of the brown citrus aphid, Toxoptera citricida (Kirkaldy), in Flor- ida in 1995 had a major impact on the U.S. citrus industry (Halbert and Brown 1996). More recently, losses related to the soybean aphid, Aphis glycines Mat- sumura, have been estimated at about $120 million in 2003 for Minnesota alone (Anonymous 2005). The origins of this work actually began in part in the 1970’s when one of us (LMR) contributed data on invasive species in the United States (e.g., Sailer 1978). Nearly 30 years later, adventive insects are still a major concern for agriculture in North America. The ob- jectives of this paper are to: 1) list the adventive aphid species (excluding adel- gids and phylloxerans) of North America and record data on first detection date (along with validation source), principal hosts, and zoogeographic area of origin; 2) analyze these data and determine if patterns exist; and 3) compare these data to other sternorrhynchan groups. MATERIALS AND METHODS We use the term adventive species as defined by D.R. Miller et al. (2002), 1.e., for those species that are non-native to North America regardless of economic harm. Other terms such as invasive, alien, exotic, and immigrant have some- times been used interchangeably (see Wheeler and Hoebeke (2001) and Miller et al. (2005) for discussion). Using Smith and Parron’s (1978) list of North Amer- ican aphids as the base, we compiled a new list for North American north of Mexico. We exclude species that have been intercepted at ports-of-entry or discovered in nurseries or greenhouses and subsequently eradicated. With the exception of Aphis chloris Koch, a bio- control agent for Hypericum perforatum PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Harris 2003), no aphid species has been purposely introduced. Geographic origins are given in gen- eral terms only, relying heavily on Black- man and Eastop (1994, 2000), supple- mented by other references, as noted in Table 1, and by opinion of the authors. The origins of a few cosmopolitan species are unknown. It is not always readily apparent whether a species is adventive or native; some species we have listed as adventive may in fact be naturally Holarctic in distribution and vice versa. For determining the earliest record of the North American adventive aphids, we relied on published records and on specimens examined in The Canadian National Collection of Insects, Nema- todes and Arachnids (CNC) and the Edith Patch Collection (CNC-Patch) housed at CNC, Ottawa, ON, Canada; the MacGillivray Collection (ACNB), Agriculture and Agri-Food Canada, Fredericton, NB, Canada; The Florida State Collection of Arthropods (FSCA), Gainesville, FL, U.S.A.; and The Na- tional Entomological Collection of the National Museum of Natural History (USNM), Beltsville, MD, U.S.A. In addition, Asa Fitch’s aphid species notes that are currently housed with the USNM aphid collection at Beltsville, MD, were also examined. This collection of written records and observations from the 1840’s—1870’s provide some of our earliest glimpses into the North Ameri- can aphid fauna. While there are some questions concerning Fitch’s “‘species concept’”” versus current thinking for some of his included species, there is little confusion given the morphology and associated host plants for others. His reference to the occurrence in 1791 of the cabbage aphid, Brevicoryne brassicae (L.) is possibly the earliest record of an adventive aphid for North America. Nomenclature followed herein reflects that of Remaudiére and Remaudiere VOLUME 108, NUMBER 3 585 Table 1. Introduced aphid species in Canada and continental United States of America, giving earliest record with source of record, geographic origin, pest status, and economic host species. Aphid names follow Remaudiere and Remaudiére (1997). Earliest records are based on literature references and on specimens in the following collections: USNM — National Museum of Natural History; CNC — Canadian National Collection of Insects, Arachnids and Nematodes; FSCA — Florida State Collection of Arthropods; ACNB — MacGillivray collection, Agriculture and Agri-Food Canada, Fredericton, NB; CNC-Patch — Edith Patch collection, housed at CNC; EMEC — Essig Museum of Entomology Collection, University of California, Berkeley. Introduced Species Earliest N.A. record (1) (Collection or Reference) Geographic Origin/ Reference (2) Status in North America/ eference (2) North American Economic Host(s) Acyrthosiphon caraganae SK, 1940 (CNC) (Cholodkovsky) Acyrthosiphon kondoi Shinji Acyrthosiphon lactucae (Passerini) Acyrthosiphon malvae (Mosley) (3) Acyrthosiphon pisum (Harris) Aloephagus myersi Essig Amphorophora ampullata Buckton Anoecia corni (Fabricius) Aphidounguis mali Takahashi (4) Aphis amaranthi Holman Aphis callunae Theobald Aphis chloris Koch Aphis craccae Linnaeus Aphis craccivora Koch (5) Aphis cytisorum Hartig Aphis epilobii Kaltenbach Aphis euphorbiae Kaltenbach (6) Aphis fabae Scopoli (7) Aphis genistae Scopoli Aphis glycines Matsumura Aphis gossypii Glover CA, 1974 (Kono 1977) MD, 1943 (USNM) NE, 1891 (USNM) KS, 1877 (Harper et al. 1978) CA, 1939 (Essig 1950) MN, 1887 (Oestland 1887) IA, 1897 (USNM) NERISS9T(EN©@) FL, 1990 (Halbert et al. 2000) BC, 1974 (Forbes and Chan 1976) BC, 1980 (Harris 2003) BC, NB, ME, 1965 (CNC, ACNB) NE, 1890 (USNM) DC, 1906 (USNM) CO, 1915 (USNM) CA, 1933 (USNM) NY, 1869 (Fitch 1870 [as A. rumicis L.]) MA, 1921 (USNM) VACHE ING Ke Yes Mile MN, MO, OH, WI, WV, 2000 (Voegtlin 2000) NY, 1852 (Russell 1968) Palearctic Palearctic (e Asia) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Afrotropical Palearctic (Europe) Palearctic (Europe) Palearctic (e Asia) Neotropical (Cuba) Palearctic (Europe) Palearctic (Europe) Palearctic Palearctic Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Oriental and Palearctic (East Asia) Palearctic pest pest Halbert et al. 2000 pest minor pest Essig 1950, regulatory issue potential pest Halbert et al. 2000 Released as weed biological control agent pest Caragana arborescens Medicago sativa Lactuca sativa Geranium, Pelargonium Leguminosae, incl. Piswm, Medicago Aloe (greenhouse) roots of Gramineae, incl. cereals edible Amaranthus Hypericum perforatum Vicia cracca polyphagous, with preference for legumes Cytisus scoparius polyphagous major pest Ragsdale Glycine max et al. 2004 major pest polyphagous 586 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Introduced Species Earliest N.A. record (1) (Collection or Reference) Geographic Origin/ Reference (2) Status in North America/ North American Economic Host(s) Aphis hederae Kaltenbach Aphis idaei van der Goot Aphis nasturtii Kaltenbach Aphis nepetae Kaltenbach Aphis nerii Boyer de Fonscolombe Aphis polygonata (Nevsky) (8) Aphis pomi De Geer Aphis praeterita Walker Aphis rumicis Linnaeus (7) Aphis salicariae Koch Aphis sambuci Linnaeus Aphis sedi Kaltenbach Aphis spiraecola Patch Aphis taraxicola (Borner) (9) Aphis thalictri Koch Aphis umbrella (Borner) Aphis urticata Gmelin Aphis vaccini (Borner) Aploneura lentisci (Passerin1) Asiphonella dactylonii Theobald Aspidaphis adjuvans (Walker) Atarsaphis agrifoliae (Ferris) Atheroides serrulatus Haliday Aulacorthum circumflexum (Buckton) Aulacorthum solani (Kaltenbach) Betulaphis brevipilosa Borner CA, 1910 (Essig 1910) BC, 1951 (Forbes et al. 1973) NE, 1894 (USNM) ID, 1957 (Gittins et al. 1976) IL, 1879 (Thomas 1879) IL, 1977 (FSCA) PA, 1844 (Haldeman 1844) ME, 1918 (Patch 1918) Unknown (7) ON, 1952 (CNC) NY, 1847 (Fitch notes [as A. sambucifoliae Fitch]) ME, 1911 (Patch 1911) CO, 1907 (Gillette 1910) QC, 1983 (Cloutier et al. 1986) BC, 1999 (CNC) NE, SD, 1891 (USNM [as 4. maliae Koch]) PA, 1913 (Olsen 1921) BC, 2000 (CNC) IL, 1879 (Thomas 1879 [as Rhizobius poae Thomas]) AZ, 1947 (USNM) CO, 1907 (Gillette 1917b) CA, 1921 (Ferris 1921) QC, 1966 (CNC) CO, 1907 (Gillette 1908) NY, 1879 (Riley and Monell 1879) BC, 1959 (Forbes et al. 1974) Palearctic (Europe) minor pest Palearctic (Europe) Palearctic pest Palearctic Palearctic (e Asia) pest Palearctic (Central Asia) Palearctic (Europe) pest Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic or Oriental Palearctic Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Mediterranean) Palearctic (e Asia) potential pest Palearctic (Europe) Palearctic (e Asia) Palearctic Palearctic pest (e Asia) or Oriental Palearctic Palearctic (Europe) major pest minor pest major pest Hedera helix Solanum tuberosum ornamentals (flowers) Malus pumila, Pyrus communis, Cydonia oblonga polyphagous Gramineae, including Triticum Cynodon dactylon Gramineae, incl. cereals polyphagous polyphagous VOLUME 108, NUMBER 3 Table 1. Continued. 587 Introduced Species Earliest N.A. record (1) (Collection or Reference) Geographic Origin/ Reference (2) Status in North America/ Reference (2) North American Economic Host(s) Betulaphis helvetica Hille BC, 1926 Ris Lambers Betulaphis quadrituberculata (Kaltenbach) Brachycaudus helichrysi (Kaltenbach) Brachycaudus cardui (Linnaeus) Brachycaudus persicae (Passerini) Brachycaudus rumexicolens (Patch) (10) Brachycaudus schwartzi (Borner) Brachycaudus spiraeae (Borner) Brachycaudus tragopogonis (Kaltenbach) Brachycorynella asparagi (Mordvilko) Brevicoryne brassicae (Linnaeus) Calaphis betulicola (Kaltenbach) Callipterinella calliptera (Hartig) Callipterinella minutissima (Stroyan) Capitophorus carduinus (Walker) Capitophorus elaeagni (Del Guercio) Capitophorus hippophaes (Walker) Capitophorus pakansus Hottes and Frison Capitophorus horni Borner Cavariella aegopodii (Scopoli) Cavariella archangelicae (Scopoli) Cavariella pastinacae (Linnaeus) Cerataphis brasiliensis (Hempel) Cerataphis lataniae (Boisduval) Cerataphis orchidearum (Westwood) (Glendenning 1926) BC, 1946 (ACNB) OR, 1893 (USNM) NY, 1887 (USNM) MD, DE, NJ, VA, MI, 1887 (Smith 1890) CT 1913 (Patch 1917) CA, 1975 (Eastop 1975 ) ON, 1956 (CNC) WA, 1993 (FSCA) NY, NJ, 1969 (Leonard 1970, Race 1970) NY, 1791 (Fitch notes) NY, 1854 (Fitch notes) OR, 1909 (USNM) BC, 1975 (Forbes and Chan 1980) GA, 1948 (USNM) SD, 1892 (USNM) DC, 1895 (USNM) IL, 1929 (Hottes and Frison 1931) ME, 1951 (Corpuz- Raros and Cook 1974) MN, 1903 (USNM) WA, 1953 (USNM) NY, 1855 (Fitch notes) OH, 1917 (USNM) NE, 1912 (USNM) DC, 1923 (USNM) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic Palearctic Palearctic (Europe) Palearctic Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe, Iceland) Palearctic (Europe) Oriental Oriental Oriental major pest pest pest locally serious pest Stoetzel 1990 pest minor pest Hajek Prunus, Compositae Prunus, Compositae Prunus Prunus persica Asparagus officinalis Brassica crops Betula and Dahlsten 1986 minor pest Hajek Betula and Dahlsten 1987 potential weed biocontrol agent Gassmann and Kok 2002 Potential virus vector pest pest pest pest Cirsum Daucus carota palms palms greenhouse orchids Nn oo co Table 1. Continued. Introduced Species Ceruraphis eriophori (Walker) Chaetosiphon potentillae (Walker) Chaetosiphon tetrarhodum (Walker) Chaitophorus leucomelas Koch Chaitophorus populialbae (Boyer de Fonscolombe) Chromaphis juglandicola (Kaltenbach) Cinara cedri Mimeur Cinara confinis (Koch) Cinara costata (Zetterstedt) Cinara cupressi (Buckton) Cinara fresai Blanchard Cinara juniperi (De Geer) Cinara laricis (Hartig) Cinara pilicornis (Hartig) Cinara pinea (Mordvilko) Cinara pruinosa (Hartig) Cinara tujafilina (Del Guercio) Coloradoa abrotani (Koch) Coloradoa absinthii (Lichtenstein) Coloradoa achilleae Hille Ris Lambers Coloradoa artemisiae (Del Guercio) Coloradoa rufomaculata (Wilson) Coloradoa tanacetina (Walker) Corylobium avellanae (Schrank) Cryptaphis poae (Hardy) Cryptomyzus galeopsidis (Kaltenbach) Earliest N.A. record (1) (Collection or Reference) UT, 1935 (Knowlton and Smith 1936) NY, 1851 (Fitch notes) CO, 1916 (USNM) MN, 1907 (USNM) CA, QC, 1966 (USNM, Quednau 1966) ON, 1955 (CNC) CA, 1896 (USNM) NY 2004 (CNC, FSCA) CO, 1920 (USNM) ON, 1905 (USNM) CA, 1916 (Bartholomew 1932) FL, 1942 (Tissot 1944 [as C. wacasassae}) NY, 1902 (USNM) MA, 1921 (USNM) ME, 1906 (Patch 1912) DC, 1919 (Wilson 1919) CO, 1910 (Gillette 1917a [as Lachnus palmerae}) CA, 1914 (Davidson 1914) NB, 1964 (ACNB) MB, BC, 1965 (CNC) ON, 1962 (CNC) BC, 1965 (CNC) NE, 1890 (USNM) RI, 1974 (Smith and Parron 1978) BC, 1983 (Forbes and Chan 1984) MB, 1965 (Robinson and Bradley 1965) ME, 1914 (Patch 1914) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON North American Economic Host(s) Geographic Origin/ Status in North America/ Reference (2) i Reference (2) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) pest Olsen 1974 Juglans regia Palearctic (Europe) Cupressus, Juniperus Palearctic Abies Palearctic Picea Palearctic Cupressus, Juniperus unknown Cupressus, Juniperus Palearctic (Europe) Juniperus Larix Picea Palearctic Palearctic (Europe) Palearctic Pinus Palearctic Picea unknown Palearctic (Central Asia) Palearctic (Central Asia) Palearctic (Central Asia) Palearctic (Central Asia) Palearctic (Central Asia) Palearctic (Central Asia) Palearctic (Europe) pest Achillea minor pest chrysanthemums Corylus avellana Palearctic (Europe) Palearctic (Europe) VOLUME 108, NUMBER 3 Tablet Continued. Introduced Species Earliest N.A. record (1) (Collection or Reference) Geographic Origin/ Reference (2) Status in North America/ Reference (2) North American Economic Host(s) Cryptomyzus ribis (Linnaeus) Ctenocallis setosus (Kaltenbach) Ctenocallis zelkowae (Shinji) Decorosiphon corynothrix Borner Diuraphis frequens Walker Diuraphis noxia (Kurdjumoy) Drepanosiphum oregonensis Granovsky Drepanosiphum platanoides (Schrank) Dysaphis apiifolia (Theobald) Dysaphis crataegi (Kaltenbach) (11) Dysaphis devecta (Walker) Dysaphis foeniculus’ (Theobald) (12) Dysaphis plantaginea (Passerini) Dysaphis pyri (Boyer de Fonscolombe) Dysaphis radicola (Mordvilko) Dysaphis tulipae (Boyer de Fonscolombe) Elatobium abietinum (Walker) Eriosoma grossulariae (Schule) (13) Eriosoma pyricola Baker & Davidson Eriosoma ulmi (Linnaeus) (13) Eucallipterus tiliae Schouteden Eucarazzia elegans (Ferrari) Euceraphis betulae (Koch) (14) Euceraphis punctipennis (Zetterstedt) (14) Eulachnus agilis (Kaltenbach) Eulachnus brevipilosus Borner Eulachnus rileyi (Williams) NY, 1854 (Fitch notes) OR, 1951 (Essig 1952) CA, 1983 (Kono 1983) NC, 1963 (Smith and Knowlton 1975) ON, 1951 (CNC) TX, 1986 (Stoetzel 1987) OR, 1914 (USNM) CA, 1908 (Wilson 1909) CA, 1934 (USNM) DC, 1915 (USNM) QC, 1966 (Quednau 1966) OR, 1949 (USNM) NE, 1890 (Williams 1911) CO, 1978 (Smith and Parron 1978) DC, 1894 (USNM) NH, 1903 (USNM) CAS DEFI9I5 (USNM) ON, 1913 (CNC) OH, 1897 (Baker and Davidson 1916) BC, 2003 (CNC) DC, 1890 (USNM) CA, 1984 (Stoetzel 1985) CO, 1909 (USNM) NY, 1847 (Fitch notes) NY, 1909 (Gillette 1909) WA, 1954 (Hottes and Essig 1955) IN, 1886 (USNM) Palearctic (Europe) Palearctic (Europe) Palearctic (E Asia) Palearctic (Europe) Palearctic (Europe) Palearctic (Central Asia) Palearctic Palearctic (Europe) Palearctic Palearctic Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) pest Pest Kono, 1983 minor pest Halbert et al. 1992 major pest Quisenberry and Peairs 1998 pest pest occasional pest pest, regulatory concern pest pest minor pest Ribes Zelkova Triticum Gramineae, incl. cereals Picea Acer Umbelliferae Umbelliferae Malus pumila Umbelliferae (including Daucus carota) Malus pumila Pyrus communis bulbs Abies Ribes, Ulmus Pyrus communis Ulmus; Ribes Tilia Labiatae Betula Betula Pinus Pinus Pinus 590 Table 1. Introduced Species Continued. Earliest N.A. record (1) (Collection or Reference) Geographic Origin/ Reference (2) Status in North America/ Reference (2) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON North American Economic Host(s) Euthoracaphis umbellariae Essig Forda formicaria yon Heyden Forda marginata Koch Geoica setulosa (Passerini) Geoica utricularia (Passerini) (15) Geopemphigus floccosus (Moreira) Glyphina betulae (Linnaeus) Greenidea ficicola Takahashi Greenidea psidii van der Goot Hayhurstia atriplicis (Linnaeus) Hyadaphis coriandri (Das) Hyadaphis foeniculi (Passerini) Hyadaphis tartaricae (Ajzenberg) Hyalopteroides humilis (Walker) Hyalopterus amygdali (Blanchard) (16) Hyalopterus pruni (Geoffroy) (16) Hyperomyzus carduellinus (Theobald) Hyperomyzus lactucae ( Linnaeus ) Hyperomyzus pallidus Hille Ris Lambers Hyperomyzus picridis (Borner) Hyperomyzus rhinanthi (Schouteden) (17) Idiopterus nephrelepidis Davis Jacksonia papillata Theobald Liosomaphis berberidis (Kaltenbach) Lipamyzodes matthiolae (Doncaster) CA, 1929 (Essig 1932) CO, 1909 (USNM) DC, 1885 (USNM) ID, 1985 (FSCA) IL, 1887 (USNM) FL, 1945 (USNM) PA, 1945 (USNM) FL, 2002 (Halbert 2004) CA, 1998 (Gill 1998a) CA, 1887 (USNM) FL, 1997 (Halbert et al. 2000) NY, 1879 (USNM) IL, 1979 (Voegtlin 1981) DC, MA, 1909 (Hayhurst 1909) IL, NJ (date ?) (16) (Smith and Parron 1978) IL, 1879 (Thomas 1879) FL, 1997 (Halbert et al. 2000) IL, 1878 (Thomas 1879) ME, 1914 (Hille Ris Lambers 1949) PA, 1942 (USNM) ME, 1951 (CNC) IL, 1907 (Davis 1909) ID, 1959 (USNM) NY, 1846 (Fitch notes) WA, 1976 (Fox 1977) Palearctic (e Asia) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Neotropical Palearctic (Europe) Oriental Oriental Palearctic (Europe) Palearctic (Central Asia) Palearctic (Central Asia) Palearctic (Central Asia) Palearctic Palearctic (e Asia) Palearctic (Europe) Palearctic (e Asia) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Neotropical Stoetzel 1990 Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) minor pest G.L. Miller et al. 2002 minor pest potential pest minor pest, regulatory pest major pest minor pest major pest Voegtlin 1981 pest pest minor pest potential pest pest Stoetzel 1990 roots of Gramineae roots of Gramineae roots of Gramineae roots of Gramineae, incl. cereals roots of Ipomoea batatas Ficus Myrtus Chenopodium including Ch. quinoa Umbelliferae, including Coriandrum, Foeniculum Lonicera ; Umbelliferae Lonicera tatarica Prunus (s.g. Amydalus) Prunus (S.g. Prunus) Ribes Ribes Ribes Ribes greenhouse ferns VOLUME 108, NUMBER 3 Table 1. Continued. 59] Introduced Species Earliest N.A. record (1) (Collection or Reference) Geographic Origin/ Reference (2) Status in North America/ Reference (2) North American Economic Host(s) Lipaphis pseudobrassicae (Davis) (18) Longicaudus trirhodus (Walker) Macrosiphoniella abrotani (Walker) Macrosiphoniella absinthii (Linnaeus) Macrosiphoniella artemisiae (Boyer de Fonscolombe) Macrosiphoniella grandicauda Takahashi and Moritsu Macrosiphoniella leucanthemi (Ferrari) Macrosiphoniella millefolii (De Geer) Macrosiphoniella sanborni (Gillette) Macrosiphoniella subterranea (Koch) Macrosiphoniella tanacetaria (Kaltenbach) Macrosiphum Iilii (Monell) (19) Macrosiphum rosae (Linnaeus) Macrosiphum stellariae Theobald Melanaphis bambusae Fullaway Melanaphis sacchari (Zehntner) Melanaphis sorini Halbert and Remaudiere Metopolophium dirhodum (Walker) Metopolophium festucae (Theobald) Microlophium carnosum (Buckton) (20) Microlophium sibiricum (Mordvilko) (20) DC, 1899 (USNM) MA, 1908 (USNM) IL, 1929 (USNM) IL, 1879 (Thomas 1879) NB, 1950 (ACNB) MD, 1997 (USNM) PA, 1948 (USNM) Cie 1957 (EMEC) NE, 1891 (USNM) PA, 1948 (USNM) CT, 1909 (CNC-Patch) NY, 1878 (Comstock 1880) MA, 1841 (Harris 1841) BC, 1980 (Forbes and Chan 1983) GA, 1914 (USNM) BS Lie (BS EA) FL via CA, 1996 (Halbert and Remaudiere 2000) UT, 1910 (USNM) CA, 1970 (USNM, Eastop 1971) NE, 1889 (Williams 1911) PA, 1963 (USNM, Pepper 1965) Palearctic (Europe) major pest Paddock Cruciferae Palearctic (Europe) Palearctic Palearctic Palearctic Palearctic Palearctic Palearctic Palearctic (e Asia) Palearctic Palearctic Palearctic? Palearctic (Europe) Palearctic Palearctic (e Asia) Oriental Oriental Halbert and Remaudiére 2000 Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) Palearctic (Europe) 1915 pest minor pest pest pest pest pest White et al. 2001 pest Halbert and Remaudiere 2000 major pest pest Rosa Leucanthemum vulgare Asteraceae (esp. chrysanthe- mums) cultivated chrysanthe- mums cultivated chrysanthe- mums (occa- sionally) Cultivated lilies Rosaceae (esp. Rosa) Caryophyllaceae (esp. Dianthus) bamboos Gramineae (esp. sugar- cane) Miscanthus Gramineae, incl. cereals; Rosa Gramineae Urtica Urtica 592 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Earliest N.A. record (1) Geographic Origin/ Status m North America/ North American Introduced Species (Collection or Reference) Reference (2) Reference (2) Economic Host(s) Mindarus abietinum IL 1879 (Thomas 1879 Palearctic (Europe) Abies Koch (21) [as Schizoneura pinicola]) Monaphis antennata ON, 1996 (CNC) Palearctic (Europe) Betula (Kaltenbach) Muscaphis musci Borner NC, 1962 (Smith and Palearctic (Europe) Knowlton 1975) Myzaphis bucktoni ME, 1959 (USNM) Palearctic (Europe) Rosa Jacob Myzaphis rosarum NY, 1906 (USNM) Palearctic (Europe) pest Rosa (Kaltenbach) Myzocallis carpini BC, 1964 (Forbes Palearctic (Europe) Carpinus (Koch) et al. 1973) betulus Myzocallis castanicola CA, 1914 (USNM) Palearctic (Europe) pest Castanea and Baker Quercus Myzocallis coryli CA, 1903 (Clarke Palearctic (Europe) pest Corylus (Goeze) 1903) avellana Myzus antirrhini MD, 1996 (USNM) Palearctic (Europe) polyphagous (Macchiati) Myzus ascalonicus NB, 1954 unknown Blackman pest polyphagous Doncaster (MacGillivray and Eastop 2000 1954b) Myzus cerasi Passerini NY 1855 (Thomas Palearctic (Europe) pest Prunus (s.g. 1879) Cerasus) Myzus certus (Walker) NB, 1952 (MacGil- Palearctic (Europe) minor pest Caryophyllaceae, livray 1954a) incl. Dianthus Myzus hemerocallis Ps AS I9SS Palearctic (e Asia) pest Hemerocallis (Schrank) (USNM) Myzus ligustri (Mosley) NY, 1915 (USNM) Palearctic (Europe) occasionally severe Ligustrum sp.; pest Mason 1940 Euonymus sp. Myzus lythri (Schrank) DC, 1916 (USNM) Palearctic (Europe) potential weed Lythrum biocontrol agent salicariae Voegtlin 1995b Myzus ornatus Laing CA, 1936 (Essig Palearctic pest polyphagous 1938) Myzus persicae (Sulzer) MA, 1841 (Harris Palearctic (e Asia) major pest polyphagous 1841) Myzus varians Davidson CA 1911 (Davidson Palearctic (e Asia) pest Clematis 1912) Nasonovia ribisnigri IL, 1879 (Thomas Palearctic (Europe) major pest; Lactuca sativa (Mosley) 1879) regulatory concern Neophyllaphis araucariae FL, 1963 (Woodruff Australasian pest Russell 1967 = Araucaria Takahashi 1963) (Australia) heterophylla Neophyllaphis podocarpi CA, 1954 (Russell Australasian pest Russell 1967, © Podocarpus Takahashi 1967) (Australia) Denmark 1969 Neotoxoptera formosana CA, 1927 (Essig Oriental pest Allium (Takahashi) 1935) Neotoxoptera oliveri CA, 1927 (Essig Oriental pest Allium (Essig) 1935) Ovatus crataegarius OR, 1893 (USNM) Palearctic (Europe) pest Mentha ; (Walker) Crataegus Paracolopha morrisoni MD 1916 (Baker Palearctic (e Asia) bamboos (Baker) 1919) Panaphis juglandis OR, 1928 (USNM) Palearctic (e Asia) pest Juglans regia (Goeze) VOLUME 108, NUMBER 3 Table 1. Continued. Introduced Species Pemphigus bursarius (Linnaeus) Pemphigus spyrothecae Passerini Pentalonia nigronervosa Coquerel Periphyllus californiensis (Shinji) Periphyllus lyropictus (Kessler) Periphyllus testudinaceus (Fernie) Phloeomyzus passerini (Signoret) (22) Phorodon humuli (Schrank) Phyllaphis fagi (Linnaeus) Pleotrichophorus chrysanthemi (Theobald) Pleotrichophorus glandulosus (Kaltenbach) Prociphilus xylostei (De Geer) (23) Pseudacaudella rubida (Borner) Pseudoregma panicola (Takahashi) Pterocallis alni (De Geer) Pterocomma pilosum Buckton Pterocomma populeum (Kaltenbach) Pterocomma salicis (Linnaeus) Rhopalomyzus lonicerae (Siebold) Rhopalosiphoninus hydrangeae (Matsumura) Rhopalosiphoninus latysiphon (Davidson) Earliest N.A. record (1) (Collection or Reference) NY, 1872 (Fitch 1872) QC, 1972 (Alleyne and Morrison 1974) DC, 1909 (USNM) WA, 1903 (Essig and Abernathy 1952) VT, 1847 (Fitch notes [as Aphis aceris L.]) BC, 1924 (Essig and Abernathy 1952) NB? 1950, ME 1970 (Smith 1974a [as P. dearborni Smith]) NY, 1863 (Walsh 1863) IL, 1879 (Thomas 1879) CA, 1910 (USNM) CO, 1915 (USNM) ME, 1914 (Patch 1917) NC, TN, 1973 (Smith and Knowlton 1975) FL, 1928 (FSAC) IL, 1910 (Davis 1910) UT, 1958 (Knowlton 1958) MT, 1904 (USNM) MN, 1886 (Oestlund 1886) VA, 1912 (USNM) BC, 1975 (Forbes and Chan 1976) CA, 1912 (USNM) Geographic Origin/ Reference (2) Palearctic (Europe) Palearctic (Europe) Unknown (either Neotropical or Oriental) Palearctic (e Asia) Essig and Abernathy 1952 Palearctic (Europe) Palearctic (Europe) Palearctic Palearctic (Europe) Palearctic (Europe) Palearctic (Asia?) Palearctic (Europe) Palearctic Palearctic (Europe) Oriental Palearctic (Europe) Palearctic Palearctic (Europe) Palearctic Palearctic (Europe) Palearctic (e Asia) Palearctic Status in North America/ Reference (2) pest pest pest pest minor pest minor pest minor pest North American Economic Host(s) Populus nigra; Lactuca sativa Populus nigra Musa (banana); ornamental Musaceae, Araceae, Zingiberaceae Acer Acer platanoides Acer Prunus: Humutlus Fagus chrysanthemums Ambrosia, Artemisia Lonicera; roots of Picea Lonicera Bulbs; Solanum tuberosum 594 Tablewk= Introduced Species Continued. Earliest N.A. record (1) (Collection or Reference) Geographic Origin/ Reference (2) Status in North America/ Reference (2) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON North American Economic Host(s) Rhopalosiphoninus staphyleae (Koch) Rhopalosiphum maidis (Fitch) (24) Rhopalosiphum nymphaeae (Linnaeus) Rhopalosiphum rufiabdominale (Sasaki) (24) Schizaphis graminum (Rondani) Schizaphis minuta (van der Goot) Schizaphis palustris (Theobald) Schizaphis rotundiventris (Signoret) Schizolachnus pineti (Fabricius) Shivaphis celti Das (25) Sipha elegans Del Guercio Sipha glyceriae (Kaltenbach) Sitobion avenae (Fabricius) Sitobion fragariae (Walker) Sitobion luteum (Buckton) (26) Smynthurodes betae Westwood Subacyrthosiphon cryptobium Hille Ris Lambers Takecallis arundicolens (Clarke) Takecallis arundinariae (Essig) Takecallis taiwanus (Takahashi) Tetraneura fusiformis Matsumura (27) Tetraneura ulmi (Linnaeus) Thecabius lysimachiae Borner (28) BC, 1953 (ACNB) NY, 1851 (Fitch 1856) NE, 1890 (USNM) NY, 1856 (Fitch 1856) MD, 1884 (Pergande 1902) FL, 1998 (Halbert et al. 2001) BC, 1974 (Forbes and Chan 1976) FL, 1988 (Halbert et al. 2000) BC, 1961 (CNC) GA, SC, 1996 (Halbert et al. 2000) NB, 1949 (ACNB) ME, 1909 (Patch 1910) NY, 1851 (Davis 1914) CA 1916 (USNM) FL, 1940 (FSCA) DC, 1895 (USNM) IA, 1966 (USNM) CA, 1903 (USNM) CA, 1911 (Essig 1917) FL, 1930 (USNM) FL, 1959 (ESCA, USNM) NE, 1890 (USNM) NY, 2000 (FSCA) Palearctic (Europe) Palearctic (e Asia) Palearctic Davidson 1917 Palearctic (e Asia) Palearctic Oriental Palearctic (Europe) Palearctic (e Asia) or Oriental Palearctic Palearctic (E Asia) and Oriental Palearctic Black- man & Eastop 2000 Palearctic Palearctic Palearctic (Europe) unknown Palearctic (Mediterranean) Palearctic (Europe) Oriental Oriental Oriental Palearctic (e Asia) / Oriental Palearctic Palearctic (Europe) minor pest serious pest pest/ biological control pest serious pest nuisance pest Halbert and Choate 1999 minor pest pest major pest Davis 1914 pest minor pest minor pest Stoetzel et al. 1996 possible pest Polyphagous on roots, including stored bulbs and rootcrops Gramineae various aquatic plants Gramineae Gramineae Celtis Gramineae Gramineae (incl. rice) Gramineae Gramineae, Rubus greenhouse orchids polyphagous on roots incl. Phaseolus, Solanum, Gossypium bamboos bamboos bamboos Ulmus?; roots of Gramineae inc. rice Populus, Lysimachia VOLUME 108, NUMBER 3 Table 1. Continued. nN \o Nn —_—_—___-.:.O0.::_.0 °c Introduced Species Earliest N.A. record (1) (Collection or Reference) Geographic Origin/ Reference (2) Status in North America/ eference (2) North American Economic Host(s) ’ << ) Hoplia clorophana Erichson, 1848 LG? 5 Hoplia coerulea (Drury, 1773) Ge 5 Gymnoloma femorata Burmeister, 1844 kG: 5 Hoplocnemis mutica Burmeister, 1844 Ge 5 Pachycnema calcarata Burmeister, 1844 ERGs 5 Pachycnema squamosa Burmeister, 1844 Ge 5 Rutelinae Rutelini Pelidnota virescens Burmeister, 1844 OG: 7. Anomalini Anomala cincta Say, 1835 OO: G: 7 Strigoderma sulcipennis Burmeister, 1844 ORG: 7 Epectinaspis mexicana (Burmeister, 1844) OVG: i Callirhinus metallescens Blanchard, 1850 O. G. 7 Dynastinae Dynastini Golofa imperialis Thomson, 1858 OG: 6 Pentodontini Tomarus sallei (Bates, 1888) O. G. 6 Orizabus clunalis (Leconte, 1856) ONG: 6 Cyclocephalini Cyclocephala lunulata Burmeister, 1847 ONG: 6 Trichiinae Trichiini Trigonopeltastes geometrica Schaum, 1841 OG; 5) Cetoniinae Gymnetini Cotinis mutabilis (Gory & Percheron, 1833) OG: 5 Goliathini ° Neoscelis dohrni (Westwood, 1855) ONG; 5 Cetoniini Euphoria basalis (Gory & Percheron, 1833) ORG: 5 Cremastocheilini Cremastocheilus knochi LeConte, 1853 ORG: =) placed inside Macrodactylini, is found Melolonthinae, Dynastinae, and Ruteli- independent and is more basal inside the nae. They all share one synapomorphy: clade that maintains the relationships of anterior edge of prementum sinuate. VOLUME 108, NUMBER 3 Table 2. Characters states of taxa used in the phylogenetic analysis. Species 11111111112222222222333333333344444444445555 12345678901234567890 1234567890 1234567890 1234567890123 Aegidium cribratum Melolontha melolontha Phyllophaga obsoleta Diplotaxis hirsuta Macrodactylus mexicanus Liogenys fuscus Ceraspis pilatei Tsonychus ocellatus Barybas aurita Clavipalpus basalis Pelidnota virescens Strigoderma sulcipennis Epectinaspis mexicana Anomala cincta Callirhinus metallescens Cyclocephala lunulata Orizabus isodonoides Tomarus sallei Golofa imperialis Neoscelis dorhni Cremastocheilus knochi Euphoria basalis Cotinis mutabilis Trigonopeltastes geometrica Hoplia festiva Hoplia asperula Hoplia clorophana Hoplia coerulea Hoplia trivialis Hoplia dispar Hoplia surata Gymnoloma femorata Pachycnema calcarata Pachycnema squamosa Hoplocnemis mutica Hoplia pollinosa 01110101010113101110111010111020111010011000011111100 01110001010113121000011010101020110010110010111110011 01110001010113101100011010101020100011110010111110001 01110001000113101100011010101020100011110010011111001 21110001001113121000011000110000110111000110011111001 01110101000113101100011000111020110011110010011111001 211101013021131200001 11000001020110111010010011100101 01110001001113101000011000110000110111010110011110101 01110001012113100001111010010020110111110110000111101 01110000001113101100111000110021110010110010010111101 00110100000113011100111011111120110011110010011011111 00110000000113001000111011111120110011110110011101101 001100000001 13001000111010111120110011110110011111001 001100000001 13001100111010111120110011110110011011101 001100000001 13001000111010111120110111110210011101101 011100000001 13101100111010111121110111110210010110001 01111101000113111100111010111101110110111210011111011 01111103000113111100111010111120110111110210010011101 01111103000113111100111010111121110111110210011111011 00001114100001101000111001111011011012101300001111101 01000210110000101000111000111021110010111200001111111 01000211101001101100111001111011011012101300010111101 00001214200001101000111001111011011012101302001111101 010000120001 13121000111000111011011011101100001111101 011101000121131001021001000110201 101111 10111011111001 01110100012113100102100100011020110111110111011111001 01110200012113100102100100011020110110110111011111001 01110200012113100102100100011020110110110111011111001 01110100003113100002100100011020110111110111011111001 01110000012113100002100100011020110111110111011111001 01110000012113100102100100011020110111110111011111001 01010200003102120101100100011010110111111211011111001 11010103003013100111100100011021121101111311011111001 11010103003013100111100100011021121101111311011111001 01010103013013000111100100111021121101101201011111111 01110200013113100002100100011020110111110111011111001 The clade Macrodactylini-Melolonthini (Fig. 1) is supported by two synapomor- phies: both mesotarsal claws cleft and presence of antennal sensilla of type K. The relationships among Diplotaxis hir- suta Vaurie and the clade containing Phyllophaga obsoleta (Blanchard) and Melolontha melolontha (Linne) (Fig. 1) is supported by a bootstrap value of 55% and two synapomorphies: procoxae transverse and sternites fused along mid- line. Finally the relationship between Phyllophaga obsoleta and Melolontha melolontha is supported by a bootstrap value of 68% and three synapomorphies: pronotum with lateral margin crenulate, antennal sensilla of type J present, and the sclerites associated with the spiculum gastrale absent. The clade of Dynastinae and Ruteli- nae (Fig. 1) is supported by three syna- pomorphies: clypeal apex entire, pro- coxae transverse, and the mandibles exposed dorsally. The Dynastinae clade is supported by a bootstrap value of 60% and one synapomorphy: distal 624 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Aegidium cribratum Trigonopeltastes geometrica 91 Cremastocheilus knochi 80 Euphoria basalis 91 Neoscelis dorhni Cotinis mutabilis Clavipalpus basalis Liogenys fuscus Diplotaxis hirsuta 55 68 Melolontha melolontha Phyllophaga obsoleta Aa Cyclocephala lunulata 63 Tomarus sallei Orizabus isodonoides Golofa imperialis Pelidnota virescens 56 Anomala cincta Epectinaspis mexicana Strigoderma sulcipennis Callirhinus metallescens 72 Macrodactylus mexicanus Isonychus ocellatus Ceraspis pilatei Barybas aurita Hoplia festiva 86 Hoplia asperula Hoplia trivialis Hoplia dispar Hoplia surata Hoplia pollinosa 65 Hoplia clorophana Hoplia coerulea Gymnoloma femorata 80 96 Hoplocnemis mutica 87 Pachycnema calcarata Pachycnema squamosa Fig. 1. Strict consensus tree of 30 equally parsimonius trees (TL = 178, CI = 0.410, RI = 0.730). Only bootstrap values over 50 % are shown. Four diagnostic characters for the hopliids are shown: one metatarsal claw; widely retractile metatarsal claws; metatarsal claws without oniquia; and basis of spiculum gastrale widened. VOLUME 108, NUMBER 3 edge of labrum curved. The clade of Tomarus sallei (Bates), Orizabus isodo- noides (Leconte) and Golofa imperialis Thomson is supported by four synapo- morphies: head projections present, fron- toclypeal suture present only at sides, clypeal apex dentate; and pygidium semicircular. The clade of Orizabus iso- donoides and Golofa imperialis is sup- ported by one synapomorphy: sensilla of type L absent. he clade of Rutelinae (Fig. 1) is supported by a bootstrap value of 56% and two synapomorphies: tegumentary diffraction present and base of epipleura thickened. The relationship between Pe- lidnota virescens Burmeister and Anom- ala cincta Say is supported by the presence of antennal sensilla of type G. This last clade is the sister group of Epectinaspis mexicana (Burmeister), Stri- goderma sulcipennis Burmeister, and Callirhinus metallescens Blanchard. The relationship of Strigoderma mexicana and Callirhinus metallescens is supported by the presence of antennal sensilla of type H. The clade Dynastinae-Rutelinae is the sister group of the clade of the assembly of macrodactylids and all the representatives of hopliids. In one clade two species of macrodactylids are found: Macrodactylus =mexicanus Burmeister and Jsonychus ocellatus Burmeister. Their relationship is supported by a boot- strap value of 72% and three synapo- morphies: both mesotarsal claws cleft, fifth sternite two times longer than fourth sternite, and distal part of the left mandible shortened. The relationships of the clade containing two species of macrodactylids (Ceraspis pilatei Harold and Barybas aurita Bates) and the clade of the hopliids are supported by two synapomorphies: pygidium covered by scales and abdominal surface covered with scales. Barybas aurita appears as the sister taxon of the hopliids, supported by one synapomorphy: lateral margin of the pronotum crenulate. 625 The clade of the hopliids (Fig. 1) is supported by a bootstrap value of 86% and seven synapomorphies: clypeal apex entire, protibiae with three well-defined teeth, antennal sensilla of type K present, one metatarsal claw, widely retractile metatarsal claws, metatarsal claws with- out onychia, and base of spiculum gastrale widened. The clade of Hoplia clorophana Erichson and Hoplia coerulea (Drury) is supported by a _ bootstrap value of 65% sharing two synapomor- phies: loss of frontoclypeal suture and prementum with anterior edge entire. South African taxa are placed in another clade, supported by a bootstrap value of 80% and five synapomorphies: body depressed, elytra covered with setae and scales, metatibiae with one apical spur, labrum with anterior edge thin, and labrum with anterior edge entire. The node of Hoplocnemis and Pachycnema 1s supported by a bootstrap value of 96% and six synapomorphies: clypeus with apex dentate, mesepimeron exposed dor- sally, metatibiae thickened, galea_ with- out teeth, molar surface of left mandible without teeth or ridges, and prementum with anterior edge membranous. The clade formed by the two species of Pachycnema is supported by a bootstrap value of 87% sharing two synapomor- phies: anterior edge of labrum notched and body shape quadrate. Our analysis demonstrated that some traditional groups of the subfamily Melolonthinae are not monophyletic. Representatives of such groups (Lio- genys, Clavipalpus, Macrodactylus, Cer- aspis) are distributed in different lineages inside the tree (Fig. 2). But the species of Macrodactylus and Isonychus, as repre- sentatives of Macrodactylini (sensu stricto), are located near the species of Ceraspis and Barybas (Macrodactylini sensu lato), and are the closest relatives of the hopliuds. This phylogenetic hypothesis supports hopliines as a subfamily of Scarabaeidae 626 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Orphininae Trichiinae Cetoniinae Clavipalpus Liogenys Diplotaxini Melolonthini Dynastinae Rutelinae Macrodactylini Macrodactylini Ceraspis Barybas Hopliinae Figs 2. and corroborate (sensu Lawrence and Newton 1995) with the same rank of Melolonthinae, Rute- linae, and Dynastinae. Future detailed analysis based on more taxa of hopliines macrodactylines 1s the iéxistences of three tribes inside Hopliinae. Based on the present analysis we confirm that: Melolonthinae “Melolonthinae” Cladogram of the relations of the hopliids with the subfamilies of Melolonthidae. necessary to two or VOLUME 108, NUMBER 3 a) hopliines constitute a monophyletic group; and b) it is a group independent from Melolonthinae and Rutelinae and closely related to some macrodactylines. ACKNOWLEDGMENTS We thank CANACOL Foundation, Bruce Gill (Canada Department of Agriculture), Henry Howden and Fran- cois Genier (Canadian Museum of Na- ture) for their valuable aid and support while studying the collections in Ottawa. Mary Liz Jameson and Brett C. Ratcliffe (UNSM) are acknowledged for their friendly help and support for studying collections conserved in Nebraska, as well as for the loan of types of the species of Hoplia described by H. W. Bates that were deposited in London (NHML). Tiburcio Laez Aponte (Instituto de Ecologia, Xalapa) took the scanning electron microscope pictures. We thank Alejandro Espinosa de los Monteros (Instituto de Ecologia, Xalapa), Javier Garcia (Instituto de Ecologia, Xalapa) and Juan José Morrone (Facultad de Ciencias, UNAM, Mexico City) for their review and contributions to this paper. We are grateful to the anonymous reviewers and to the editor for their helpful suggestions on the manuscript. HCR was supported by a scholarship #157789 (CONACYT). This paper is a contribution to the project “Coleop- teros Lamelicornios de America Latina”’ supported by Instituto de Ecologia (ac- count 902-08-011). LITERATURE CITED Arnett, R. H. Jr. 1973. The beetles of the United States. A manual for identification. The American Entomological Institute, Ann. Ar- bor. 1,112 pp. Balthasar, V. 1963. Monographie der Scarabaeidae und Aphodiidae der Palarktischen und Orien- talischen Region (Coleoptera: Lamellicornia). Band 1-3. Tschechoslowak Akademie der Wissenschaften, Praha. 287 pp. Bates, H. W. 1887-1889. Biologia Centrali-Amer- icana, Insecta, Coleoptera, Vol. II, part 2. Taylor and Francis, London. 432 pp. 627 Boyer, L. B. 1940. A Revision of the species of Hoplia ocurring in America north of Mexico (Coleoptera: Scarabaeidae). Microentomology 51): 1-31. Burmeister, H. C. C. 1844. Handbuch der En- tomologie, Vol. 4. pt. | (Coleoptera, Lamelli- cornia, Anthobia et Phyllophaga Systello- chela). Berlin. 780 pp. Dalla-Torre, K. W. 1912-1913. Coleopterorum Catalogus, Scarabaeidae, Melolonthinae (IV) vol. XX, pars 45, 450 pp. Endrédi, S. 1966. Monographie der Dynastinae (Coleoptera: Lamellicornia) I Teil. Entomolo- gische Abhandlungen der Staatlichen Museum fur Tierkunde, 33: 1-457. Evans, A. V. 2003. Checklist of the New World chafers (Coleoptera: Scarabaeidae: Melo- lonthinae). Zootaxa 211: 1-458. Gemminger, M. and E. von Harold. 1869. Catalo- gus Coleopterorum, hucusque descriptorum synonymicus et systematicus, Vol. 3, Scarabaei- dae. E. H. Gummi, Monachu, 753-958 pp. Goloboff, P. 1999. NONA ver. 2. Published by the author, Tucuman, Argentina. Iablokoff-Khnzorian, S. M. 1977. Uber die Phylo- genie der Lamellicornia. Entomologische Ab- handlungen der Staatliche Museum fiir Tier- kunde 41: 135-199. Jameson, M. L. and B. C. Ratcliffe. 2002. Series Scarabaeiformia Crowson 1960 (= Lamelli- cornia) Superfamily Scarabaeoidea Latreille 1802. Introduction, pp. 1—S. Jn Arnett, R. H. reine @ahomassu lw sees kc lleyam anda ama Frank, eds. American Beetles, Vol. 2. Poly- phaga: Scarabaeoidea through Curculionoi- dea. CRC Press, Boca Raton, Florida. 861 pp. Janssens, A. 1949. Table synoptique et essai de classification pratique des Coleopteres Scar- abaeidae. Bulletin Institute Royal Sciences Naturelles 25(15): 1—30. Kohlmann, B. and M. A. Moron. 2003. Analisis hist6rico de la clasificaci6n de los Coleoptera Scarabaeoidea o Lamellicornia. Acta Zoolo- gica Mexicana (n.s.) 90: 175-280. Lacordaire, T. 1856. Histoire Naturelle des In- sectes. Librairie Encyclopedique de Roret, Paris. 594 pp. Lacroix, M. 1998. Insectes Coléopteres Hopliidae (2epartie). Faune de Madagascar 88(2): 401-755. Lawrence, J. F. and A. F. NewtonJr. 1995. Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names), pp. 779-1006. Jn Paka- luk, J. and S. A. Slipinski, eds. Biology, Phylogeny and Classification of Coleoptera. Papers celebrating the 80th birthday of Roy A. Crowson. Muzeum I Instytut Zoologii PAN, Warszawa. (Reprinted with permission 1n: 628 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Publicaciones Especiales No. 3, Centro de Estudios en Zoologia, Universidad de Guada- lajara, Mexico). Maddison, W. P., M. J. Donogue, and D. R. Maddison. 1984. Outgroup analysis and par- simony. Systematic Zoology 33: 83-103. Medvedev, S. I. 1976. Sistematike 1 Filogeni Plastinchatouyx Zhukov Palearktiki. Entomo- logickoe Oboshchestva 55(2): 400-409. Meinecke, C. C. 1975. Riechsensillen und System- atik der Lamellicornia (Insecta: Coleoptera). Zoomorphologie 82: 1-42. Mico, B. E. 2001. Los escarabeidos antofilos de la peninsula Ibérica (Col. Scarabaeoidea: Hoplii- nae, Rutelidae, Cetontidae); taxonomia, filo- genia y biologia. Tesis Doctoral (no publi- cada). Departamento de Ciencias Ambientales y Recursos Naturales. Universidad de Ali- cante, Espana. 519 pp. Moron, M. A. 2004. Escarabajos, 200 millones de evolucion. Segunda edicion. Instituto de Eco- logia, A.C. y Sociedad Entomologica Arago- nesa, Zaragoza, Espana. 204 pp. Moron, M. A., B. C. Ratcliffe and C. Deloya. 1997. Atlas de los escarabajos de Mexico. Coleop- tera: Lamellicornia, Vol. I Familia Melo- lonthidae. Comision Nacional para el Con- ocimiento y Uso de_ la _ Biodiversidad (CONABIO) y Sociedad Mexicana de Ento- mologia, México. 279 pp. Mulsant, E. 1842. Histoire naturelle des coleopteres de France, pt. 2. Lamellicornes, Paris. 623 pp. Mulsant, E. and C. Rey. 1871. Histoire naturelle des coléopteres de France. Lamellicornes et Pectinicornes. Paris. 736 pp. Nixon, K. C. and J. M. Carpenter. 1993. On outgroups. Cladistics 9: 413-426. Nixon, K. C. 2002. WinClada ver. 1.00.08. Published by the author, Ithaca, New York. Paulian, R. 1959. Coleopteres scarabéides. Faune de France. Lechevalier, Paris. 22 edicion. 298 pp. Paulian, R. and J. Baraud. 1982. Faune des Coléopteres de France, II. Lucanoidea et Scarabaeoidea. Lechevalier, Paris. 477 pp. Peringuey, L. 1902. Descriptive catalogue of the Coleoptera of South Africa. Lucanidae and Scarabaeidae. Transactions of the South Afri- can Philosophical Society, XII, 564-920 Reitter, E. 1902. Bestimmungstabelle der Melo- lonthiden aus der europaischen Fauna und den angrenzenden Landern. IV. Enthaltend die Rutelini, Hoplini und Glaphyrini. Verhan- dlungendas Naturforschendan Briin 41: 27-158. Vereines in APPENDIX I List of the characters used for the phylogenetic analysis. Numbers in pa- & rentheses indicate the state assigned for each character. 1. Body proportions, determined as the total length measured from the apex of clypeus to the apex of elytra divided among the maximum elytral width. Two times longer than wide (0), 1.5 times longer than wide (1), 3 times longer than wide (2). Tegumentary diffraction, considered as iridescent, vitreous or metallic shine of cuticular surface. Present (0), absent (1). 3. Dorso-ventral body — shape. pressed (0), not depressed (1). 4. Position of head with regard to longitudinal axis of body. Down- ward (0), raised (1). 5. Head projections, as horns, tubercles or keels. Absent (0), present (1). 6. Frontoclypeal suture. Complete (0), printed only at sides (1), absent (2). 7. Laterobasal clypeal notch. Absent (O)}s present @)- 8. Clypeal apex. Entire(0), sinuate (1), bilobate (2), dentate (3), ornate (4). 9. Base of pronotum. Rounded (0), straight (1), lobate (2), dentate (3). 10. Lateral margin of pronotum. Entire (0), crenate (1). 11. Elytral vestiture. Without setae or scales (0), with setae (1), with scales (2), with setae and scales (3). 12. Mesoepimera. Dorsally exposed (0), not dorsally exposed (1). 13. Posthumeral notch of elytra. Present (0), absent (1). 14. Setae on posthumeral notch of élytra. Scarce; less than’ 20 ‘setae (0), abundant, more than 20 setae (1), not apply (2). 15. Base of epipleural fold. Thick (0), thin (1). 16. Pygidium proportions, determined by dividing maximum = pygidial width upon total length of pygi- dium. Semitriangular, from 0.60 0.95 times wider than long (0), N De- 24. Metatarsal . Abdominal sternites. . Mandibles. o>) eS) VOLUME 108, NUMBER 3 semicircular, from 1.00—1.91 times wider than long (1), ovate, from 2.20—2.92 times wider than long (2) . Pygidial vestiture. With scales (0), without scales (1). . Number of well defined teeth on pro- tibial external border. 1—2 (0), 3—4 (1). . Metatibia proportions, determined by dividing total length of metatibia upon it maximum width. Narrowed, 4-9 times longer than wide (0), thick- ened, 1—3 times longer than wide (1). 20. Apical spurs of metatibiae. 2 spurs (0), 1 spur (1), without spurs (2). 21. Apices of mesotarsal claws. Cleft in both claws (0), entire in one or both claws (1). . Number of metatarsal claws. One (O), two (1). . Movement of metatarsal claws. Widely retractile (0), scarcely retrac- tile or moveless (1). onychia. Present (Q), absent (1). 25. Shape and position of procoxae. Conical (0), transverse (1). 26. Mesosternal process. Absent (0), present (1). 27. Abdominal vestiture. With scales (0), without scales (1). Fused along midline (0), not fused (1). . Length of fifth abdominal sternite. Two times longer than preceeding (0), with same length of preceeding or shorter than this (1). Not exposed dorsally (0), exposed dorsally (1). _ Form of distal part of left mandible. Shortened (Fig. 4) (0), elongated (Fie Gi) curved (Figs: 355,25) (2): . Apex of left mandible. With teeth (Figs. 3, 5, 25) (0), without teeth (Fig. 6) (1). Conjunctivus of left mandible. Large (Figs. 4, 6) (0), reduced (Figs. 3, 5, 25) Gy): 40. 41. 55: + Dise’ “of 629 . Molar surface of left mandible. With teeth or blades (Fig. 3) (0), with ridges (Figs. 4, 25) (1), without teeth, blades or ridges (Figs. 5—6) (2). . Apex of galea. With teeth (Figs. 7, 26) (0), without teeth (Fig. 8) (1). Lacinia. Reduced (Fig. 8) (0), fused with galea (Figs. 7, 26) (1). Consistency of anterior edge of prementum. Membranous (0), scler- otized (1): . Form of anterior edge of prementum. Entire (Fig. 11) (O), sinuate (Figs. 9— 10,27): (l); motchedi(Fig. 12) @): mentum. Longitudinally furrowed (Fig. 10) (0), without fur- row (Figs. 9,27) Go: Lateral edge of labrum. Elongated (Fig. 16) (0), shortened (Figs. 13, 28) (CD): Thickeness of anterior edge of la- brum. Thick (0), thin (1). 2. Form of anterior edge of labrum. Lobate (Fig. 13) (0), sinuate (Figs. 16, 28) (1), curved (Fig. 14) (2), notched (Fis. 15) (GC): . Consistency of distal edge of labrum. Membranous (0), sclerotized (1). . Basis of spiculum gastrale. Narrowed (0), widened (1), absent (2). . Sclerites associated with spiculum gastrale. Present (0), absent (1). . Sensilla type A. Present (Fig. 17) (0), absent (1). ~ Sensilla’ Lype’ Fe) Present (Pigs? is, 23) (0), absent (1). . Sensilla Type G. Present (Figs. 19, 23) (0), absent (1). . Sensilla Type H. Present (Fig. 20) (O), absent (1). Sensilla Type J. Present (Fig. 21) (0), absent (1). . Sensilla Type K. Present (Fig. 22) (0), absent (1). 2. Sensilla Type EL. Present (Figs, 17— 18. 20-24) (0); ‘absent (1): Sensilla Type H. Present (Fig. 24) (0), absent (1). 630 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 5 Figs. 3-6. Left mandible, adult, dorsal view. 3, Diplotaxis hirsuta. 4, Macrodactylus mexicanus. 5, Anomala cincta. 6, Cotinis mutabilis. C, conjunctivus; M, molar area; S, scissorial area. Scale bar =1 mm. VOLUME 108, NUMBER 3 631 fi 8 Figs. 7-8. Left maxilla, adult, ventral view. 7, Golofa imperialis. 8, Trigonopeltastes geometrica Schaum, 1841. G, galea; L, lacinia. Scale bar =1 mm. 632 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 9-12. Labium, adult, ventral view. 9, Diplotaxis hirsuta. 10, Macrodactylus mexicanus. 11, Hoplia clorophana. 12, Cotinis mutabilis (Gory & Percheron, 1833). Scale bar =1 mm. VOLUME 108, NUMBER 3 633 16 Figs. 13-16. Labrum, adult, ventral view. 13, Diplotaxis hirsuta. 14, Tomarus sallei. 15, Cotinis mutabilis. 16, Trigonopeltastes geometrica. Scale bar =1 mm. 634 Figs. 17-18. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘ Sensilla on inner surface of penultimate antennal segment, 2,500. 17. Cotinis mutabilis. 18. Euphoria basalis. Placoid sensilla types ““A”, ““F” and basiconical sensilla type “L” VOLUME 108, NUMBER 3 635 Figs. 19-20. Sensilla on inner surface of penultimate antennal segment, 2,500. 19. Pelidnota virescens. 20. Callirhinus metallescens. Placoid sensilla types ‘““G”’, ““H”’ and basiconical sensilla type “L”. 636 ocellatus, 22. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 21 ys) 22. Sensilla on inner surface of penultimate antennal segment, Hoplia dispar. Placoid sensilla types “J”, “kK” and basiconical sensilla type 2,500. 21. Isonychus “] s VOLUME 108, NUMBER 3 637 Figs. 23-24. Sensilla on inner surface of penultimate antennal segment, 2,500. 23. Tomarus sallei. 24. Aegidium cribratum. Placoid sensilla types “F”, “G”, basiconical sensilla type “L”’, and trichoid sensilla type “M” 638 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Zi 28 Figs. 25-28. Mouth parts of Hoplia festiva Bates. 25. Left mandible, dorsal view. 26. Left maxilla, ventral view. 27. Labium, ventral view. 28. Labrum, ventral view. Scale bar =1 mm. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 639-645 TEWARIUS REINERT, A NEW GENUS OF AEDINI (DIPTERA: CULICIDAE) JOHN F. REINERT Center for Medical, Agricultural and Veterinary Entomology (CMAVE), United States Department of Agriculture, Agricultural Research Service, 1600/1700 SW 23rd Drive, Gainesville, FL 32608-1067, U.S.A., and collaborator Walter Reed Biosystematics Unit (WRBU), National Museum of Natural History, Smithsonian Institution, Washington, DC (e-mail: jreinert@gainesville.usda.ufl.edu) Abstract.—A new genus, Tewarius Reinert, is described in tribe Aedini and the primary diagnostic characters of females, female genitalia, males, male genitalia, pupae, and fourth-instar larvae are provided. Four species are included in the genus: Tewarius agastyai (Tewari and Hiriyan) (type species), Te. nummatus (Edwards), Te. pseudonummatus (Reinert), and Te. reubenae (Tewari and Hiriyan). Two species groups are designated, the Agastyai and Nummatus Groups. Key Words: mosquito, Diptera, Culicidae, Aedini, Tewarius, new genus Edwards (1923) described Aedes num- matus as a new species and placed it in subgenus Aedimorphus Theobald, but indicated it was an extremely distinct species and strikingly different from all other species of the subgenus. His de- scription was based on only adult hab- itus characters. Edwards (1932) later included Ae. nummatus in the monotypic group B (nummatus-group) of Aedimor- phus. Barraud (1928, 1934) also included Ae. nummatus in subgenus Aedimorphus and stressed it was an aberrant species 1n markings and structure, and differed in many features from all others of the subgenus. He also provided partial illus- trations and brief descriptions of the male genitalia and the damaged larval exuviae. Reinert (1973) redescribed and illustrated all known stages of Ae. nummatus, and those of a new species, Ae. pseudonummatus, and transferred them to subgenus Diceromyia Theobald, but in a separate species group. He provided a comparison of morphological features of the known stages and their structures with subgenera Aedimorphus and Stegomyia Theobald. Tewari and Hiriyan (1992) described and illustrated two new species, Ae. (Dic.) agastyai and Ae. (Dic.) reubenae, of this group. In this important paper they described and illustrated the pupal and fourth-instar larval stages in detail, as well as the adults and genitalia, revealing numerous important morphological characters of the pupa and fourth-instar larva for the first time. After evaluation of all known life stages of the four species mentioned above, and a comparison with other genera of tribe Aedini, it is obvious that they do not belong in any known aedine genus (see Reinert et al. 2004). They are, therefore, transferred to a new genus, Tewarius, which is described below. Tewarius is masculine and named in honor of Satish C. Tewari, Centre for Research in Medical Entomology, Tamil Nadu, India. The proposed two-letter 640 abbreviation for Tewarius is Te. The morphological terminology used follows Harbach and Knight (1980, 1982), except for terms proposed by Reinert (1990, 1999, 2000). The classification of Aedini proposed by Reinert et al. (2004) is used. Tewarius Reinert, new genus Type species: Aedes agastyai Tewari and Hiriyan, 1992 Previous usage.—Aedes (Aedimorphus) of Edwards (1923), in part of Barraud (1928, 1934), Stone et al. (1959); Aedes (Aedimorphus) Group B (nummatus- group) of Edwards (1932); Aedes (Dicer- omyia) in part of Reinert (1973), Knight and Stone (1977), Kaur (2003); and Aedes (Diceromyia) nummatus group of Tewari and Hiriyan (1992). Females.—Head: Vertex with broad, decumbent scales: occiput with several erect, forked scales; ocular line narrow, with broad, pale scales; interocular space with 2 long setae; eyes contiguous above antennal pedicels; clypeus bare; antennal pedicel with several broad scales and short. tine! ‘setae “on” mesal’ ‘Surtiace; maxillary palpus 0.19-0.22 length of proboscis, dark-scaled; proboscis 0.78— 1.10 length of forefemur, dark-scaled. Thorax: Scutum with dark integument, covered with narrow, curved, decumbent Scales except’ bare *prescutellar Varea, scales dark except for large white-scaled patch covering most of anterior approxi- mately 0.50, similar white scales on lateral margins of prescutellar area mesal to setae in Te. agastyai and Te. reubenae; dorsocentral setae present (few anterior and few posterior); acrostichal setae absent; setae few on antealar area, numerous on supraalar area and several on prescutellar area; scutellum trilobed, broad scales on midlobe, lateral lobes with either broad or narrow, curved scales; antepronota widely separated, with scales and several setae; postprono- tum with narrow, curved scales dorsally PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and few broad scales ventrally, few posterior setae; upper proepisternum with broad, white scales, 4-10 setae; lower proepisternum bare; postspiracu- lar area with broad, white scales, 2—5 setae; subspiracular area normally with few, broad, white scales (none seen in Te. nummatus); mesokatepisternum with large patch of broad, white scales ex- tending from upper area over posterior area (small break at about middle of patch in Te. reubenae), few upper and numerous posterior setae; paratergite narrow, with broad, white scales (Te. agastyai and Te. reubenae) or bare (Te. nummatus and Te. pseudonummatus); prealar area with broad, white scales on lower area (Te. agastyai and Te. reube- nae) or without scales (Te. nummatus and Te. pseudonummatus), upper area with 5— 16 setae; mesepimeron with patch of broad, white scales on upper area, several setae on posterodorsal area, lower setae absent; metameron bare. Legs: Fore-, mid-, and hindcoxae with patch of broad, pale scales, several setae: fore- and midfemora with anterior sur- face dark-scaled with basal pale-scaled area, posterior surface mainly white- scaled with dark-scaled areas; hindfemur mainly white-scaled but with distal part of dorsal margins of anterior and poste- rior surfaces dark-scaled; tibiae dark- scaled or with pale-scaled areas on posterior surfaces; tarsi dark-scaled or with pale-scaled areas on posterior sur- faces of some tarsomeres; posttarsi each with 2 equal ungues, fore- and midun- gues each with one tooth, hindungues simple. Wing: Veins dark-scaled (Te. pseudonummatus with small white-scaled patch at base of costa); upper calypter with numerous setae on posterior mar- gin: alula with narrow scales on margin. Abdomen: Terga I-VII each dark-scaled with white-scaled patch basolaterally. Genitalia: Tergum VIII with base and apex slightly concave, numerous broad scales on much of surface, several long, VOLUME 108, NUMBER 3 stout setae on apical margin, setae on distal 0.35—0.42; sternum VIII with base slightly concave, apex with small, median emargination separating small lobe on each side of midline, numerous scales and setae covering most of surface; tergum IX relatively short, apex with small, median emargination, 3—11 setae distally on each side of midline, index 0.56—0.72; insula tonguelike, long, with 1 or 2 small tuberculi on distal 0.20; upper vaginal lip narrow, sclerite large; lower vaginal lip narrow, without sclerite; postgenital lobe moderately long, mod- erately broad, apex rounded, several setae distally: cercus short, moderately wide, apex rounded with few long, stout setae on margin, dorsal surface with several broad scales; 3 spermathecal capsules, one slightly larger than others. Males.—Head: Antenna with whorls containing numerous long setae directed dorsally and ventrally, apical 2 flagello- meres disproportionately long compared to others; maxillary palpus 5-segmented, approximately 0.42—0.50 length of pro- boscis, slender throughout, palpomere 5 short, very few relatively short setae on distal palpomeres, palpomere 5 also with 2 or 3 longer setae; proboscis long, slender. Legs: Fore- and midtarsi with ungues unequal, each with one tooth; hindtarsus with ungues equal, simple. Genitalia: Tergum IX with broad lobe on each side of midline of posterior margin, each lobe bearing 4-11 moderately long, slender setae; gonocoxite short, moder- ately wide, broad scales on lateral and ventral surfaces and outer part of dorsal surface, dorsal surface without apical and basal lobes, with several long, stout setae on outer margin for most of length and numerous moderately long, slender setae especially on proximal part of mesal area, ventral surface with short, slender setae; gonostylus attached to apex of gonocoxite, proximal part broad, forked near midlength into 2 narrow, slightly curved arms, lateral arm with 641 one minute, slender, seta at apex, mesal arm with one short, dark, stout, bluntly pointed gonostylar claw at apex; clasp- ette comprised of narrow, basal setose plaque bearing | or 2 short, dark, stout setae apically and few, short, slender setae subapically, basal plaque connected with its mate by narrow, spiculate aedeagal guide; proctiger moderately long, paraproct moderately to heavily pigmented, with 3 bluntly pointed teeth apically, without cercal setae; phallo- some with aedeagus comprised of 2 lateral sclerites each bearing numerous elongate, curved, pointed, caudally di- rected teeth, paramere moderately long and narrow; sternum IX moderately long, with 2-4 setae on caudomesal area. Pupae.—Cephalothorax: Seta 3-CT single to 2-branched, very long, notice- ably longer than 1-CT; 2-CT branched, short; 5-CT noticeably longer than 4-CT; 6-CT short; 7-CT single to 5-branched, long; 10,12-CT branched; 11-CT single. Trumpet: Moderately long; pinna mod- erately wide; tracheoid area weakly de- veloped. Abdomen: Seta 3-I single, very long, noticeably longer than 6,7-I; 1-II-— IV branched, slender; 2-II single, rela- tively long, inserted lateral to 1-I] and slightly mesal to 3-II; 3-II,III single, noticeably longer than following terga; 5-II inserted lateral to 4-II; 6-II single, very long but shorter than 3-II; 5-IV—-VI very long, single, each longer than combined length of following 2 terga; 6-V longer than 6-III,1V,VI; 2-VI notice- ably mesal to 1,3-VI; 6-VII branched, shorter than and inserted posterior to 9- VII: 9-VII branched, aciculate, normally 1 or 2 branches noticeably longer than others, inserted on posterolateral corner of segment. Paddle: Ovoid, apex sharply rounded, midrib extends to near apex; hairlike, marginal spicules absent; seta 1|- Pa single (rarely forked at tip), very long, about length of paddle. Fourth-instar larvae.—Head: Seta 1-C single, long, slender, attenuate; 4-6-C 642 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON inserted close together, points of inser- tion forming a small triangle; 4-C short, branched, inserted mesal and _ slightly posterior to 6-C; 5-C long, with 2-5 aciculate branches, inserted posterolater- al to 4-C, posteromesal to 6,7-C; 6-C single (rarely 2-branched), very long (noticeably longer than dorsal apotome), inserted lateral to 4,5-C and _ slightly posteromesal to 7-C; 7-C branched, aci- culate, moderately long; 13-C branched, inserted lateral to 12-C; 14-C single. Antenna: Long; with few small spicules, primarily on proximal 0.50; seta 1-A with 2 or 3 attenuate branches; 2,4-A both long, approximately equal in_ length. Mouthparts: Dorsomentum long and rel- atively narrow, with several teeth. Thorax: Seta 1,3-P branched, shorter than 2-P; 2-P single; 4,5,7,8-P branched; 6-P single; 1,4- M, 1,2,6-T branched; 13-T branched, long. Abdomen: Setae 1-I-III branched, slender; 3-I,]] branched; 6-I—III branched, long, 6-I,II aciculate, 6-I] shorter than 6- III; 7-I branched, long; 7,8-II branched, short; 13-III-V branched; 1-VII single, long; 3,7,10,12-VII and 2,4-VIII bran- ched; comb with 20-32 scales; saddle incomplete ventrally; 1-X branched, short, inserted on saddle; 2,3-X long, single (2-X occasionally 2-branched in Te. numma- tus); ventral brush (4-X) with several long, multiple-branched setae with short stems attached to grid with transverse bars, 2 or 3 short, branched, precratal setae. Siphon: Pecten comprised of numerous, long, slender, slightly curved, evenly spaced spines with tiny spicules on ventral margin from base to near apex; seta 1-S multiple- branched, inserted distal to pecten; 6,8-S short. Eggs.—Unknown for all species. Included species (known stages in pa- rentheses).—-Tewarius agastyai (female, male, pupa and larva), Te. nuwmmatus (female, male, and larva), Te. pseudonum- matus (female) and Te. reubenae (female, male, pupa, and larva). Distribution.—Known_ species’ have been reported from the Oriental Region, 1.e., Te. agastvai and Te. reubenae from Western Ghats, India, Te. nummatus from Assam, India, and Te. pseudonum- matus from Chiangmai, Thailand. Bionomics.—Immature stages have been collected from treeholes, bamboo stumps, cut bamboo, and once from a broken bottle in the jungle. Tewarius agastyai immatures were collected in natural forest in high rainfall areas at elevations ranging from 350 to 750 m whereas Te. reubenae immatures were collected in wet evergreen forests at elevations ranging from 850 to 1,600 m (Tewari and Hiriyan, 1992). Diagnosis and discussion.—See Te- wari and Hiriyan (1992) for descriptions and illustrations of the female, female genitalia, male, male genitalia, pupa, and fourth-instar larva of the type species, Te. agastyai. Pupal setae in their fig. 4 are corrected as listed: 1-CT = 5-CT; 2- CT = 1-CT; and 5-CT = 2-CT. These authors also provide a key to the four species I include in Tewarius. The most distinctive characters of each life stage of genus Tewarius follow. Females: Head with vertex covered with broad, decumbent scales, erect forked scales restricted to occiput; eyes contig- uous; maxillary palpus and proboscis dark-scaled; scutum with large, white- scaled patch covering most of anterior approximately 0.50; supraalar area dark- scaled; acrostichal setae absent; dorso- central setae present; scutellum with broad scales on midlobe; mesokatepis- ternum with one large patch of broad, white scales extending from upper area over posterior area (rarely with small break in patch at about midlength); mesepimeron without lower setae; fore- and midungues equal, each with one tooth, hindungues equal, simple; tergum VIII and sternum VIII each with numer- ous broad scales; insula tonguelike with 1 or 2 small tuberculi on distal 0.20; VOLUME 108, NUMBER 3 upper vaginal sclerite large; cercus short, apex rounded, with several broad scales on dorsal surface; and 3 spermathecal capsules. Males: Maxillary palpus 5- segmented, approximately 0.42—0.50 length of proboscis, slender throughout, with few setae; fore- and midtarsi with ungues unequal, each with one tooth, hindtarsus with ungues equal, simple; posterior margin of tergum IX with broad lobe on each side of midline bearing several, moderately long, slender setae; gonocoxite short, without apical and basal lobes on mesal margin of dorsal surface; gonostylus forked near midlength into 2 narrow arms, lateral arm with minute seta apically, mesal arm with short, stout, bluntly pointed gonos- tylar claw apically; claspette comprised of basal setose plaque bearing | or 2 short, stout setae apically and several short, slender setae subapically; cercal setae absent; and aedeagus comprised of 2 lateral sclerites bearing numerous elongate teeth. Pupae: Seta 3-CT single or 2-branched, very long, noticeably longer than 1-CT; 3-I single, very long, noticeably longer than 6-1; 3-II,III single, noticeably longer than following terga; 6-II single, very long but shorter than 3- II; 5-IV—VI single, very long, each longer than combined length of following 2 terga; and paddle ovoid, apex sharply rounded, hairlike spicules on margin absent, seta 1-Pa single (rarely forked at tip), very long, about length of paddle. Fourth-instar larvae: Seta 1-C_ singie, long, slender, attenuate; 4-C short, branched, inserted mesal and _ slightly posterior to 6-C; 5-C long, with 2-5 aciculate branches, inserted posterolater- al to 4-C and posteromesal to 6,7-C; 6-C single (rarely 2-branched), very long (noticeably longer than dorsal apotome), inserted lateral to 4,5-C and _ slightly posteromesal to 7-C; 13-C_ branched; antennal shaft with few small spicules; setae 2,4-A both long; 1,3-P branched, shorter than 2-P; 4-M, 6-T branched; 13- 643 T branched, long; 6-II shorter than 6-III; 1- VII single, long; 2,4- VIII branched; comb with 20-32 scales; saddle incomplete ven- trally; ventral brush comprised of several, multiple-branched, long setae with short stems attached to grid with well developed transverse bars and 2 or 3 short precratal setae; setae 2,3-X both single (2-X occa- sionally 2-branched), long; pecten com- prised of numerous, long, slender, slightly curved, evenly spaced spines with tiny spicules on ventral margin from base to near apex; and seta 1-S multiple-branched, inserted distal to pecten. The large patch of white scales on the anterior half of the scutum is somewhat similar to species of the Gubernatoris Group of Finlaya s.1. Theobald from the Oriental Region. Another Finlaya_ s.l. species, gi//i (Barraud), also has the anterior half of the scutum pale-scaled. These species differ in numerous features of the adult habitus, female and male genitalia, and immature stages from Tewarius. Especially different are char- acters of the male genitalia, e.g., dev- elopment of the gonostylus (long, nar- row, curved), claspette (columnar stem with long claspette filament at apex), aedeagus (tubelike without teeth) and proctiger (with cercal setae), and female genitalia, e.g., insula (liplike with setae laterally). Pupae of Finlaya gilli also have seta 1-Pa very long and single but are easily separated from pupae of Tewarius by having seta 6-VII inserted anterome- sal to seta 9-VII and seta 9-VIII inserted far anterior to the posterolateral corner of segment VII. The larvae differ having a short seta I-C with blunt apex, setae 5,6-C both single and very long, and seta 2-X multiple-branched and noticeably shorter than seta 3-X. The forked gonostylus of Tewarius is somewhat similar to some Afrotropical species of Diceromyia, however, these species have the inner branch of the gonostylus much shorter than the outer branch and the gonocoxite of most of 644 these species also has a patch of long, broad, fusiform scales distally on the mesal margin of the ventral surface. The moderately long and thin male maxillary palpus somewhat resembles those of some Scutomyia Theobald, e.g., Sc. albolineatas (Theobald) and Sc. platylepida (Knight and Hull). However, males of Tewarius differ in a number of important adult habitus, genital, pupal, and larval characters from Scutomyia. Species of Tewarius can be separated into two groups, Agastyai Group con- taining Te. agastyai and Te. reubenae and the Nummatus Group containing Te. nummatus and Te. pseudonummatus. Only adult female characters are used for distinguishing the two groups since the male and immature stages are not known for Te. pseudonummatus and only two damaged larval exuviae are known for Te. nummatus. Also, the pupa is un- known for the latter species. Females of the Agastyai Group have the prescutellar area with narrow, curved, white scales on lateral margins mesal to setae, parater- gite with broad, white scales, upper proepisternum with 4 or 5 setae, and lower prealar area with broad, white scales. Females of the Nummatus Group have the prescutellar area with narrow, curved, dark scales on lateral margins mesal to setae, paratergite bare, upper proepisternum with 9 or 10 setae, and lower prealar area bare. ACKNOWLEDGMENTS Appreciation is expressed to Kenneth J. Linthicum (CMAVE) for providing research facilities; to James E. Pecor and Thomas V. Gaffigan (WRBU) for the loan of specimens from the National Museum of Natural History collection; and to Ralph E. Harbach (The Natural History Museum, London, United King- dom) and Graham B. White (Depart- ment of Entomology and Nematology, University of Florida, Gainesville, FL) for reviewing the manuscript. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Barraud, P. J. 1928. A revision of the culicine mosquitoes of India. Part XXIII. The genus Aedes (sens. lat.) and the classification of the subgenera. Descriptions of the Indian species of Aedes (Aedimorphus), Aedes (Ochlerotatus), and Aedes (Banksinella), with notes on Aedes (Stegomyia) variegatus. Indian Journal of Medical Research 14: 653-669 + pl. LVI-LIX. —— —. 1934. The fauna of British India, including Ceylon and Burma. Diptera, Vol. V. Family Culicidae. Tribes Megarhinini and Culicini. London, UK: Taylor and Francis, 463 pp. + pls. I-VIII. Edwards, F. W. 1923. Mosquito notes. —IV. Bulletin of Entomological Research 14: 1—9. 1932. Diptera Fam. Culicidae. Jn Wyts- man, P., ed. Genera insectorum. Bruxelles, Belgique: Louis Desmet-Verteneuil, Impri- meur-Editeur, 258 pp. + pls. I-V. Harbach, R. E. and K. L. Knight. 1980. Taxono- mists’ Glossary of Mosquito Anatomy. Plexus Publishing, Incorporated, Marlton, New Jer- sey. 415 pp. ——, 1982. Corrections and additions to Tax- onomists’ Glossary of Mosquito Anatomy. (1981). Mosquito Systematics 13: 201—217. Kaur, R. 2003. An update on the distribution of mosquitoes of the tribe Aedini in India (Diptera: Culicidae). Oriental Insects 37: 439_455. Knight, K. L. and A. Stone. 1977. A catalog of the mosquitoes of the world (Diptera: Culicidae), Vol. VI. The Thomas Say Foundation, College Park, Maryland. 611 pp. Reinert, J. F. 1973. Contributions to the mosquito fauna of Southeast Asia. —XVIII. A recon- sideration of Diceromyia Theobald with the inclusion of Aedes Edwards and Aedes pseudonummatus new species (Diptera: Culicidae). Contributions of the American Entomological Institute (Ann Arbor) 19(1): 22-40. —. 1990. Medical entomology studies—X VII. Biosystematics of Kenknightia, a new subgenus of the mosquito genus Aedes Meigen from the Oriental Region (Diptera: Culicidae). Contri- butions of the American Entomological In- stitute (Gainesville) 26(2): 1-119. . 1999. Descriptions of Zavortinkius, a new nummnatus subgenus of Aedes, and the eleven included species from the Afrotropical Region (Diptera: Culicidae). Contributions of the American Entomological Institute (Gainesville) 31(2): 1—105. . 2000. Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part I. Introduc- VOLUME 108, NUMBER 3 tion, preparation techniques, and anatomical terminology. Contributions of the American Entomological Institute (Gainesville) 32(2): 1-17. Remert, Je E> R2 E. Harbach; and’ I. Kitching: 2004. Phylogeny and classification of Aedini (Diptera: Culicidae), based on morphological characters of all life stages. Zoological Journal of the Linnean Society 142: 289-368. 645 Stone, A., K. L. Knight, and H. Starcke. 1959. A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae), Vol. VI. The Thomas Say Foundation, Washington, DC. 358 pp. Tewari, S. C. and J. Hiriyan. 1992. Description of two new species of Aedes (Diceromyia) from south India (Diptera: Culicidae). Mosquito Systematics 24: 154-175. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 646-650 THREE NEW SPECIES OF HYGIA (PTEROCOLPURA BLOTE) (HEMIPTERA: HETEROPTERA: COREIDAE: COREINAE: COLPURINI FROM SUMATRA HARRY BRAILOVSKY Departamento de zoologia, Instituto de Biologia, UNAM, Apdo Postal No. 70153, Mexico 04510 D. F, MEXICO (e-mail: coreidae@servidor.unam.mx) Abstract.—Three new species of the subgenus Hygia (Pterocolpura Bloéte), from Sumatra (Indonesia) are described and illustrated: H. (P.) anthrax, H. (P.) diaphora, and H. (P.) sylvestris. Key Words: Sumatra This paper is a sequel to my earlier papers on the subgenus Hygia ( Ptero- colpura Blote) (Brailovsky 2002, Brai- lovsky and Barrera, in press). A consid- erable quantity of additional Colpurini material collected in 1999 by A. V. Gorochov and members of the Zoolog- ical Institute in St. Petersburg, Russia, has become available, and three new species have been identified. The sub- genus now contains 25 species. The following abbreviations are used for the institutions cited in this paper: UNAM (Instituto de Biologia, Universi- dad Nacional Autonoma de México); ZMAS (Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia). All measurements are in millimeters. Hygia (Pterocolpura) anthrax Brailovsky, new species (Fig. 2) Description.—Measurements: Male: Head length 1.52; width across eyes 1.40; interocular space 0.74; interocellar space 0.30; preocular distance 0.96; length antennal segments: I, 1.42; II, Insecta, Heteroptera, Coreidae, Colpurini, new species, Indonesia, 2.12: Ti. “1365 EV". 229) Pronoume: Length 1.64; maximum width of anterior lobe 1.60; maximum width of posterior lobe 2.72. Scutellar length 1.28; width 1.24. Body length 9.12. Female: Head length? 1272: width across) cyeseul-o2: interocular space 0.80; interocellar space 0.35; preocular distance 1.04; length antennal segments: I, 1.46; II, 2.16; II, 1.46; IV, 1.36. Pronotum: Length 2.08; maximum width of anterior lobe 1.92; maximum width of posterior lobe 3.28. Scutellar length 1.52; width 1.50. Body length 10.64. Male (holotype). Color: Black with following areas yellow to brownish yellow: antennal segment IV (basal third black), dorsal view of postocular tuber- cle, apex of scutellum, an irregular discoidal spot on inner third of apical margin of endocorium, posterior margin of connexivum, rostral segments I to IV, trochanters, anterior lobe of metatho- racic peritreme, and posterior margin of pleural abdominal sterna; anterior half of costal border of corium dark orange, and posterior half black; hemelytral membrane dark brown; femora black VOLUME 108, NUMBER 3 with brownish-orange marks, and basal joint brownish yellow; tibiae black with two yellow rings, one subbasal the other subdistal; tarsi pale chestnut orange; posterior border of abdominal sternite VII brownish orange. Structure: Ros- trum reaching posterior margin of ab- dominal sternite V; buccula with spiny proyection; each frontal angle produced forward as a medium-sized conical lobe, directed forward and slightly outward, and not reaching the postocular tubercle; humeral angles rounded, obtuse, not exposed; hemelytra macropterous, reach- ing apex of last abdominal segment; femora unarmed. Genital capsule: Posteroventral edge tetralobulate, lateral lobes short and conical, submedial lobes broad, and short, and mesally weakly expanded, almost straight (Fig. 2). Female. Coloration similar to male holotype. Connexival segments VIII and IX, dorsal abdominal segments VIII and IX, and genital plates black to reddish brown; outer angle of gonocoxae I yellow. Variation.— Basal entirely black. Type material.—Holotype male, Indo- nesia: Sumatra, Prov. Jambi, 35 km N of Sungaipenuh, env. of National Park Kerinci-Seblat, Mt. Kerinci, 1,500— 2,000 m, 18-22-XI-1999, A. V. Gorochov (ZMAS). Paratypes: 6 6,5 %, same data as holotype (UNAM, ZMAS). Discussion.—This species is similar to Hygia (Pterocolpura) armillata (Bred- din) with the tibiae black to reddish brown and two yellow rings, the rostrum reaching the abdominal sternite V, and the frontal angles produced forward as medium-sized conical lobes, not reaching the postocular tubercles. In H. (P.) armillata, the basal joint of antennal segments II and III are yellow, the femora are yellow with distal thirds black to reddish brown and _ scattered with discoidal and irregular reddish- third of femora 647 brown spots, and the posteroventral edge of male genital capsule is clearly penta- lobulate with median lobe expanded (Figs). anviioCP)) anthrax, antennal segments II and III are entirely black, the femora are almost entirely black with brownish orange marks, and the poster- oventral edge of the genital capsule is tetralobulate, with the space between the submedial lobes weakly expanded to almost straight (Fig. 2). Etymology.—From the Greek “‘an- thrax’ (coal black), referring to the overall dark coloration. Hygia (Pterocolpura) sylvestris Brailovsky, new species (Figs. 3, 8, 10) Description.—Measurements: Male: Head length 1.52; width across eyes 1.50; interocular space 0.72; interocellar space 0.28; preocular distance 0.97; length antennal segments: I, 1.04; II, 1.36; Ei o1003 V5 12085 sPronobum: Length 1.60; maximum width of anterior lobe 1.80; maximum width of posterior lobe 2.80. Scutellar length 1.32; width 1.32. Body length 9.08. Female: Head length 1.58; width across eyes 1.56; interocular space 0.74; interocellar space 0.33; preocular distance 1:02: length antennal segments: I, 1.04; HI, 1.48; II, L208: 1V. L165 Pronotum: Lengthsikso: maximum width of anterior lobe 1.92; maximum width of posterior lobe 3.08. Scutellar length 1.48; width 1.44. Body length 10.12. Male (holotype). Color: Black with following areas yellow to brownish yellow: basal joint of antennal segment III, antennal segment IV (basal third black), apex of scutellum, an irregular discoidal spot on inner third of apical margin of endocorium, posterior margin of connexivum, rostral segments I to IV, trochanters, basal third of hind femur, and posterior margin of abdominal pleural sterna; hemelytral membrane 648 dark brown; tarsi pale chestnut orange. Structure. Rostrum reaching basal mar- gin of abdominal sternite VI; buccula rounded without anterior spiny projec- tion; eyes relatively small and globose; each frontal angle produced forward as small conical lobe; humeral angles rounded, obtuse, not expanded; hemely- tra macropterous, reaching apex of last abdominal segment; femora unarmed. Genital capsule: Posteroventral edge pentalobulate, lateral lobes strongly pro- duced, stout, and conical, submedial lobes broadly quadrate with inner angle curved inward, and mesial lobe evenly trapezoid, with apex slightly bifurcate (Fig: 3). Female.—Coloration similar to male holotype. Connexival segments VIII and IX black with posterior margin dark yellow; dorsal abdominal segments VIII and IX black; genital plates black with outer angle of gonocoxae I dark yellow. Female genitalia as in Figs. 8, 10. Variation.—Anterior lobe of metatho- racic peritreme dark yellow. Type material.—Holotype male, Indo- nesia: Sumatra, Prov. Jambi, 35 km N of Sungaipenuh, env. of National Park Kerinci-Seblat, Mt. Kerinci, 1,500— 2,000 m, 18-22-XI-1999, A. V. Gorochov (ZMAS). Paratype: 1 2°, same data as holotype (ZMAS). Discussion.—Closely related to Hygia (Pterocolpura) tomokunii Brailovsky and Barrera (in press) with the posteroven- tral edge of male genital capsule pentalo- bulate. In H. (P.) tomokunii, the buccula has an anterior spine projection, the hemelytral membrane is pale brown, the fore and middle femora are brown to black with their basal third yellow, the hind femur has the anterior half yellow and posterior half brown, the lateral lobes of male genital capsule are elongate and acute, and the median lobe is clearly bifid (Fig. 4). In H. (P.) sylvestris, the buccula is rounded without an anterior spiny projection, the hemelytral mem- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON brane is dark brown, the fore and middle femora are black, the hind femur is black with only the basal joint yellow, the lateral lobes of the male genital capsule are strongly robust, and the median lobe is trapezoidal with the apex truncated or weakly bifid (Fig. 3). Etymology.—From the Latin “sylves- tris’ (wood), referring to the fact that the species 1s found in association with forests. Hygia (Pterocolpura) diaphora Brailoysky, new species (Figsi ls 7559) Description.—Measurements: Male: Head length 1.96; width across eyes 2.04; interocular space 0.84; interocellar space 0.40; preocular distance 1.20; length antennal segments: I, 1.96; II, 2:92: WMT, 1-96; IV 1-765 Pronotume Length 2.48; maximum width of anterior lobe 2.40; maximum width of posterior lobe 3.68. Scutellar length 1.92; width 1.80. Body length 13.50. Female: Head length 2.08; width across eyes 2.08; interocular space 0.88; interocellar space 0.40; preocular distance 1.22; length antennal segments: I, 2.04; II, 3.20; III, 2.06; IV, 1.80. Pronotum: Length 2.64; maximum width of anterior lobe 2.48; maximum width of posterior lobe 4.08. Scutellar length 2.12; width 2.08. Body length 13.43. Male (holotype). Color: Head black; antennal segments I to III black and IV orange with basal thid black; pronotum black with frontal angles and anterior half of anterolateral border yellow to brow- nish orange; humeral angles and posterior margin with reddish-black marks; scutel- lum reddish black with apex yellow; clavus and corium reddish black with costal border dark orange and small yellow discoidal spot close to apical margin of endocorium; hemelytral membrane brown with veins darker; connexivum reddish black with posterior margin of connexival VOLUME 108, NUMBER 3 649 Figs. 1-10. 2, H. (P.) anthrax. 3, H. (P.) sylvestris.4, H. (P.) tomokuni. 5, H. (P.) pentafurcata. 6. H. (P.) armillata. 7-10, Female genitalia. 7-8. Lateral view. 7, H. (P.) diaphora. 8, H. (P.) sylvestris. 9-10, Caudal view. 9, H. (P.) diaphora. 10, H. (P.) sylvestris. segments III to VI yellow; dorsal abdom- inal segments dark reddish with orange reflections. Ventral coloration: Black with dark reddish reflections; rostral segments I to IV, trochanters, inner face of basal third of hind femur, and posterior margin of pleural abdominal sterna III to VI yellow; posterior margin of propleura and Hygia (Pterocolpura) spp. 1-6, Male genital capsule in caudal view. 1, H. (P.) diaphora. metapleura, and anterior and posterior lobe of metathoracic peritreme brownish orange; femora dark orange brown; tibiae pale orange brown; tarsi yellow with chestnut reflections. Structure: Eyes hemi- spheric, and globose; rostrum reaching posterior border of abdominal sternite V; buccula with spiny projection; each fron- 650 tal angle produced forward as _ slender triangular lobe, apically rounded; humeral angles rounded, obtuse, not exposed; macropterous, reaching apex of last ab- dominal segment; femora with double row of small tubercles. Genital capsule: Posteroventral edge pentalobulate, lateral lobes stout and quadrate, submedial lobes elongate, foli- ate, directed upward and slightly out- ward, and median lobe triangular with apex rounded (Fig. 1). Female.—Coloration similar to male holotype. Connexival segments VIII and IX black with dark reddish reflections, and with posterior border brownish yellow; dorsal abdominal segments VIII and IX and genital segments black with dark reddish reflections; outer angle of gonocoxae | yellow. Female genitalia as im figs) 7.) 9: Variation.—1, Dorsal view of postocu- lar tubercle brownish orange; 2, apical third of rostral segment I dark brown; 3, humeral angles brownish orange; 4, dorsal abdominal segments dark reddish brown. Type material—Holotype male, Indo- nesia: Sumatra, Prov. Jambi, 35 km N of Sungaipenuh, env. of National Park Ker- inci-Seblat, Mt. Kerinci, 1,500—2,000 m, 18-22-XI-1999, A. V. Gorochov (ZMAS). Paratypes: 2 ¢, 2 2, same data as holotype (UNAM, ZMAS). Discussion.—This new species resem- bles Hygia (Pterocolpura) pentafurcata Brailovsy (2002) by the shape of the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON frontal angles, the small yellow dis- coidal spot on the endocorioum, the rostrum nearly reaching the posterior margin of abdominal sternite V, and the tibiae uniformly paler than the femora. In H. (P.) .diaphora; the fore and middle femora are reddish brown, the median lobe of the posteroventral edge of the male genital capsule is triangular, not “bifid likesin va) pentafurcata (Figs. 1, 5), and the fore and middle femora are reddish brown with their anterior half or anterior third yellow. Etymology.—From the Greek “‘dia- phoros” (different), referring to the fact that the species differs from the other ones. ACKNOWLEDGMENTS I thank I. M. Kerzhner (ZMAS) for the loan of these specimens, and E. Barrera (UNAM) for the illustrations. Comments on the manuscript from the anonymous reviewers are greatly appre- ciated. LITERATURE CITED Brailovsky, H. 2002. Two new genera and five new species of Malaysian and Australasian Corei- dae (Insecta: Hemiptera: Heteroptera). Reich- enbachia 34(28): 241—255. Brailovsky, H. and E. Barrera. In press. Seven new species of Hygia (Pterocolpura) Bléte (Hetero- ptera: Coreidae: Colpurini) from Malaysia (Borneo). Journal of the New York Entomo- logical Society. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 651-654 A NEW GENUS AND SPECIES OF POEMENIINI (HYMENOPTERA: ICHNEUMONIDAE) FROM CHINA MaAo-LING SHENG The Key Laboratory for Silviculture and Conservation of Ministry of Education, Beying Forestry University, Beijing 100083, China; General Station of Forest Pest Management, State Forestry Administration, Shenyang, Liaoning, 110034, China (e-mail: shengmaoling@163.com) Abstract.—Dolichotrochanter Sheng, n. gen., and Dolichotrochanter peculiaris Sheng, n. sp., belonging to the tribe Poemeniini of the subfamily Poemeniinae (Ichneumonidae), from Henan Province, China, are described. Key Words: Hymenoptera, Ichneumonidae, Poemeniinae, Poemeniini, Dolicho- trochanter, new genus, new species Poemeniini, a small tribe belonging to the subfamily Poemeniinae of the Ich- neumonidae, comprises eight genera (Wahl and Gauld 1998). Six genera have been reported in China (Sheng 1998, Wang and Gupta 1995, Yu and Horst- mann 1997). In this article, I describe one new genus and its type species collected in China. The morphological terminology fol- lows Wahl and Gauld (1998). Dolichotrochanter Sheng, new genus Description.—Forewing about 10 mm long. Clypeus small, nearly quadrate, about 1.1X as wide as long, basal half strongly convex, rest impressed, apex nearly truncate. Apex of mandible with single chisel-like tooth. Gena in profile about as long as eye, its upper half coarsely scabrous. Occipital carina com- plete. Notauli strong. Forewing vein Icu- a basad of 1/M, distance between them about 0.6 as long as Icu-a. Areolet present. Second trochanter of hind leg very long, ventral side (the longest side) about 0.5 to 0.6 as long as hind femur. Posterior tarsal claw of hind leg with a strong ventral tooth, anterior tarsal claw simple. Metasoma with tergite 1 about 2 as long as posteriorly broad. Tergites 1 to 5 with dense punctures. Last tergite wide, depressed and elon- gate. Ovipositor sheath about as long as front wing. Ovipositor slender, strongly compressed, lower valve tip with about 7 strong subvertical ridges. Type species.—Dolichotrochanter pe- culiaris Sheng, n. sp. Species and distribution.—There is a single Chinese species, described below. Autapomorphies.—Second trochanter of hind leg very long, its ventral side (the longest side) about 0.5 as long as hind femur; posterior tarsal claw of hind leg with a strong ventral tooth, anterior tarsal claw simple. Etymology.—The genus name is based on the very long second trochanter of the hind leg. Gender: masculine. 652 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. femur, ventral view. 4, Tergites 1—3. Dolichotrochanter peculiaris Sheng, new species (Figs. 1-4) Diagnosis.—This species can be dis- tinguished from other poemeniines by Dolichotrochanter peculiaris. 1, Wings. 2, Front tibia, lateral view. 3, Hind trochanters and the characters given above in the generic description. Female.—Body 14mm long. Inner margin of eyes ventrally convergent. Supraclypeal area strongly convex, with punctures, about 9.5 rather sparse VOLUME 108, NUMBER 3 wider than narrowest length. Clypeal suture straight, short and deep. Basal portion of clypeus smooth, apical por- tion with brown hairs, apical margin simple. Malar space very narrow, about 0.1 as long as basal width of mandible, with fine granular surface. Vertex in- flated, with uneven fine punctures, smooth behind interocellar area. Supra- antennal area nearly smooth, front por- tion concave. Antenna filiform with 34 flagellomeres. Occiput mediodorsally flat. Front portion of pronotum smooth, shiny, front and hind margin with dense punctures. Mesoscutum with dense punctures and oblique longitudinal wrin- kles centrally. Front and lower portion of mesopleuron with punctures, middle part with oblique wrinkles and _ short sculpture, speculum and portion under mesopleural fovea smooth. Scutellum and _ postscutellum nearly flat, with dense punctures. Metapleuron swollen, with dense punctures. Forwing vein 2m- cu with two bullae. Hind wing veins I- cu 0.25X as long as cu-a, the latter strongly reclivous. Front side of front tibia with 4 large spines (Fig. 2). Propo- deum with dense punctures, middle portion with fine transverse wrinkles. Propodeal spiracle at midpoint, elliptical and very close to pleural carina. Meta- somal tergites with dense punctures, but gradually weak and unclear from tergite 5 to tergite 7. Tergite 1 without distinct lateral longitudinal carina, mid- dle portion with transverse wrinkles, fused with sternite 1. Tergite 2 with- out groove, about 1.2 as long as its apical width. Tergites 3 to 5 nearly as long as its width. Tergite 6 and 7 slightly transverse. Color: Black; palpi and tegula yellow. Front and middle legs, except blackish brown front and middle Sth tarsal segments, and ventral sides of first trochanters of hind legs, yeliowish brown. Hind coxa, ventral second trochanters, femora except black- sides of 653 ish dorsal sides reddish brown: rest blackish brown. Wings (Fig. 1) brownish hyaline. Stigmata light brown. Veins blackish brown. Male.—Unknown. Type material—Holotype °, China: Ganshan Park, 1000 m, Shanxian Coun- ty, Henan Province, May 31, 2000, Mei- cai Wei and Yi-hai Zhong. Deposited in the General Station -of Forest) Pest Management (GSFPM), State Forestry Administration, Shenyang, Liaoning, China. Host.—Unknown. Etymology.—The species name is based on the peculiar second trochanter of the hind leg. Discussion.—This new genus can be distinguished from other genera of the tribe Poemeniini by: Icu-a basad of 1/ M by about 0.6 its length; areolet present; and second trochanter of hind leg very long, ventral side (the longest side) about 0.5 as long as the hind femur. In the key to the tribes and genera of Poemeniinae (Wahl et Gauld 1998), the new genus can be inserted as follows: 6. Forewing vein 3rs-m present, enclosing an Obliquelysnhiombicsancole tere tienen 7 — Forewing vein 3rs-m completely absent, WAIN INO) WRCS OFAROIS 5 sc posto oso soec 8 7. Forewing vein 2m-cu with two bullae... 7’ — Forewing vein 2m-cu with one _ bulla. Afrotropical Region (South Africa) | Guptella 7’. Posterior tarsal claw of hind leg with a strong ventral tooth. Gena in profile about as long as eye. Icu-a basad of 1/M_ by about 0.6 its length. Eastern Palaearctic Region (China) ... = Dolichotrochanter, n. gen. — Claws of hind leg simple. Gena in profile about 0.4 as long as eye. Icu-a basad of 1/M by about 0.3 its length. Neotropical Region Ganodes ACKNOWLEDGMENTS I express sincere thanks to Prof. Dr. K. Sch6nitzer, Zoologische Staatssamm- lung Miinchen, Germany, for his kind- 654 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ness in offering valuable material. Fi- nancial support from the National Nat- ural Science Foundation of China (NSFC, No. 30471397) is thankfully acknowledged. LITERATURE CITED Sheng, M.-L. 1998. Study on the genus Neoxorides from Northeastern China (Hymenoptera, Ich- neumonidae). Acta Zootaxonomica Sinica 23: 201-204. Wahl, D. B. and I. D. Gauld. 1998. The cladistics and higher classification of the Pimpliformes (Hymenoptera: Ichneumonidae). Systematic Entomology 23: 265-298. Wang, S. F. and V. K. Gupta. 1995. Studies on the tribe Neoxoridini from China (Hymenoptera: Ichneumonidae: Pimplinae). Oriental Insects 29: 175-184. Yu, D. S. and K. Horstmann. 1997. A catalogue of world Ichneumonidae (Hymenoptera). Mem- oirs of the American Entomological Institute. SSele SS oEpp: PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 655-660 A CAVERNICOLOUS SPECIES OF THE GENUS ANELPISTINA (ZYGENTOMA: NICOLETHDAE) FROM SAN SEBASTIAN CAVE, OAXACA, MEXICO Luis ESPINASA AND ADRIANA FISHER Natural Sciences Department, Shenandoah University, 1460 University Drive, Winchester, VA 22601, U.S.A. (e-mail: lespinas@su.edu) Abstract.—Anelpistina acanthocrus, n. sp. (Insecta: Zygentoma: Nicoletiidae) from a cave in Oaxaca, Mexico, is described. The new troglobitic species differs from other described species by the presence of pegs or spines on the legs. Key Words: Nicoletiidae, new species, cave, Mexico The subfamily Cubacubaninae (In- secta: Zygentoma: Nicoletiidae) 1s dis- tributed throughout the New Word (Mendes 1988). Within this subfamily, individuals of the genus Anelpistina Sulvesiri, 1905; are characterized by the presence of 1 + 1 articulated sub- median appendages on urosternum IV in adult males and/or a deep point of insertion of parameres in urosternum IX with the internal face of the coxal processes with macrochaetae more or less sclerotized (Espinasa 1999). How- ever, the monophyly of the group has recently been questioned (Espinasa 2005). Of the 14 species assigned to Anelpis- tina, six inhabit caves, but only A. puertoricensis Espinasa and Baker A\I- pheis, 2003 and A. decui (Wygodzinsky and Hollinger 1977) (see Espinasa 1999 for genus reassignment) have the elon- gated appendages and body of truly cave-adapted Nicoletiids. The new spe- cies described here also shares these adaptive characteristics and probably is a troglobite. MATERIALS AND METHODS Individuals were originally collected by J. Reddell, D. Mckenzie, M. McKen- zie, and S. Murphy from the cave of ““Grutas de San Sebastian” on the 31st of Decembers in’ 1972." in) the stater on Oaxaca, Mexico, and deposited in the collection of the American Museum of Natural History in New York. The four individuals used in this study were found in a vial with ethanol from this collec- tion. Dissections were made with aid of a stereo microscope. The male holotype and a female paratype were mounted in fixed preparations with Hoyer’s solution. The remaining male and female samples were left in a vial with ethanol. All illustrations were made with aid of a camera lucida attached to a compound microscope. Anelpistina acanthocrus Espinasa and Fisher, new species (Figs. 1A-K, 2A—G) Type material.—Male holotype, male paratype, two female paratypes. Grutas de San Sebastian, Oaxaca, Mexico. 31- 656 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON _0.5mm_ 3.0 mm 0.5mm 4 I Tl / fh ay “Ip itt lps = Ar1\h / Ta ‘ Dh VOLUME 108, NUMBER 3 12-72. J. Reddell, B. Mckenzie, M. McKenzie, and S. Murphy cols. De- posited in the American Museum of Natural History, New York, NY. Description.—Maximum body length 16 mm. Maximum conserved length of antenna 18 mm, although broken. Cau- dal appendages also broken. Body pro- portions as in Fig. lA. General color light yellow to white. Head with macrochaetae and micro- chaetae as shown in Figs. 1B—C, with approximately 8 + 8 macrochaetae on border of insertion of antenna. Basal article of antenna in 2/3 as long as first article and with unicellular glands on ventral surface, clustered approximately in four groups and with a row of microchaetae bordering them in form of ae. (rico) On, outside lateral border an extra longitudinal row of four undefined clusters (Fig. 1G). Female basal articles of antenna simple. Mouthparts very long, maxilla as shown in Fig. 1E. Ultimate article of maxillary palp approximately equal in length to penultimate article. Apex of galea with two conules of different widths (Fig. 1D). Two teeth on lacinia plus a third thin hyaline one in between them. Labial palp as in Fig. II, apical article slightly longer than wide and shghtly longer than penultimate article. Penultimate article with not too promi- nent bulge containing two macrochaetae. Labium and first article of labial palp with macrochaetae. Mandible chaeto- taxy as in Fig. 1H, with approximately five macrochaetae. Mesonotum with approximately four 657 Fig. 1K. Hind tibia approximately six times longer than wide and approximate- ly 3/4 length of tarsus. Claws relatively long and with elongate setae. Trochanter of the second leg with two spines inserted on tubercles (Fig. 2A) in both male and female. Abdominal terga and sterna as in other members of genus. Abdominal sterna II—VII subdivided into coxites and sternites. Sterna VIII and IX of male entire. Chaetotaxy of sternites with median portion of sternites with 2 + 2 sublateral macrochaetae at posterior hind borders, but without | + | macro- chaetae near suture at about middle of segment. Appendages of urosternum IV of adult male narrow, more than six times longer than wider and curved outward, similar to A. ruckeri Silvestri, 1905, but very much shorter, reaching only two thirds of stylet of this segment (Fig. 2B). Apex pointed, but without a hook. Urosternum VIII of male emarginate on posterior margin, angles of emargination pointed (Fig. 2E). Ur- osternum IX of male as in Figs. 2E and 2G without a row of sensory cones. Point of insertion of parameres in urosternum IX deep, internal face of coxal processes spiniform and _ extremely elongated, reaching almost apex of parameres (Fig. 2G). Urosternum IX centrally and on coxal processes with some slightly thicker setae, but otherwise not more sclerotized. Stylets [X larger than others, with two macrochaetae and an extra subapical pair. Terminal spine with small teeth. Stylets without modifications in male macrochaetae on lateral borders apart and female. Urotergite X shallowly from several setae of varied sizes emarginate in both sexes, posterior (Fig. 1J). Legs elongate as shown in angles with 2 + 2 macrochaetae and = Fig. 1. Anelpistina acanthocrus. A, Adult male paratype. B-K, Adult male holotype. A, Body propor- tions. B, Head. C, Macrochaetae on border of insertion of antenna. D, Apex of maxilla. E, Maxilla. F—G, Pedicellus. H, Mandible. I, Labium. J, Mesonotum. K, Hind leg. 658 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘ae pan PAN V,i7a7 Peat el fil A Sie = as — Pea Z a Ary oie /- awd SE AAD =~ Sey Wy 74 3, AS OAs 3 Wane ] SSS y f, | Tihs I Fp yes —) Yj (N bot we 7 ISS ~S ie SS —_ ea SS La SES 7) oe ee on se cf SOS SSRs = a —- im a ce2 7 ay ea PeLbow a WE Seaiepr See Sy bdr eds 0.5mm _ ; : Ay SVEN Sahm Cog ~ hy WV] 3s ee Fs , \ Ve hi: Pe Gy VOLUME 108, NUMBER 3 a few relatively strong setae, length of inner macrochaetae almost equal to distance between them (Fig. 2F). Penis and parameres as shown in Figs 2E and 2G. Parameres attaining slightly less than 2 length of stylets IX in adult. Parameres subtriangular and slightly pointed: wide basally and narrow at the apex. Subgenital plate of female rounded to parabolic (Fig. 2C). Oviposi- tor in adult females surpassing apex of stylets IX by % length of stylets (Fig. 2C). Gonapophyses with approxi- mately 19 articles. Cercus of adult male with a longer than wide basal article sometimes fol- lowed by a wider than long, then a very long article bearing numerous spines, followed by numerous short articles of simple chaetotaxy. Spines on inner sur- face consist of a strong, subacute one inserted on a tubercle, a very small one, and a long, acute and slightly curved one also inserted on a tubercle (Fig. 2D). Female cercus simple. Postembryonic development.—Mostly unknown; all samples ranged only within 15 mm to 16 mm. As with other species, younger individuals may have a reduction of secondary sexual characters. It is also likely that in larger males, the appen- dages of urosternite IV may be larger. Distribution.—Known only from the type locality. Etymology.—From the Greek akantha = spine, and the Latin crus = leg (nominative singular). In reference to the spines or pegs in the legs of both males and females. Remarks.—Espinasa and Baker AI- pheis (2001) subdivided Ane/pistina based Fig. 2. 659 on the morphology of the appendages of urosternum IV (Fig. 2B) of the adult males into the following groups: a) species that lack the appendages (A. decui and A. inappendicata Espinasa, 1999); b) species with simple appendages of medium size, their length being slightly longer than the length of urosternum IV and less than five times longer than wide (A. anophtalma Bilimek, 1867, A. wheeleri Silvestri, 1905, A. miranda Silvestri, 1912, A. boneti Wygodzinsky, 1946, A. bolivari Wygod- zinsky, 1946, A. cuaxilotla Espinasa, 1999 and A. puertoricencis); c) species that in addition to the appendages, have two projections on either side of insertion of stylets (A. weyrauchi Wygodzinsky, 1959 and A. ariasae Espinasa, 2005): d) species with very long, simple appendages, their length being approximately equal to twice the length of urosternum IV and more than six times longer than wide (A. ruckeri, A. carrizalensis Wygodzinsky, 1946 and A. doradoi Espinasa and Baker Alpheis, 2001). The new species does not belong to any of the aforementioned groups be- cause, although its appendages have a similar appearance to those of the last group (six times longer than wide), they are extremely short and do not surpass the apex of the stylets (Fig. 2B). Beside the morphology of the appendages of urosternum IV, adult males can _ be differentiated from all other species within the genus by the appearance of the genital area, in particular the internal face of the coxal processes which are spiniform and _ extremely’ elongated, reaching almost to the apex of the parameres (Fig. 2G), and by the tro- Anelpistina acanthocrus. A-B, D—G, Adult male holotype. C, Female paratype. A, Trochanter of the second leg with spines inserted on tubercles. B, Urosternum IV. C, Ovipositor and subgenital plate. D, Cercus (spines on inner surface are shown in dark on the lower-right portion of the figure); E, Male genital area. F, Urotergite X. G, Urosternum IX, coxal process and paramere. 660 chanter of the second leg with two spines inserted on tubercles (Fig. 2A). Females usually are more difficult to differentiate within the genus because the change in ovipositor length during post- embryonic development might create confusion. In the new species the pres- ence of two spines on the trochanter of the second leg makes it easy to differen- tiate from all other species described. The presence of spines or pegs of this kind on any part of the body is typically restricted to adult males. It is interesting that what typically constitutes male sexual secondary characters is also found in females in this new species. ACKNOWLEDGMENTS We thank Randall T. Schuh, George Willett Curator and Chair of the Di- vision of Invertebrate Zoology of the American Museum of Natural History, for kindly making the material available for study. We also thank Monika Baker, Issa J. Rishmawi, Graeme Smith, and Luis F. Mendes for reviewing the man- uscript, and Dr. Tracy Fitzsimmons and Dr. Calvin Allen for the support to publish this manuscript. Partial support was also given to the project by CEAM- ISH-UAEM. LITERATURE CITED Bilimek, D. 1867. Fauna der grotte Cacahuamilpa in Mexiko. Verhandlungen der kaiserlich- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON k6niglichen zoologisch-botanischen Gesell- schaft in Wien 17: 901—908. Espinasa, L. 1999. Two new species of the genus Anelpistina (Insecta: Zygentoma: Nicoletiidae) from Mexican caves, with redescription of the genus. Proceedings of the Biological Society of Washington 112(1): 59-69. . 2005. A new species of the genus Anelpis- tina (Insecta: Zygentoma: Nicoletiidae) from Peru. Proceedings of the Biological Society of Washington 118(2): 337-343. Espinasa, L. and M. Baker Alpheis. 2001. A new species of the genus Ane/pistina (Insecta: Zygentoma: Nicoletiidae) from the Biosphere Reserve Sierra de Huautla. Proceedings of the Biological Society of Washington 114(2): 489-496. Espinasa, L. 2003. A new cavernicole species of the genus Anelpistina (Insecta: Zygentoma: Nicoletiidae) from the Guanica subtropical dry forest, Puerto Rico. Proceedings of the Biological Society of Washington 116(2): 503-508. Mendes, L. F. 1988. Sur deux nouvelles Nicoletii- dae (Zygentoma) cavernicoles de Gréce et de Turquie et remarques sur la systematique de la famille. Revue Suisse de Zoologie 95(3): 751-772. Silvestri, F. 1905. Materiali per lo studio dei Tisanuri. VI. Tre nuove specie di Nicoletia appartenenti ad un nuovo sottogenero. Redia (Firenze) 2: 111-120. 1912. Tisanuri finora noti del Messico. Bolletino del Laboratorio di Zoologia general e agraria di Portici 6: 204-221. Wygodzinsky, P. 1946. Sobre Nicoletia (Anelpis- tina) Silvestri 1905 e Prosthecina Silvestri, 1933. Ciencia 7: 15-25. Wygodzinsky, P. and A. M. Hollinger. 1977. A study of Nicoletiidae from Cuba (Thysanura). Resultats des Expeditions Biospéleologiques Cubano-Roumaines a Cuba 2: 317-324. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 661-671 DESCRIPTION OF THE THIRD-INSTAR LARVA AND PUPA OF LYCOMEDES HIRTIPES ARROW (COLEOPTERA: SCARABAEIDAE: DYNASTINAE: AGAOCEPHALINI) WITH NOTES ON ITS BIOLOGY AND DISTRIBUTION IN COLOMBIA Luts CARLOS PARDO-LOCARNO AND MIGUEL ANGEL MORON (LCPL) Centro Internacional de Agricultura Tropical (CIAT), Cali, Colombia; (MAM) Departamento de Entomologia, Instituto de Ecologia, A.C. Apdo. Postal 63, Xalapa, Veracruz 91000, México (e-mail: moron_ma@ecologia.edu.mx) Abstract.—The larva and pupa of the dynastine scarab beetle Lycomedes hirtipes Arrow from Colombia are described. This larval description 1s the first for the tribe Agaocephalini. A key to the larvae is provided for the genera of Dynastinae in the New World. Some data on the biology and distribution of the species in Colombia are also included. Resumen.—Se describen la larva y la pupa de Lycomedes hirtipes Arrow. Esta representa la primera descripcion larvaria para las especies de la tribu Agaocephalini. Se incluye una clave para separar las larvas hasta ahora conocidas de los géneros de Dynastinae del nuevo mundo. También se presentan algunos datos sobre la biologia y la distribucio6n de esta especie en Colombia. Key Words: Lycomedes, Agaocephalini, immature stages, taxonomy, morphology and Je eValle Colombia, The tribe Agaocephalini contains 11 genera and 43 species distributed from southern Mexico to Argentina (Endrodi 1985, Ratcliffe 2003). Ten species. of Aegopsis Burmeister, Horridocalia End- rédi, Lycomedes Bréme, and Spodistes Burmeister are cited from Colombia (Restrepo-Giraldo et al. 2003). Adults of these species are sporadically collect- ed at lights, on rotten fruits, under loose bark, or in forest soil. Most of the species are associated with humid Andean tropical forests and_ planta- tions located from 1000 to 2100 m altitude. Larvae or pupae of the species of this tribe have not been described, and the habits of immatures are unknown. During field work of the first author (LCPL) in coffee plantations of Cauca departments, a number of dynastid larvae were col- lected under the litter layer. After breed- ing in the laboratory, some of these were identified as pupae and adults of Agao- cephalini. In this paper we describe for the first time the third-instar larva and the pupa of Lycomedes hirtipes Arrow, with ob- servations on the habits, phenology and precise distribution of the species. Also, we present a key to the known third- instar larvae of the tribes and genera of Neotropical Dynastinae. Technical terms are those of Ritcher (1966), Moron (1987), and Costa et al. (1988). Studied specimens are deposited in the private collection of Familia Pardo Locarno (CFPL), Palmira, Valle, Colombia. 662 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Key TO THIRD-INSTAR LARVAE OF AMER- ICAN GENERA OF DYNASTINAE Based on Ritcher 1966, Moron 1987, Moron and Ratcliffe 1990, Moron and Pardo-Locarno 1994, Morelli and Alzu- garay 1994, Morelli 1997, Alvarez-Cas- tillo et al. 1998, Vincini et al. 2000, Morelli and Moron 2003, Ratcliffe 2003, Onore and Moron 2004; Ramirez-Sali- nas et al. 2004, Ocampo and Moron 2004, Ratcliffe and Moron 2005. N Ww 6. Last segment of antenna with | dorsal sensory spot. Most of maxillary stridulato- ry teeth with anteriorly projecting points. Abdominal spiracles of segments 1-4 sim- ilar in size; those of segments 5—8 pro- pres hy SeM@ME «5s poe obla Hoo oem O ME yp) Last segment of antenna with 2 or more dorsal sensory spots. Most of maxillary stridulatory teeth truncate or rounded, without anteriorly projecting points. Size of abdominal spiracles otherwise. ....... 3 Maximum width of head capsule 5—6 mm. Respiratory plate with 13-25 oval holes across any diameter. Maxilla with a row of S—lOkstrdulatonyatcethhe.- ems ee eee Orizabus Fairmaire Maximum width of head capsule 3-4 mm. Respiratory plate with 12-20 elongate, irregularly shaped holes across any diame- ter. Maxilla with a row of 9-12 stridulatory (UETEL nd este Sueeatinn ey ia cola he fo Aphonus LeConte Last segment of antenna with 2—5 dorsal SENSOLYESD OLS. 5 ete eee cpio ee eee eee 4+ Last segment of antenna with 6 or more GOTSAalISENSOMASPOUS ss een tner een anes ee 21 Raster with septula or pallidia. ........ 5) Raster without septula or palidia. ...... 7 Ocelli present. Septula narrowed, not ex- tending across lower anal lip. Palidia monostichous. Maximum width of head capsule 4.0-4.5 mm. ..... Euetheola Bates Ocelli vague or absent. Septula widened, extending across lower anal lip. Palidia polystichous. Maximum width of head Capsule s— lO mmiIme Pa aaeee: Cera Nee 6 Each palidium consisting of 5—7 irregular rows of long pali. Last segment of antenna With)2iGorsall SenSOnveSpOtS. sien eenele : as CE See ote Tomarus Erichson (in part) Each palidium consisting of 2—3 irregular rows of short, spinelike pali. Last segment of antenna with 5 dorsal sensory spots. = fo tae Heterogomphus (s.str.) Burmeister Te 10. ake . Frons Chaetoparia of epipharynx with numerous sensilla among setae, Chaetoparia of epipharynx with few or no Sensillasamon tise lac iene einen mene 10 Head blackish brown. Each acanthoparia with 4-6 setae. Maximum width of head Capsule /=smmim ha ee Head yellow or reddish yellow. Each acanthoparia with 8—13 setae. Maximum width of head capsule 3—S mm. Lateral margins of labrum sharply angulate posteriorly. Surface of cranium slightly roughened. Inner margin of both mand- ibles with a distinct premolar tooth. .... Reta oe ee Dyscinetus Harold Lateral margins of labrum broadly round- ed posteriorly. Surface of cranium reticu- late, shinny. Inner margin of both mand- iblesssmooth) not toothed iene Cyclocephala Latreille Dorsa of abdominal segments 8 and 9 each with 2 widely separated, transverse rows of long setae; short, stout setae absent. .... 11 Dorsa of abdominal segments 8 and 9 each with variable number of short, stout setae and 2 widely separated, transverse rows of long-sctae me rgnis dN aa ee eee 14 Last segment of antenna with 4-5 dorsal sensory spots. Frons with a uniform cov- ering of setae; primary setae of frons difficult to distinguish. Maximum width ofpheadiicapsulei7—Simimns) 4 5) eae SAS aes het Sols es bss Philoscaptus Bréthes Last segment of antenna with 2 dorsal sensory spots. Frons without uniform covering of setae; primary setae easy to distinguish or absent. Maximum width of headicapsules—ommimsaa ss ae Sci en eee 1 without primary setae. Each acanthoparia with 8—9 setae. Head reddish brown. Ancognatha Erichson Xyloryctes Hope Frons with at least | pair of primary setae. Each acanthoparia with 10—11 setae. Head light brown or yellowish brown. .Frons with 1 exterior seta on each side. Posterior frontal setae and anterior frontal absent. Surface of cranium with numerous small pores. Ocelli vague. ... . sah ereuiSgalheie seid RS oe eee eee Oxyerylius Casey Frons with 1-2 exterior frontal setae and | posterior seta on each side. Anterior frontal setae absent. Surface of cranium slightly roughened, reticulate. Ocelli present. ee ONGRD Tae geen oe Tomarus Erichson (in part) setae . Lateral margins of labrum broadly round- ed, NOtaAneulates esis ace caer ene 15 At least one lateral margin of labrum amgulate..o {Als (s U S Lees See 18 VOLUME 108, NUMBER 3 IS). 16. LHe 18. 12 2 tN o>) 24. Foretarsal claw longer than middle and Iniiavel zis GENS 5 S36 se ones ee o'er 16 Foretarsal claw similar in size to middle anG/ormbindutarsaliclaws: = o.4 4. 5 aa45 0. 17 Epicranial setae abundant (more than 15). Right anterior process of hypopharyngeal sclerome much shorter than basal segment of labial palpus. Actinobolus Westwood Epicranial setae scarce (less than 10). Right anterior process of hypopharyngeal scler- ome equal in length to basal segment of labial palpus. Homophileurus Kolbe Ocelli present. Lateral third of frons with deep, coarse, coalescent punctures. ..... 5 OS 6: 6 leo peer erens eee: Trioplus Burmeister Ocelli absent. Anterior two thirds of frons with scattered, small, shallow pits. sada sling Phileurus Latreille (in part) Left lateral margin of labrum angulate... 19 Left and right lateral margins of labrum Al SULALCM wav tea: pagent aatpotes of FEAR 4 20 Frons with 6 anterior frontal setae. Last segment of antenna with 4 dorsal sensory spots. Archophileurus Kolbe Frons without anterior frontal setae. Last segment of antenna with 2 dorsal sensory spots. Phileurus Latreille (in part) . Frons with 2 anterior frontal setae. Hap- tomeral process of epipharynx sinuate. . . ar ee ee NE Phileurus Latreille (in part) Frons without anterior frontal setae. Hap- tomeral process of epipharynx truncate. J cau. 2 Ud te 09 RAC eee eae Hemiphileurus Kolbe Each tarsal claw with 2 setae. Each tarsal claw with 3-4 setae (if tarsal claw with 2 setae, then with lateral margins of epipharynx slightly angulate, Megasoma actaeon L.) . Lateral margins of epipharynx rounded, not angulate. Left lateral margin of epipharynx or both lateral margins of epipharynx slightly angulate. N Nn .Ocelli absent. Spiracles of abdominal seg- ments | to 8 similar in size. Last segment of antenna with 12 dorsal sensory spots. Maximum width of head capsule 10— POINT wer Palate k es cts Wis hs Enema Hope Ocelli present. Spiracles of abdominal segments | to 8 of different sizes. Last segment of antenna with 6-10 dorsal sensory spots. Maximum width of head Gay sule 73 Temi; «oc cha so db es eos Head with numerous pits and 2 anterior frontal setae, without posterior frontal setae. Maxilla with a row of 6 stridulatory teeth. Last segment of antenna with 6—7 dorsal sensory spots. .... Lycomedes Breme i) Nn Head dense and finely punctate, without anterior frontal setae, with 1 posterior frontal seta on each side. Maxilla with a row of 9-10 stridulatory teeth. Last segment of antenna with 8 dorsal sensory spots. Diloboderus Reiche . Inner margin of left mandible with | large tooth. Dorsum of abdominal segment 7 with many short, spinelike setae. Spiracular bulla convex, without knoblike — pro- cess. Dynastes Kirby Inner margin of left mandible with 2 large teeth. Dorsum of abdominal segment 7 without short, spinelike setae. Spiracular bulla with large and acute knoblike pro- cess. Golofa Hope . Each acanthoparia with 9-10 setae. Dorsa of abdominal segment 7 with many short, stout setae. Megasoma Kirby Each acanthoparia with 10—15 setae. Dorsa of abdominal segment 7 without short, stout setae, or with scattered short, spine- like setae. 27 .Spiracles of abdominal segments | to 5 larger than spiracles of segments 6 to 8. Head without anterior frontal setae. Max- imum width of head capsule 14-15 mm . Heterogomphus (Dineterogomphus Prell) Spiracles of abdominal segments 1 to 8 usually similar in size (except in Strategus splendens (Beauvois) with spiracle of ab- dominal segment 8 much smaller). Head with 2-8 anterior frontalsetae. Maximum width of head capsule 9-12 mm. Strategus Hope LARVA OF LYCOMEDES BREME This larval description of Lycomedes hirtipes Arrow is the first for the genus and the tribe Agaocephalini. Based on current knowledge of Dynastinae larvae, the larvae of Lycomedes are most similar morphologically to those of Diloboderus abderus Sturm. The known larvae of Lycomedes have the head capsule with numerous pits and 2 anterior frontal setae, without posterior frontal setae; inner margin of left mandible with 2 teeth; maxilla with a row of 6 stridula- tory teeth; the last segment of the antenna with 6—7 dorsal sensory spots; spiracles of abdominal segments | to 4 progressively larger, spiracles of seg- ments 5 to 7 of similar size, and spiracle of abdominal segment 8 smaller than the 664 preceeding. Larvae of Diloboderus have the head capsule dense and finely punc- tate, without anterior frontal setae, with 1 posterior frontal seta on each side; inner margin of left mandible without tooth; the maxilla with a row of 9-10 stridulatory teeth; the last segment of the antenna with 8 dorsal sensory spots; spiracles of abdominal segments 2 to 6 of similar size, larger than spiracle of segment I, and spiracles of abdominal segments 6 to 8 progressively smaller. Larvae of both genera have defined ocelli; lateral margins of epipharynx rounded, not angulate; each acantho- paria with 9-11 setae; distance between 2 lobes of respiratory plate shorter than dorsoventral diameter of bulla; spiracu- lar bulla convex, without knoblike pro- cess; each tarsal claw with 2 setae; and do not have pallidia or septula. Lycomedes hirtipes Arrow (Figs. 1—15) Description of the third-instar larva is based on three larvae collected in rich organic soil with litter from a coffee plantation shadowed with trees of /nga spp. “guamo” and Erythrina poeppigiana (Walp) ““cachimbo” (Leguminosae); and five third-instar larvae collected under unidentified rotten logs from a coffee plantation or remnants of primary hu- mid forest. Locality data: Colombia: Department of Cauca, Caldono, Pesca- dor, 21-I-2001, 1450 m elevation, M. Trujillo (1 larva); same data except 26- H-2001, M. Trujillo and L.C. Pardo- Locarno (2 larvae); Cauca, Palmira, La Buitrera, 19-XI-2002, 1550 m elevation, L. C. Pardo-Locarno (2 larvae); and Valle, Quindio, Armenia, — II-2003, 1400 m elevation, L. C. Pardo-Locarno (3 larvae)(CFPL). Description of the male pupa is based on one pupa reared from a third-instar larva collected in soil of a shade coffee plantation and two exuvia of pupae PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON reared to adult males. Locality data: Colombia: Department of Cauca, Cal- dono, Pescador, 21-I-2001, 1,450 m ele- vation, M. Trujillo (1 pupa); same data except 26-H-2001, M. Trujillo and L. C. Pardo-Locarno (2 exuvia) (CFPL). Description of the female pupa is based on one pupa and one exuvium of a pupa reared from a third-instar larvae collected in soil of shade coffee planta- tion; and one pupa reared from a third- instar larva collected in organic matter under a rotten log. Locality data: Co- lombia: Department of Cauca, Caldono, Pescador, 21-I-2001, 1,450 m elevation, M. Trujillo (1 pupa and 1 exuvium); Department of Quindio, Armenia, II- 2003, 1400 m elevation, L. C. Pardo- Locarno (1 pupa) (CFPL). Third-instar larva.—Approximate dor- sal body length: 49-65 mm. Head dark to reddish brown. Mandibles_ reddish brown with apical third black. Body yellowish white, with sparse vestiture of setae, and anterior third slightly nar- rowed (Fig. 1). Legs yellowish with apex brownish. Head (Figs. 1-2): Maximum width of head capsule 7.5—-8.2 mm. Cranium shiny, with deep punctures less abundant toward epicranial suture. Frons punctate and finely rugose, with 2 short, anterior frontal setae and single anterior angle seta on each side. Dorsoepicranium with single, long seta on each side. C/ypeus: Shape subtrapezoidal with anterior an- gles widely rounded, and single exterior seta on each side; surface with scattered, large punctures. Preclypeus weakly scler- otized, without setae. Labrum: asymmet- rical, with anterior border slightly tri- lobed, without clithra; irregularly punctate, with row of 4 basal single lateral seta on each side, and 4 anterior, long setae. Epipharynx (Fig. 3): Corypha with 5-7 stout, short setae. Haptomeral process beaklike, strongly sclerotized, with apex entire; 3-4 macro- scopic sensilla behind process. Acantho- setae, VOLUME 108, NUMBER 3 665 none paria with 10-11 short setae. Chaetoparia with 28—34 setae on each side. Gymno- paria narrowed. Dexiotorma moderately wide and long, with short pternotorma. Laeotorma short and narrowed, with large pternotorma. Nesia with sensorial cone elongate and with large sclerotized plate. Haptolachus with 4-5 sensilla bellow sensorial cone. Crepis weakly defined. Mandibles: Right mandible (Figs. 4-5) with | large scissorial tooth anterior to scissorial notch and 3 scissor- ial teeth posterior to notch. Stridulatory area irregular, length 3 times width, consisting of 32—35 transverse ridges of which proximal 8 ridges much broader. Mesad of stridulatory area is a group of Lycomedes hirtipes, third instar larva. Scale line = 5 mm. 7-8 closely appressed setae. Molar area with 4 lobes. Calx wide, prominent. Brustia formed by 3-4 setae. Lateral face with 4 prebasal setae and 2 preapical setae. Left mandible (Figs. 6-7) with 1 large scissorial tooth anterior to scissor- jal notch, IF larse tooth” posterior to notch, and | wide tooth, with sinuate border, on premolar area. Stridulatory area elongate, length 3 times width, consisting of 30-33 transverse ridges of which proximal 7 ridges much broader. Mesad of stridulatory area is a group of 6—7 closely appressed setae. Molar area with 3 lobes. Calx narrow, prominent. Acia narrowed, with rounded apex. Brustia formed by 4-5 setae. Lateral 666 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-7, Lycomedes hirtipes, third instar larva. 2, Head, frontal view. 3, Epipharynx. 4, Right mandible, ventral view. 5, Right mandible, dorsal view. 6, Left mandible dorsal view. 7, Left mandible, ventral view. Scale lines = | mm. face with 2-3 prebasal setae and 1 _ subterminal, conical unci. Surface with preapical setae. Mavxilla: Galea and _ irregular rows of stout setae. Stridulato- lacinia fused (Fig. 8) forming mala. ry area with row of 6 short, truncate Mala with large uncus at apex and 2 teeth and a distal, transverse, truncate VOLUME 108, NUMBER 3 667 Figs. 8-13. Lycomedes hirtipes, third instar larva. 8, Right maxilla, dorsal view. 9, Hypopharynx, dorsal view. 10, Last antennal segment, dorsal view. 11, Last antennal segment, ventral view. 12, Prothoracic spiracle. 13, Raster. Scale lines = 1 mm, except figs. 10-12 = 0.5 mm. process. Labium (Fig. 9): Glossa slightly longer than wide, with 12—16 scattered setae and 6-8 sensilla at middle, and irregular, lateral rows formed by 5-8 setae on each side. Labial palpus short, with distal segment narrowed toward apex. Hypopharyngeal sclerome with wide, erect, truncate process on right 668 14 Figs. 14-15. Lycomedes hirtipes, female pupa. 14, Dorsal view. 15, Ventral view. Scale lines = 5 mm. side, and group of 7—8 setae on left side; both lateral lobes with 5-6 setae ar- ranged in | row. Antenna: First and third segments of same length, second segment 1.2 times longer than first, and last segment 1.6 times longer than _ third. Surface of last segment (Figs. 10-11) with 6—7 dorsal and 8—10 ventral sensory spots. Ocelli clearly defined (Fig. 2). Thorax (Fig. 1): Prothoracie lateral sclerome irregularly shaped, with | erect seta. Pronotum, mesonotum and meta- notum each with 2 long setae. Thoracic spiracle (Fig. 12) with irregular C-shaped respiratory plate 0.7—0.8 mm high and 0.5—0.6 mm wide; distance between 2 lobes of respiratory plate 0.25 times shorter than dorsoventral diameter of bulla; respiratory plate with 23-28 holes across diameter at middle; holes irregu- larly oval. Abdomen (Fig. 1): Spiracles of abdom- inal segments | to 4 progressively larger (0.90-1.13 mm high and 0.87—1.05 mm wide), spiracles of segments 5 to 7 of similar size (1.02-1.05 mm high and 0.96-0.97 mm wide), and spiracle of abdominal segment 8 smaller than the preceeding (0.81 mm high and 0.80 mm PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON wide). Dorsal areas of segments I-V each with sparse vestiture distributed in irreg- ular, transverse rows as follows: each prescutum with | row of 5—10 short setae; each scutum with | row of 14-22 short setae and | row of 4-6 long setae; each scutellum with | row of 10—18 short setae. Dorsal area of segment VI with 1 transverse row of 11 short setae, 1 row of 8—10 short setae, 1 row of 22 short setae, and | row of 5—6 long setae. Dorsal area of segment VII with | transverse row of 11 short setae, 1 row of 6-10 short setae and | row of 6 long setae. Dorsal area of segment VIII with 1 transverse row of 5 long setae. Dorsal area of segment IX with | transverse row of 4 long setae widely separated. Dorsal area of segment X with 36 short setae irregularly distrib- uted, and 24 long setae mixed with 90— 100 short setae distributed near anal slit. Ventral areas of segments I-VI each with transverse row of 4-5 slender, medium size setae, widely separated. Ventral areas of segments VII-IX each with transverse row of 5—7 slender, long setae. Pleural lobes of abdominal segments I-— VIII with sparse short setae. Raster (Fig. 13) without palidia or septula. Teges formed by 55—60 stout, short setae. Campus with 4 slender, long setae. Lower anal lip with 45—50 stout, short setae and | transverse row of 10—12 long setae on border of anal slit. Pupa.— Male: Body length: 35-40 mm. Form elongate, robust, exarate. Orange yellow to reddish yellow, with velvety, golden yellow microtrichia on abdominal segments, mainly toward last segment. Head: Reflexed downward; antenna, mouth parts and ocular canthus well differentiated. Compound eyes _nar- rowed. Fronto-clypeal region with short, wide projection (1.5—-1.8 mm _ length) apically truncated. Thorax: Pronotum wider than long, convex, with a transverse small process or tubercle on middle of anterior half; lateral borders weakly angled; basal VOLUME 108, NUMBER 3 border nearly straight. Meso- and meta- nota differentiated. Mesoscutellum wide- ly rounded posteriorly. Metascutellum widely sinuated posteriorly. Ecdysial suture running from anterior border of pronotum to posterior border of meta- notum. Pteroteca narrowed, free, with apex rounded, compressed around body; hind wing teca slightly longer than elytron teca. Protibia with 3 pigmented, small areas on external border; meso- and metatibiae without projections or pigmented areas on external borders but with apical spurs differentiated; all tarsomeres weakly defined: protarsus with apex slightly enlarged and pigmen- ted. Abdomen: Tergites I-VIII convex, with 5 pairs of well sclerotized and pigmented dioneiform organs between segments II-VI and transverse sclero- tized carina on anterior border of seg- ment VII. Each dioneiform organ 2.5mm wide. Tergo-lateral tubercles absent on all segments. Spiracle I elon- gated, not prominent. Spiracles H-—IV large, with ringlike, sclerotized peri- treme, slighly directed forward. Spiracles V—-VII closed, slightly prominent, sur- rounded by fine rugae. Spiracle VIII weakly indicated by fine rugae. Sternites II-VII convex. Last segment without urogomphi, with lateroventral fleshy lobes and prominent genital ampulla on middle of ventral surface. Female: Body length: 34-36 mm (Figs. 14-15). Similar to male except on head, thorax, and protarsus. Head: Fronto-clypeal area swollen, with round- ed borders and deep hole in middle. Thorax: Pronotum broadly convex, with lateral borders weakly angled, without DEOCESSs OG, Cubercle, jon. ~middle, of anterior half. Protarsus with same pro- portions of meso- and metatarsi, not enlarged. Biology and distribution.—Third-in- star larvae of Lycomedes hirtipes were observed in rich organic soils with an 669 abundance of litter in mountain tropical forests and old coffee plantations under dense shade. Larvae of other species of Dynastinae frequently found in these conditions are Podischnus agenor Olivier, Strategus aloeus (Linné), Golofa eacus Burmeister and Aspidolea fuliginea Bur- meister. Larvae of L. hirtipes exhibit thanatosis when they are exposed by turning the head downward, retracting the legs, and curving tightly the body, and remaining quiet for a long time, thus simulating death until the disturbance 1s reduced. Then they dig into the soil with slow movements. The larvae of other sympatric species of Dynastinae are very active when exposed, and quickly crawl out of the disturbed area and return to soil. Adults of L. hirtipes are dawn flyiers. They sometimes feed on mature fruits and are active mainly during the sum- mer-autumn rainy season in Colombia. According to Reyes-Usuga (1995) they were attracted to black lght traps located in Pance, Cali, Department of Valle during May (2), September (4), October (38), November (24), and De- cember (1). Also, they were attracted to fruit baited traps located in Nirvana reserve, La Buitrera, Palmira, Depart- ment of Valle during August (8), Sep- tember (9), October (39), November (6), and December (2) of 2000 (Serna and Pardo-Locarno personal communica- tion). No eggs were obtained from adults in captivity, so the time for a complete life cycle remains unknown. Field rec- ords for adults and rearing of third- instar larva collected in January to obtain pupae in July and adult emer- gence in August suggest a one year life cycle. Restrepo et al. (2003) cited L. hirtipes from the Colombian departments of Boyaca, Caldas, Cauca, Huila, Tolima, and Valle. Recently, the first author (LCPL) collated data from national 670 collections and found specimens collect- ed at the following localities: Antioquia (Supia), Cauca (Caldono, Popayan, La Sierra), Huila (Gigante), Narino (La Union), Quindio (Armenia, La Tebaida), Tolima (Ibagué, China Alta), Risaralda (Santa Rosa, La Florida), and Valle (Calusé, Felidia, Calima-Darién, San Antonio-Jamundi, including Pance, To- cota, Saladito and Yanaconas-Cali dis- tricts). All the above localities are between 1,200 and 2,100 m elevation along the slopes of the western and central Andes in Colombia (approx. 1°— TAN 74 =7 7 IW) sto ‘ther owest vol “he Magdalena River. A record from de- partment of Boyaca (Togui) need to be confirmed. ACKNOWLEDGMENTS The first author (LCPL) thanks stu- dents Andrés Felipe Serna, Luis Carlos Reyes Usuga, technicians Hoover and Manuel Trujillo (Caldono), and Federico Botero (owner of the Reserva Nirvana, La Buitrera, Palmira) for their help during field work. Claudia Patricia Vélez-Garcia (Palmira) assisted with rearing of larvae. Jhon César Neita (Facultad de Agronomia, Universidad Nacional, Bogota), John Albeiro Quiroz (Museo Francisco Luis Gallego, Univer- sidad Nacional, Medellin) and Carmen Elisa Pozo (Universidad del Valle) pro- vided valuable assistance during the study of collections in their care. Thanks also to professor James Montoya (Uni- versidad del Valle) for his valuable suggestions. Special acknowledgment to the Centro Internacional de Agricultura Tropical (CIAT, Cali) and to the project “Plagas subterraneas en Suramérica”’ (Convenio CIAT-GTZ) for job and bibliographical support. This paper pre- sents results of “‘“Contribucion al con- ocimiento sistematico y bioldgico de los Scarabaeoidea (Coleoptera) de Colom- bia’’, a doctoral dissertation project of the first author at the Universidad del PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Valle. Also, this paper is a contribution to the project ““Coleopteros Lamelicor- nios de América Latina’? supported by the Instituto de Ecologia, A.C. Xalapa (account 902-08-011). LITERATURE CITED Alvarez-Castillo, H. A., A. M. Vincini, A. N. Lopez, D. M. Carmona, and P. L. Manetti. 1998. Descripcion de los estados inmaduros de Heterogomphus pauson (Perty, 1830) (Coleop- tera: Scarabaeidae: Dynastinae). Elytron 12: 97-103. Costa, C., S. A. Vanin, and S. A. Casari-Chen. 1988. Larvas de Coleoptera do Brasil. Museo de Zoologia, Universidade de Sao Paulo, Sao Paulo, 282 pp. Endr6édi, S. 1985. The Dynastinae of the World. Dr. W. Junk Publishers, Dordrecht, 800 pp. Morelli, E. 1997. Descripcion de los estados inmaduros y notas sobre la biologia de Diloboderus abderus (Sturm, 1826) y Philos- captus bonariensis (Burmeister, 1847) (Coleop- tera: Melolonthidae, Dynastinae). Acta Zool- ogica Mexicana (n.s.) 71: 57—70. Morelli, E. and R. Alzugaray. 1994. Descripcion de la larva de Cyclocephala testacea Burmeister, 1847 y clave para la determinacion de larvas de cuatro especies del género Cyc/locephala en el Uruguay (Coleoptera: Dynastinae). Revista Brasileira de Biologia 54(1): 77-84. Morelli, E. and M. A. Moron. 2003. Descriptions of Xv/oryctes Hope larvae with a key to species based on the third stage larvae (Coleoptera: Scarabaeidae: Dynastinae). The Coleopterists Bulletin 57(3): 289-295. Moron, M. A. 1987. Los estados inmaduros de Dynastes hyllus Chevrolat (Coleoptera: Melo- lonthidae; Dynastinae) con observaciones so- bre su biologia y el crecimiento alométrico del imago. Folia Entomologica Mexicana 72: 33-74. Moron, M. A. and L. C. Pardo-Locarno. 1994. Larvae and pupae of two species of Golofa Hope (Coleoptera: Melolonthidae: Dynasti- nae) from Colombia. The Coleopterists Bulle- tin 48(4): 390-399. Moron, M. A. and B. C. Ratcliffe. 1990. Descrip- tions of Strategus larvae with a key to species based on the larvae (Coleoptera: Scarabaeidae: Dynastinae). Elytron 4: 53-66. Ocampo, F. C. and M. A. Moron. 2004. De- scription of the third instar larva of Hemi- phileurus dispar Kolbe (Coleoptera: Scarabaei- dae: Dynastinae: Phileurini). Proceedings of the Entomological Society Washington 106(2): 412-416. VOLUME 108, NUMBER 3 Onore, G. and M. A. Moron. 2004. Dynastes neptunus Quenzel (Coleoptera: Scarabaeidae: Dynastinae); descriptions of the third instar larva and pupa, with notes on biology. The Coleopterists Bulletin 58(1): 103-110. Ramirez-Salinas, C., M. A. Moron, and A. Castro- Ramirez. 2004. Descripcion de los estados inmaduros de tres especies de Anomala, An- cognatha y Ligyrus (Coleoptera: Melolonthi- dae: Rutelinae y Dynastinae) con observa- ciones de su _ biologia. Acta Zoologica Mexicana (n.s.) 20(3): 67-82. Ratcliffe, B. C. 2003. The Dynastine scarab beetles of Costa Rica and Panama (Coleoptera: Scarabaeidae: Dynastinae). Bulletin of the University of Nebraska State Museum 16: 1—S06. Ratcliffe, B. C. and M. A. Moron. 2005. Larval descriptions of eight species of Megasoma Kirby (Coleoptera: Scarabaeidae: Dynastinae) with a key for identification and notes on biology. The Coleopterists Bulletin 59(1): 91-126. 671 Restrepo-Giraldo, H., M. A. Moron, F. Vallejo, L. C. Pardo-Locarno, and A. Lopez-Avila. 2003. Catalogo de Coleoptera Melolonthidae (Scar- abaeidae-Pleurosticti) de Colombia. Folia En- tomologica Mexicana 42(2): 239-263. Reyes-Usuga, L. C. 1995. Contribucion al inven- tario y estudio ecologico de las plagas riz6fagas (Coleoptera: Melolonthidae: Dynastinae) de la Cuenca Alta y Media del Rio Pance, Parque Nacional Natural Los Farallones de Cali, Valle del Cauca, Colombia. Tesis de grado (no publicada) Universidad Nacional de Co- lombia-Palmira, 81 pp. Ritcher, P. O. 1966. White grubs and their allies. A study of North American Scarabaeoid larvae. Oregon State University Press, Corvallis, 219 pp. Vineini, A. M., A. N. Lopez, P. L. Mannetti, H. Alvarez-Castillo, and D. M. Carmona. 2000. Descripcion de los estados inmaduros de Dyscinetus rugifrons (Burmeister, 1847) (Co- leoptera: Scarabaeidae: Dynastinae). Elytron 14: 91-98. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 672-676 NEW COMBINATIONS IN SHARPSHOOTER LEAFHOPPERS (HEMIPTERA: CICADELLIDAE: CICADELLINAE) STUART H. MCKAMEY Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Institution P.O. Box 37012, MRC-168, Washington, D.C. 20560, U.S.A. (e-mail: smckamey(@sel.barc.usda.gov) Abstract.—Nine species-group name changes in Cicadellinae are proposed so that all species in the subfamily can be listed in a single classification that is aligned with the International Code of Zoological Nomenclature. Bothrogonia tangmaiana Yang and Li and Mareba panamensis Young are fixed as the correct original spellings. Five new combinations are proposed: Hadria alayoi and H. zayasi (Dlabola and Novoa), Caribovia intensa nigrinervis (Schréder), and Cicadella viridis suffusa (Salmon), and C. intermedia (Rao). The combination Cardioscarta flavifrons transversa Melichar 1s reinstated. Diedrocephala bimaculata (Gmelin) is reinstated as the valid name for the species of the genus most often cited in literature and collections as D. variegata (Fabricius). Key Words: Homoptera, Auchenorrhyncha, Caribovia, taxonomy In the course of preparing a database of world leafhoppers, it became apparent that many purely nomenclatural changes were needed so that all species could be included in a single classification. These changes must be made separately, prior to WWwW-based dissemination of the database, to avoid unwitting prolifera- tion of unpublished taxonomy or of taxonomy that is not in accordance with the International Code of Zoological Nomenclature (ICZN; International Commission on Zoological Nomencla- ture 1999). For example, over 20 generic homonyms were discovered. Most of these generic changes were already made (McKamey 2003, McKamey 2006), in- cluding their affected species and one tribe. This is the first installment of species-level changes and is restricted to Cicadellinae sensu stricto, i.e., the tribes Cicadellini and Proconiini, commonly called sharpshooters. Given the recency of comprehensive revisionary work and other descriptions, the required changes in Cicadellinae are fortunately few. World sharpshooters were revised by Young (1968, 1977, 1986) and since the relevant portions of Young’s revision, dozens of New World species have been added by various authors, principally by workers in Costa Rica (e.g., Godoy and Nielson 2000) and Brazil (e.g., Mejdalani et al. 2000, Cavichioli 2003). Old World additions include dozens more species, from more countries and by more authors (e.g., Ara and Ahmed 1988, Hayashi and Arai 1990, Chiang and Knight 1991, Ge and Zhang 1991, Huang 1992, Li and Wang 1992, Kuoh and Zhang 1992, Cai and Kuoh 1995, Yang and Zhang 2001). All this work together, however, indicates VOLUME 108, NUMBER 3 that wherever there are active workers, new species are still being discovered at a rapid pace, suggesting a vast number still await discovery. This probability is especially impor- tant considering the role of sharpshoo- ters in transmitting pathogenic bacteria such as Xylella fastidiosa, the species responsible for Citrus Variegated Chlo- rosis in oranges (Milanez et al. 2003), Crespera disease in coffee (Rodriguez et al. 2001), and Pierce’s Disease in grapes (Almeida and Purcell 2003). TAXONOMY Bothrogonia tangmaiana Yang and Li, fixed spelling Bothrogonia tangmaiana Yang and Li 1980: 203 [n. sp.]. Mareba panamensis Young, fixed spelling Mareba_ panamensis 1968: 64 [n.sp.]. Young The principle of first reviser (ICZN Art. 24) is applied here to fix the spelling of both species, which were spelled in two ways within the original descrip- tions. The spelling “‘tongmaiana”’ is deemed incorrect because the species was collected in Tangmai, China. This fixation is not meant to reflect its taxonomic status. Young (1986) syno- mized B. tongmaiana [sic] Yang and Li (1980) under B. curvata Yang and Li (1980). Young (1968: 63) misspelled the Panamanian species as “‘panamenis.” Hadria alayoi (Diabola and Novoa), new combination Arezzia alayoi Diabola and Novoa 1976: 2in.spel: Hadria zayasi (Dlabola and Novoa), new combination Arezzia zayasi Diabola and Novoa 1976: 4 [n.sp.]. 673 Dlabola and Novoa’s (1976) new Cuban cicadellines were understandably not included in Young’s (1977) revision of New World Cicadellini. Both species above were described under the genus Arezzia Metcalf and Bruner, which Young (1977) synonymized under Ha- dria Metcalf and Bruner. Dlabola and Novoa (1976) also referred to Arezzia the species Hortensia conciliata Metcalf and Bruner, which Young (1977) placed in Hadria. Caribovia intensa nigrinervis (Schréder), new combination Tettigella intensa_ nigrinervis Schréder 1959: 66 [n.subsp.]. Young (1977) referred the nominate species, Tettigonia intensa Walker to his new genus Caribovia but was apparently unaware of the subspecies Tettigella intensa nigrinervis Schroder (1959). It merits mention that Caribovia species are known to feed on coffee. The generic placement of this El Salvadorean sub- species deserves reevaluation, as all other taxa in the genus, including C. intensa, are endemic to the Greater Antilles. Cicadella viridis suffusa (Salmon), new combination Tettigella viridis suffusa Salmon 1954: 64 [n.subsp.]. Cicadella intermedia (Rao), new combination Rao 1980: 190 Tettigella intermedia [n.sp.]. Salmon (1954) described and illustrat- ed the “variety” Tettigella viridis suffusa from Great Britain in a_ publication seemingly overlooked by the major compendia of Auchenorrhyncha: Met- calf (1964), Nast (1972), Oman et al. (1990), as well as Young (1968, 1977, 1986). Because Salmon (1954) did not unambiguously reveal that it was infra- 674 subspecific in rank, and the variety was described before 1961, it is deemed to be of subspecific rank (ICZN Art. 45.6.4). At the time Salmon described the sub- species, Tettigella viridis was the valid name of the nominate form. Tettigella China and Fennah was suppressed by the International Commission of Zoological Nomenclature (Opinion 647) making Cicadella the valid name (China 1963). Rao (1980) described a new species from India in the suppressed name Tettigella (see above). Cardioscarta flavifrons transversa Melichar, reinstated combination Cardioscarta flavifrons transversa Meli- char 1932: 313 [nesubsp.|: Poeciloscarta flavifrons transversa (Meli- char): Metcalf 1965: 67. Poeciloscarta _flavifrons transversa (Melichar, 1932) is another subspecies not included in Young’s (1977) revision. It is here referred to the genus to which Young (1977) referred the nominate species, Cardioscarta flavifrons (Sign- oret). Both the nominate form and the subspecies are known only from Brazil. Diedrocephala bimaculata (Gmelin), reinstated validity Cicada bimaculata Gmelin 1789a: 2106 [nom. nov. for Cicada variegata Fab- ricius 1775: 684]. Fabricius (1775: 684) described Cicada variegata and two pages later (Fabricius 1775: 686) described another Cicada variegata. These two nominal species are indeed different taxonomic species, the former a species in the cicadelline genus Diedrocephala Spinola (Membra- coidea: Cicadellidae) and the latter now Cixius variabilis Metcalf (Fulgoroidea: Cixidae). In the absence of any other treatment, either homonym would need to be replaced under the current Code, as they PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON have not been considered congeneric after 1899 (Art. 23.9.5). Nevertheless, the name has already been replaced. Gmelin (1789), acting as first reviser (ICZN Art. 24.2), regarded the cica- dellid Cicada variegata as a junior hom- onym and proposed Cicada bimaculata as its new name. Unfortunately this name has been overlooked and the name Diedrocephala_ variegata is in common usage. Unfortunately the Zoological Code does not specifically mention homonyms published in the same work (and which are therefore neither senior nor junior homonyms), or when one of these is replaced and becomes by default the junior homonym. Although ICZN Art. 23.9.1 does mention prevailing usage, its intention is to help decide which of two homonyms is the junior and which is the senior, when each are described in different works.. In the case of [ibe cicadellid Cicada variegata there is no problem as the decision on priority has already been made (see above). In any case, even if the same prevailing usage criteria were used, although Diedroce- phala variegata is a well-known species, its mention in the literature falls far short of the Code’s requirements of prevailing usage to enable its maintenance as the valid junior homonym of the cixiud Cicada variegata or as the valid senior objective synonym of the leafhopper Cicada bimaculata. Hence, the valid name of the primary junior homonym is Diedrocephala bimaculata (Gmelin). Diedrocephala bimaculata is the most common and widely distributed species of the genus, occurring from Mexico southward through Paraguay and Ar- gentina (Young 1977). ACKNOWLEDGMENTS I thank F: C. Fhompson, E> Deitz: J. Brown, and two anonymous reviewers for helpful comments on previous drafts of the manuscript. VOLUME 108, NUMBER 3 LITERATURE CITED Alimetda hen ean Gee ACE Eeuncell s2003¢ Transmission of Xylella fastidiosa to grape- vines by Homalodisca coagulata (Hemiptera: Cicadellidae). Journal of Economic Entomol- ogy 96: 265-271. Ara, A. and M. Ahmed. 1988. Some new species of leafhoppers (Cicadellidae: Homoptera) from Pakistan. Pakistan Journal of Zoology 20(3): 289-297. Cai, P. and C. [L.]. Kuoh. 1995. A new record genus and five new species of Cicadellidae from China (Homoptera: Cicadelloidea). Acta Entomologica Sinica 38(4): 460-466. Cavichiohi, R. R. 2003. Fingeriana dubia gen. nov. e sp. nov. de Cicadellini (Hemiptera, Auchenor- trhyncha, Cicadellidae) do sudeste e sul do Brasil. Revista Brasileira de Zoologia 20(2): 247-249. Chiang, C. C. and W. J. Knight. 1991. Mileewanini of Taiwan. Journal of Taiwan Museum 44(1): iV 2As China, W. E. 1963. Cicadella Latreille, 1817 (Insecta, Hemiptera); validation under the plenary powers. Bulletin of Zoological No- menclature 20(1): 35-38. Dlabola, J. and N. Novoa. 1976. Dos nuevas especies del género Arezzia Metcalf y Bruner, 1936 (Homoptera: Auchenorrhyncha) y revision de otras especies Cubanas. Poeyana 158: 1—27. Fabricius, J. C. 1775. Ryngota. Systema Entomol- giae, Sisten insectorum classes, ordines, genera, species, adiectis synonymis, locis, descriptioni- bus, observationibus, Korte, 816 pp. Ge, Z. L. and Z. M. Zhang. 1991. A new genus and two new species of Cicadellidae (Homoptera: Cicadelloidea). Acta Zoologica Sinica 37(4): 367-370. Gmelin, J. F. 1789. Insecta Hemiptera. Caroli a Linné Systema Naturae 1(4): 1517—2224. Godoy, C. and M. W. Nielson. 2000. A review of the leafhopper genus Acrulogonia Young with descriptions of new species (Homoptera: Cica- dellidae: Cicadellinae). Entomological News 111(2): 107-115. Hayashi, M. and K. Arai. 1990. Five new species of the genus Pagaronia Ball (Homoptera, Cica- dellidae-Cicadellinae) from central Honshu. Esakia. Special Issue 1, 5—13. Huang, K. W. 1992. Evacanthini of Taiwan (Homoptera: Cicadellidae: Cicadellinae). Bul- letin of National Museum of Natural Science 3: 159-185. International Commission on Zoological Nomen- clature, International Code of Zoological Nomenclature. 4th ed. International Trust for Zoological Nomenclature, London, xxix + 306 pp. 675 Kuoh, C. L. and Z. Zhang. 1992. [Three new species of the genus Anatkina (Homoptera: Cicadelloi- dea).] Entomotaxonomia 14(2): 111-115. Li, Z. Z. and L. Wang. 1992 [1991]. Agriculture and Forestry Insect Fauna of Guizhou (Homoptera: Cicadellidae.), Vol. 4. Guizhou Science and Technology Publishing House, Guiyang, 304 pp. McKamey, S. H. 2003. Some new generic names in the Cicadellidae (Hemiptera: Cicadellidae: Deltocephalinae, Selenocephalinae). Proceed- ings of the Entomological Society of Washing- ton 105(2): 447-451. . 2006. Further new genus-group names in the Cicadellidae (Hemiptera). Proceedings of the Entomological Society of Washington 108: 502-510. Mejdalani, G., M. Felix, and D. M. Takiya. 2000. Description of a new species of Amblyscartidia Young from southeastern Brazil (Hemiptera: Cicadellidae: Cicadellinae). Bollettino di Mu- seo Regionale di Scienze Naturali Torino 17(1): 131-139. Melichar, L. 1932. Monographie der Cicadellinen. IV. Musei Nationalis Hungarici Annals 27: 285-328. Metcalf, Z. P. 1964. General Catalogue of the Homoptera. Fascicle VI. Cicadelloidea. Bibli- ography of the Cicadelloidea (Homoptera: Auchenorrhyncha). United States Department of Agriculture, Agricultural Research Service, 349 pp. 1965. General Catalogue of the Homo- ptera. Fascicle VI. Cicadelloidea. Part 1. Tetti- gellidae. United States Department of Agri- culture, Agricultural Research Service, 730 pp. Milanez, J. M., J. R. P. Parra, I. A. Custédio, D. C. Miagrinn Gas Cerasrand suse San leoOpesue2 005» Feeding and survival of citrus sharpshooters (Hemiptera: Cicadellidae) on host plants. Florida Entomologist 86(2): 154-157. Nast, J. 1972. Palearctic Auchenorrhyncha (Homo- ptera): An Annotated Check List. Institute of Zoology, Polish Academy of Sciences, Polish Scientific Publisher, Warsaw, 550 pp. Oman, P. W., W. J. Knight, and M. W. Nielson. 1990. Leafhoppers (Cicadellidae): a Bibliogra- phy, Generic Check-list and Index to the World Literature 1956-1985. CAB Interna- tional Institute of Entomology, Wallingford, UK, [iv] + 368 pp. Rao, K. R. 1980. Studies on cicadellid collections from Manipur (Hemiptera: Cicadellidae). Re- cords of the Zoological Survey of India 76: 189-194. Rodriguez, C. M., J. J. Obando, W. Villalobos, L. Moreira, and C. Rivera. 2001. First report of Xylella fastidiosa infecting coffee in Costa 676 Rica. Plant Disease D-2001-0723-02N (on-line; URL: http://www.apsnet.org/pd/+toc/2001/ dseOltc.htm). Salmon, M. A. 1954. A new variety of Tettigella viridis (L) (Cicadellidae) and other Auchenor- hyncha [sic] from central south Wales. Ento- mologist 87: 64-68. Schréder, H. 1959. Taxonomische und tiergeogra- phische Studien an Neotropischen Zikaden (Cicadellidae, Tettigellinae). Abhandlungen Senckenbergischen NaturfOrchenden Gesell- schaft 499: 1—93. Yang, C. and F. Li. 1980. Notes on Bothrogonia (Homoptera, Cicadellidae) with descriptions of 22 new species from China. Entomotaxonomia 2(3): 191-213. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Yang, M. and Y. Zhang. 2001 [2000]. Two new genera and two new species of Cicadellinae (Homoptera: Cicadellidae) from Yunnan Prov- ince, China. Entomotaxonomia 23(3): 169-174. Young, D. A. 1968. Taxonomic study of the Cicadellinae (Homoptera, Cicadellidae). Part 1. Proconiini. United States National Museum Technical Bulletin 261, 287 pp. . 1977. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New World Cicadellini and the genus Cicadella. North Carolina Agricultural Experiment Station Technical Bulletin 239, 1,135 pp. . 1986. Taxonomic study of the Cicadelli- nae. Part 3. Old World Cicadellini. North Carolina Agricultural Research Service Tech- nical Bulletin 281, vi + 639 pp. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 677-683 ADDITIONS TO THE FAUNA OF DRAPETIS MEIGEN (DIPTERA: EMPIDIDAE) FROM CHINA DING YANG AND PATRICK GROOTAERT (DY) Department of Entomology, China Agricultural University, Beijing 100094, China, and Key Lab of Insect Evolution & Environmental Changes, Capital Normal University, Being 100037, China (e-mail: dyangcau@yahoo.com.cn); (PG) De- partment of Entomology, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, B-1000 Brussels, Belgium (e-mail: grootaert@kbinirsnb.be) Abstract.—The following three new species of the genus Drapetis Meigen (Diptera: Empididae) are described and illustrated: D. digitata, D. elongata, and D. ventralis. An updated key to the species of the genus from China 1s presented. Key Words: Empidoidea, Empididae, Tachydromiinae, Drapetis, China, new species The genus Drapetis Meigen, a member of the subfamily Tachydromiinae (Dip- tera: Empididae), is very similar to Elaphropeza Macquart, but can be sep- arated from the latter by the hind tibia lacking anterodorsal setae and the usu- ally black body (Chvala 1975, Steyskal and Knutson 1981, Yang et al. 2004). The genus is distributed worldwide, and when last cataloged, two species were known from the Oriental Region, en- demic to Taiwan (Smith 1975), three from the Australasian and Oceanian regions (Smith 1989), and 20 from the Palaearctic Region (Chvala and Kovalev 1989). Ten species are presently known to occur in mainland China (Yang et al. 2004). Some other major references dealing with Drapetis include Collin (1961), Kovalev (1972), Chvala (1975), and Rogers (1983). Biological information about species of Drapetis is scant, with general refer- ence to adults being found scurrying on tree trunks (Kovalev 1972), or in rodent burrows and ant nests (Rogers 1983). Bartak and Rohacek (2000) suggested that one Palearctic species, Drapetis infitialis Collin, is a tyrphobiont (strictly dependent on peat bogs). Few papers specifically address the biology of this genus, but Hagler (2002) found a Nearc- tic species near Drapetis divergens Loew to be the most abundant predator of adult whiteflies (Aleyrodidae, specifically Bemisia argentifolii Bellows and Perring) in Arizona cotton fields, providing de- tails of its action as an ambush predator. Hagler (2002) found that in laboratory trials, adults of this species strictly attack adult whiteflies, leaving eggs and nymphs alone even when all stages were abundant. In this paper, three species are de- scribed as new to science, based on specimens collected with sweep nets in Guangdong Province of southern China. An updated key to the species of the genus from China is presented. The holotypes and some paratypes of the new species are deposited in the Entomological Museum of China Agricultural University (CAU), Beijing; some paratypes are also deposited in the Royal Belgian Institute of Natural 678 Sciences (RBINS), Brussels, and _ the National Museum of Natural History, Smithsonian Institution (USNM), Wash- ington, DC. Basic terminology follows McAlpine (1981) and Cumming et al. (1995) for genitalia. The following abbreviations for setae are used: acr-acrostichal, ad- anterodorsal, av-anteroventral, d-dorsal, dc-dorsocentral, np/-notopleural, — oc- ocellar, pd-posterodorsal, ppn-postpro- notal, psa-postalar, pyv-posteroventral, sa-supraalar, sc-scutellar, v-ventral and vt-vertical. UPDATED KEy TO CHINESE SPECIES OF DRAPETIS 1. Thorax yellow, usually with black spots... 9 =e horaxcblack xaevtecs. cca PMs oe, Oe amt 2 2. Hindfemur entirely yellow (Guang- don?) tic Ses Goes 2 are D. nan- lingensis Yang, Gaimari, and Grootaert, 2004 — Hindfemur partly blackish or black ..... 3} 3. Hindfemur with basal third yellow ...... 8 — Hindfemur with basal half yellow 4. Hindtibia blackish (Henan) eg eae D. apiciniger Yang, An, and Gao, 2003 Hindtibia at most with dark brown apex... 5 5. First flagellomere slightly elongated (<3 times longer than wide); right epandrial lobe shallow, at most with shallow cleft on left side — First flagellomere much elongated (>4 times longer than wide); right epandrial lobe wide, with wide cleft on right side... 6 6. Thoracic pleura entirely black; hindfemur with | very long basalmost av; left surstylus narrowing basally and with long hairs on ventral margin (Fig. 9) (Guangdong) EIS GO Ac b POMENE Oh of crate ERS D. ventralis, n. sp. Thoracic pleura partly yellow; hindfemur lacking an elongated basalmost av; left surstylus wide basally and with some short hairs on ventral margin (Fig. 6) (Guang- dong) D. elongata, n. sp. 7. First flagellomere >2.5 (2.7) times longer than wide; left surstylus obtuse apically (Guangdong) Yang, eS ee D. guangdongensis Gaimari, and Grootaert, 2004 First flagellomere 2 times longer than wide; left surstylus with short, fingerlike apex (Fig. 3) (Guangdong) D. digitata, n. sp. 8. Hindtibia with | preapical spine; R4,5 and M convergent apically (Hainan, Taiwan) plas cede, SARE D. nigrispina Saigusa, 1965 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON — Hindtibia without preapical spine; and M divergent apically (Taiwan) ..... EE AES Enon On toto iecs & D. femorata Melander, 1918 9. Mesonotum with black spots ......... 11 — Mesonotum without spots ........... 10 10. Scutellum and postnotum yellow; thoracic pleuron entirely yellow (Yunnan) ...... ME caiobsLEe D. abnormalis Yang and Yang, 1990 — Scutellum brownish yellow, postnotum brown; thoracic pleuron partly brown (Yunnan)... D. obtusa Yang and Yang, 1990 11. Mesonotum with three black spots (Yun- nan)... .D. trimaculata Yang and Yang, 1990 — Mesonotum with two black spots 12. Fore- and midtibiae and tarsi brownish yellow (Guizhou) ie ee eee TBs eee D. maolana Yang and Yang, 1994 — Legs entirely yellow (Yunnan) a Pcs eha i D. kunmingana Yang and Yang, 1990 Drapetis digitata Yang and Grootaert, new species (Figs. 1-3) Diagnosis.—Ist flagellomere slightly elongated (2.0 times longer than wide). Hindfemur with apical half blackish; foretibia dark brown. Left surstylus with short, fingerlike apex. Male.—Body length 2.9mm, wing length 2.9 mm. Head: Black with gray pollen; hairs and setae blackish except lower occiput with pale hairs; eyes nearly contiguous on face; ocellar tubercle with 2 long, strong oc and 4 short posterior hairs; 2 pairs of vt curved inward, outer vi distinctly shorter than inner vf. Antenna blackish; scape bare, shorter than pedi- cel; pedicel with circlet of black apical hairs; Ist flagellomere long, conical, 2.0 times longer than wide, short blackish pubescent; arista long (3.5 times longer than Ist flagellomere), blackish, short blackish pubescent. Proboscis dark brown with blackish hairs; palpus dark brown with blackish hairs. Thorax. Black with thin gray pollen; pleuron subshiny; hairs and setae black- ish; ppn absent; 2 long np/; 1 short sa; 1 long psa; acr multiseriate and not sepa- rated from dc; 1 long dc; scutellum with two pairs of sc (basal pair very short, VOLUME 108, NUMBER 3 679 Figs. 1-9. 1-3, Draptis digitata, male. 1, Genitalia, dorsal view. LC = left cercus; LS = left surstylus; LTL = left epandrial lobe; RC = right cercus; RTL = right epandrial lobe. 2, Right epandrial lobe. 3, Left surstylus. 4-6, Drapetis elongata, male. 4, Genitalia, dorsal view. 5, Right epandrial lobe. 6, Left surstylus. 7-9, Drapetis ventralis, male. 7, Genitalia, dorsal view. 8, Right epandrial lobe. 9, Left surstylus. 680 hairlike, about 1/4 length of apical pair). Legs yellow, except hindfemur with apical half black, tibiae and tarsi brown- ish yellow with foretibia, extreme bases of mid- and hindtibiae, and apex of hindtibia dark brown; hairs and setae brown. Forecoxa with 6 d, apically with 3 setae; midcoxa with 2 outer setae, apically with 3 setae; hindcoxa with 1 outer seta. All femora weakly thickened, subequal in thickness; forefemur with one row of av and one row of py, basalmost pv very long; midfemur with one row of short pv, basalmost pv very long; hindfemur with 5 short ad at base, one row of short av and one row of short pv. Foretibia apically with 1 long pv; midtibia with one row of short spinelike black v, apically with 1 pv; hindtibia apically with | av. Wing hyaline, slightly tinged with grayish; veins dark brown. Calypter dark brown with pale hairs. Halter yellow. Abdomen: Blackish with thin gray pollen; hairs and setae blackish; tergite 1 membranous with linear sclerite; ter- gites 3—S each with group of short black spines laterally, but tergites 4-5 with separated anterior portion weakly scler- otized, short pubescent, and with trans- verse concavity. Male genitalia (Figs. 1— 3): Left epandrial lobe rather narrow basally, its surstylus somewhat quadrate with short, fingerlike apex; right epan- drial lobe fused with its surstylus, much longer than wide, and with wide, obtuse apex; left cercus short and broad with fingerlike apex; right cercus very short, fingerlike. Female.— Unknown. Type material.—Holotype: 3, China, Guangdong Province, Nanling National Nature Reserve, Mangshan, 1,300 m, 8 May 2004, Ding Yang. Deposited in CAU. Etymology.—The name refers to the digitiform tip of the right cercus. Remarks.—The new species is similar to D. guangdongensis from Guangdong, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON but may be separated from the latter by the short’ first flagellomere (2 times longer than wide). In D. guangdongensis, the first flagellomere is 2.7 times longer than wide (Yang et al. 2004). Drapetis elongata Yang and Grootaert, new species (Figs. 4-6) Diagnosis.—Thoracic pleuron partly yellow. Ist flagellomere very elongated (4.3 times longer than wide). Hindfemur with apical half brownish, lacking a very long basalmost av; fore- and midtibiae brown, hindtibia yellow with narrow apex dark brown.. Left surstylus wide basally and with some short hairs on ventral margin. Male.— Body length 3.3 mm. Head: Black with gray pollen; hairs and setae black except lower occiput with pale hairs; eyes nearly contiguous on face; ocellar tubercle with 2 long, strong oc and 4 short posterior hairs; 2 pairs of vt curved inward, outer vf distinctly shorter than inner vt. Antenna yellow with Ist flagellomere dark brown; scape bare, shorter than pedicel; pedicel with circlet of black apical hairs; Ist flagellomere long, conical, 4.3 times longer than wide, short black pubescent; arista long (1.4 times longer than Ist flagellomere), blackish, short black pu- bescent. Proboscis brownish yellow with black hairs; palpus yellow with black hairs and | black apical seta. Thorax: Black with thin gray pollen; mesonotum with yellow anterolateral spot; pleuron subshiny and yellow, but anepisternum with large blackish poste- rior spot, katepisternum with large blackish ventral spot, and metapleuron entirely black; hairs and setae pale; ppn absent; 2 long np/; 1 short sa; 1 long psa; acr multiseriate and not separated from dc; | long de; scutellum with two pairs of sc (basal pair very short, hairlike, about length 3.2mm, wing VOLUME 108, NUMBER 3 Ys length of apical pair). Legs yellow, except hindfemur with apical half brownish, fore- and midtibiae brown, hindtibia with narrow apex dark brown, tarsi brownish yellow; hairs and setae blackish. Forecoxa with 4 d, apically with 3 setae; midcoxa with 2 outer setae, apically with 3 setae; hindcoxa with 1 outer seta. All femora weakly thickened, subequal in thickness; forefemur with one row of av and one row of py, basalmost pv very long; midfemur with one row of pv, basalmost pv very long: hindfemur with 3 short ad at base, one row of av (10-11 av on apical half long and located closely, lacking an elongated basalmost av) and one row of py. Foretibia apically with | long pv; mid tibia with one row of short spinelike black vy, apically with 1 short av; hindti- bia apically with | short thick av. Wing hyaline, slightly tinged with grayish; veins dark brown. Calypter dark brown with blackish hairs. Halter dark brown. Abdomen: Blackish with thin gray pollen; hairs and setae black; tergite 1 membranous with very narrow striplike sclerite; tergites 3—5 enlarged, each with group of short black spines laterally, but tergite 4 with separated anterior portion distinctly sclerotized, short pubescent, and with transverse concavity and tergite 5 similarly but indistinctly sclerotized. Male genitalia (Figs. 4-6): Left epandrial lobe rather narrow basally, left surstylus wide basally and with some short hairs on ventral margin; right epandrial lobe large and more or less fused with large surstylus with small, curved apicolateral process; left cercus very long and large with obtuse apex; right cercus very short, thin, and finger-like. Female.—Unknown. Type material—Holotype: 3, China, Guangdong Province, Ruyuan, Nanling National Nature Reserve, Xiaohuang- shan, 1,500 m, 9 May 2004, Ding Yang (CAV). Paratype: | 3, Nanling National Nature Reserve, Mangshan, 1,500 m, 8 681 May 2004, P. Grootaert (sample 24009, in 70% ethanol, RBINS). Etymology.—The name refers to the distinctly elongated first flagellomere. Remarks.—The new species is very similar to D. ventralis, but it may be separated from D. ventralis by the partly yellow thoracic pleuron, the hind femur lacking an elongated av at its extreme base, and the left surstylus wide basally and with some short hairs on the ventral margin. Drapetis ventralis Yang and Grootaert, new species (Figs. 7—9) Diagnosis.—Ist flagellomere very elongated (4.1 times longer than wide). Fore- and midfemur with apical % brown; hindfemur with apical half black, with basalmost av very long; fore- and midtibiae dark brown, hindtibia yellow with narrow apex dark brown. Left surstylus narrowing basally and with long hairs on ventral margin. Male.—Body length 3.0—3.3 mm, wing length 3.0—3.1 mm. Head: Black with gray pollen; hairs and setae black except lower occiput with pale hairs; eyes nearly contiguous on face; ocellar tubercle with 2 long, strong oc and 4 short posterior hairs; 2 pairs of vt curved inward, outer vt distinctly shorter than inner vt. Antenna dark yellow with Ist flagellomere brown; scape bare, shorter than pedicel; pedicel with circlet of black apical hairs; Ist flagellomere long, conical, 4.1 times longer than wide, short black pubescent; arista long (1.4 times longer than Ist flagellomere), black, short blackish pu- bescent. Proboscis dark brown with black hairs; palpus yellow with blackish hairs and | black apical seta. Thorax: Black with thin gray pollen; pleuron subshiny; hairs and setae pale; ppn absent; 2 long np/; 1 short sa; 1 long psa; acr multiseriate and not separated 682 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from dc; 1 long dc; scutellum with two pairs of sc (basal pair very short, hair- like, about 1/4 length of apical pair). Legs yellow, except fore- and midfemora with apical 1/3 brown, hindfemur with apical half black, fore- and midtibiae dark brown, hindtibia with narrow apex dark brown, tarsi brown except hindtar- sus brownish yellow; hairs and setae blackish. Forecoxa with 4 d, apically with 3 setae; midcoxa with 2 outer setae, apically with 3 setae; hindcoxa with 1 outer seta. All femora weakly thickened, subequal in thickness; forefemur with one frow of ay and one row of py, basalmost pv very long; midfemur with one row of pv, basalmost pv very long; hindfemur with 3 short ad at base, one row of av, basalmost av very long, 5 av on apical “%3 long, and one row of py. Foretibia apically with | long py; mid- tibia with one row of short spinelike black v, apically with 1 short av; hindti- bia apically with | short thick av. Wing hyaline, slightly tinged with grayish; veins dark brown. Calypter dark yellow with brownish yellow hairs. Halter dark brown with dark yellow base. Abdomen: Blackish with thin gray pollen; hairs and setae black: tergite 1 membranous with very narrow and striplike sclerite; tergites 3-5 enlarged, each with group of short black spines laterally, but tergites 4-5 with separated anterior portion weakly sclerotized, short pubescent, and with transverse concavity. Male genitalia (Figs. 7-9): Left epandrial lobe rather narrow basally, left surstylus narrowing basally and with long hairs on ventral margin; right epandrial lobe large and more or less fused with large surstylus with small, curved apicolateral process; left cercus very long and large with obtuse apex; right cercus very short, thin, and fingerlike. Female.—Body length 3.3 mm, wing length 3.0 mm. Type material.—Holotype: 3, China, Guangdong Province, Ruyuan, Nanling National Nature Reserve, Xiaohuang- shan, 1,500 m, 9 May 2004, Ding Yang (CAU). Paratypes: 5 ¢, same data as holotype (CAU); 1 ¢g, 1 ¢, Nanling National Nature Reserve, Mangshan, 1,300 m, 8 May 2004, Ding Yang (USNM); 3 3, 1 ¢, Nanling National Nature Reserve, Xiaohuangshan, 1,500 m, 9 May 2004, P. Grootaert (sample 24012, in 70% ethanol, RBINS). Etymology.—The name refers to the position of the very long anteroventral seta at the extreme base of the hind femur. Remarks.—The new species is very similar to D. elongata, but it may be separated from D. elongata by the entirely black thoracic pleuron, the hind femur having the basalmost av very long, and the left surstylus narrowing basally and with long hairs on the ventral margin. DISCUSSION Thirteen species of the genus Drapetis are now known to occur in China, all of which are distributed in the Oriental part. One species is endemic to the central region of China, five are endemic to the southwestern region, and seven species are endemic to the southern region. ACKNOWLEDGMENTS Our sincere thanks are due to Dr. Stephen D. Gaimari (California Depart- ment of Food and Agriculture, Sacra- mento) and Mrs. Mengqing Wang (Bel- ing) for their help during this study. The research was funded by the National Natural Science Foundation of China (No. 30070100, No. 30225009) and the Cultivation Fund of the Key Scientific and Technical Innovation Project, Minis- try of Education of China (No. 704006). LITERATURE CITED Bartak, M. and J. Rohacek. 2000. The species of the family Hybotidae (Diptera: Empidoidea) of the six peat-bogs in the Sumava Mts. (Czech Republic). Studia Dipterologica 7(1): 61-177. VOLUME 108, NUMBER 3 Chvala, M. 1975. The Tachydromiinae (Dipt. Empi- didae) of Fennoscandia and Denmark. I. Fauna Entomologica Scandinavica, Volume 3. Scandi- navian Science Press, Copenhagen. 336 pp. Chvala, M. and V. G. Kovalev. 1989. Family Hybotidae, pp. 174-227. In Sods, A. and L. Papp, eds. Catalogue of Palaearctic Diptera, Volume 6. Elsevier Science Publishers and Aka- demiai Kiado, Amsterdam and Budapest. 435 pp. Collin, J. E. 1961. Empididae. /n Verrall, G. H., ed. British flies, Vol. 6. Cambridge University Press, London. 782 pp. Cumming, J. M., B. J. Sinclair, and D. M. Wood. 1995. Homology and phylogenetic implications of male genitalia in Diptera—Eremoneura. Entomologica Scandinavica 26: 121-151. Hagler, J. R. 2002. Foraging behavior, host stage selection and gut content analysis of field collected Drapetis nr. divergens: a predatory fly of Bemisia argentifolii. Southwestern Entomologist 27(4): 241-249. Kovalev, V. G. 1972. Diptera of the genera Drapetis Mg. and Crossopalpus Bigot (Diptera, Empididae) from the European part of the USSR. Entomologicheskoe Obozrenie 51: 173-196. (in Russian; English translation, 1972. Entomological Review 51:109-121). McAlpine, J. F. 1981. Morphology and Terminol- ogy—Adults, pp. 9-63. In McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. 683 Vockeroth, and D. M. Wood, coords, eds. Manual of Nearctic Diptera, Vol. 1. Research Branch, Agriculture Canada, Ottawa, Mono- graph 27. 674 pp. Rogers, E. 1983. The Neotropical species of Drapetis Meigen (Diptera: Empididae). Sys- tematic Entomology 8: 431-452. Smith, K. G. V. 1975. Family Empididae, pp. 185-211. Im Delfinado, M. D. and D. E. Hardy, eds. A Catalog of the Diptera of the Oriental Region, Vol. 2. University Press of Hawai, Honolulu. 459 pp. . 1989. Family Empididae. pp. 382-392. In Evenhuis, N. L., ed. Catalog of the Diptera of the Austrasian and Oceanian Regions. Bishop Museum Press and E. J. Brill, Honolulu and Leiden. 1,155 pp. Steyskal, G. € and) L. Vo Kmnutsone 198i: Empididae, pp. 607-624. In McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, coords, eds. Manual of Nearctic Diptera, Vol. 1. Research Branch, Agriculture Canada, Ottawa, Mono- graph 27. 674 pp. Yang, D., S. D. Gaimari, and P. Grootaert. 2004. Review of the species of Drapetis Meigen from China (Diptera: Empidoidea: Tachydromii- nae). Journal of the New York Entomological Society 112(2—3): 106-110. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 684-688 A NEW NEOPHYLAX (TRICHOPTERA: UENOIDAE) FROM MIDDLE TENNESSEE Davip A. ETNIER Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville, TN 37996-1610, U.S.A. (e-mail: dipnet@utk.edu) Abstract.—Neophylax lewisae, n. sp., is described based on adults and larvae from three localities in the Nashville Basin physiographic province of Clay, Cannon, and Williamson counties, Tennessee, and from two localities from the Barrens Plateau (Highland Rim physiographic province) of Moore County, Tennessee. A larva presumably representing the same species is known from an additional locality in Smith County, Tennessee. Males are most similar to those of N. atlanta Ross and N. toshioi Vineyard and Wiggins, differing from these and all nominal Neophylax of eastern North America in having the posterior margin of the inferior appendages sharply keeled in caudal view. Key Words: Trichoptera, Uenoidae, Neophylax, middle Tennessee In May 1997 numerous Neophylax McLachlan larvae were collected from Kelley Creek, Williamson County, Ten- nessee, in the Nashville Basin physio- graphic province. When carefully exam- ined in 1998, several of these were found to differ from those of the known Tennessee species of Neophylax. Suspect- ing that they might represent an unde- scribed species, the site was revisited on 30 September—1 October 1998, and two mature pupae from this collection con- firmed that an undescribed species was involved. Very similar larvae located at two additional Nashville Basin streams and two Barrens Plateau (Highland Rim physiographic — province) Moore County, Tennessee, were sub- sequently reared, and found to represent the same species. METHODS Larvae, prepupae, and pupae were preserved in 70% isopropanol. Adults streams of from the five known localities were obtained by rearing prepupae and pupae collected in late summer or early fall. These were returned to the laboratory in sealed 4-o0z jars filled with creek water and placed in a cooler containing crushed ice. Prepupae and pupae were left in their pebble cases, 1—3 specimens barely cov- ered with stream water per loosely cov- ered 4-o0z jar, and placed in an incubator set to approximate water temperatures at estimated emergence time. Adjusting photoperiod in the incubator was not necessary for successful emergence of a high percentage of specimens. Three additional adults were hand-picked from white sheets suspended vertically near a black light at the Wet Mill Creek site on 29 October 2002. Neophylax lewisae Etnier, new species (Figs. 1—5) Types.—Holotype: Mature ¢@ pupa, Kelley Creek approximately 0.8 km up- VOLUME 108, NUMBER 3 stream (7% mile) from Blowing Spring Branch, Paul Sloan residence, 5565 Tay- lor Cemetery Road, 20.6 air km west- southwest of Franklin, 35°53.368'N, 87°5.741'W, Williamson County, Ten- nessee, 30 September—1 October 1998, J. T. Baxter, deposited in the National Museum of Natural History, Smithso- nian Institution, Washington DC (USNM). Allotype 2, same locality, 29 March 2000, reared by C. R. Parker, emerged 29 September 2000, USNM. Paratypes: Same data as holotype, Uni- versity JO Lenmessee (UT) .7.580, 1 mature 3 pupa; same data as allotype, Royal Ontario Museum (ROM), 1 &, emerged 10 October 2000. Wet Mill Creek at Tennessee Highway 53, BGpzor7 ON, ) 85° 33-.0' W,° Clay - County, Micnmessce. UT 72633... 10” September 2000, preserved 12 October 2000, 1 prepupa, 1 mature ¢ pupa, 1 early ¢ pupa, 3 early 2 pupae; Illinois Natural History Survey (INHS), 1 mature 2 pupa, 1 ¢ pupa; USNM, 29 October 2002 black light, all pinned, 1 cleared 2, 2a al cleared. (Connell Creek ‘at | Nen- nessee Highway 53, 3.9 road km (2.4 rd m1) south of Dekalb County line, 53°29.7'N, 85°33.0’W, Cannon County, @icamessce; 9’ Miay 1999, UT 7.534, 4 prepupae, | each to Clemson University (CUAC), INHS, ROM, and USNM;; 6 September 1999: ROM, | 2, emerged 21 October, 2 mature ¢ pupae, emerged 18 October, 2 November. Additional material.—Spring run trib- utary to Mulherrin Creek at 244 Ten- nessee Highway 53, just north of In- terstate 40, 36°10.83’N, 85°58.46’W, Smith County, Tennessee, 18 April 2000, UT 7.583, 1 prepupa. Shipman Creek at Shipman Creek Road, 0.3 road kena (O22 dim), north of antersection Ledfords Mill Road, 35°24'15.87’N, 86°16'38.55”W, Moore County, Tennes- see, 24 May 1988, UT 7.500, 3 prepupae; same locality, 5 October 2005, UT 7.835, incubated adults emerged 7 October (¢ 685 pupa), 17 October (2 pupa), 29 October (dead @ pupa). Cave Spring Hollow at Tennessee Highway 55, 8.4 road km (5.2 rd mi) southwest of Coffee County line, 35°20'34.70"N, 86°20'52.82”W, Moore County, Tennessee, 10 May 2005, UT 7.777, 6 prepupae; same locality, 5 October 2005, UT ~7:8375 smncubated) Gadulits emerged 11 October through 21 October (5.653: 2.2.2 pupae): Diagnosis.—Males and females con- form to the diagnosis of Neophylax pro- vided in Vineyard and Wiggins (1987). Males are similar to N. aniqua Ross, N. atlanta Ross, N. consimilis Betten, N. oligius Ross, N. securis Vineyard and Wiggins, and N. toshioi Vineyard and Wiggins in having internal branches of segment X elongate, horizontal, and extending noticeably more _ posteriad than other genitalic structures (Fig. 1). Of these species, only N. atlanta and N. toshioi share with N. lewisae the sharply hooked, beak-shaped profile of the in- ferior appendage (Fig. 1). Neither of these species has the inferior appendages with the sharp posterior keel, most notice- able in caudal view (Fig. 2), of N. lewisae. Description.—Male. Length 12.0— 13.2 mm, wing length 10.9-12.0 mm (N = 4). Spurs 1,2,4 in 6 specimens (1,2,3 in ROM @), with medial proximal spur of hind tibia vestigial to 2/3 length of lateral spur; medial distal spur of hind tibia curved, with swollen base, and at mid- length with tuft of filaments on concave surface that converge toward tip of spur (shield of Vineyard et al. 2005). Head, thorax, antennae, legs, and palps brown- ish yellow. Forewing brownish yellow with brown irrorations, outer margin slightly scalloped, yellow areas along posterior margin forming an hourglass from dorsal view when wings folded. Margins of hourglass darker brown. Body and wings covered with yellow setae except setae dark brown margining hourglass, and at wing margin at ends of veins, M1, M2, and from M3 to hind 686 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Xeb Xib Xeb llia mlia Xib 5 Figs. 1-5. Neophylax lewisae. 1, Male genitalia, lateral; 2, Male genitalia, caudal: 3, Male genitalia, dorsal; 4, Male genitalia, ventral; 5, Female genitalia, ventral. Abbreviations: X, segment ten; Xeb, external branch of segment X; Xib, internal branch of segment X; Xifb, inferior branch of segment X; llia, lateral lobe of inferior VOLUME 108, NUMBER 3 angle of wing. Segment VII with sharply pointed ventromedian posterior process that is three times as long as its basal width. Segment IX with shallow, ventral, posteromedian emargination. Male gen- italia (Fig. 1—4) with internal branches of segment X (Xib) long and slender, about twice length of inferior branches (Xifb). Lateral lobes of inferior appendages (Ilia) with beak-shaped ventromesal pro- jection (lateral view), and with sharp posterior keel in caudal view. Median lobes of inferior appendages (mlia) nar- rowly separated on midline, weakly separated from segment IX, and with horizontal, sclerotized, striations anteri- or to ventromesal projection of Ilia. Female: length 12.8-15.2 mm, wing length 11.4-14.0mm (N = 5). Spurs consistently 1, 2, 4, median spurs on hind tibia subequal to lateral partner in length and shape. Coloration as in males, but with fringe of setae margining outer end of wing more uniformly dark. Female genitalia (Fig. 5) with sclerotized outer margins of segment X continuing anteriad to form a sclerotized invagination dorsal to the posterior corners of segment VIII. Posterior margin of VIII sclerotized, and setose on the more heavily sclerotized outer one-third. Posterolateral corners of segment IX not produced posteriad. Accessory lobes of vaginal sclerite paral- lel, rounded on outer margin, straight on inner margin, and in contact with vulval lobe for about half their length. Vaginal sclerite acutely narrowed anteriad (ap- pearing pointed but tip absent or not sclerotized), with wing-shaped lateral projections anterior to vaginal opening. Segment VII with short, blunt, median projection near its posterior margin. Females key to couplet 28, N. toshioi, in Vineyard et al. (2005), but differ from that species in having posterior corners 687 of segments VIII + IX lightly sclerotized, and in having the accessory lobes of the vaginal sclerite as described above (con- vergent at their tips, rounded both laterally and medially, and not in direct contact with the tip of the vulval lobe in N. toshioi). Larva: Legs and head dark brown, head with about 7 pale muscle scars on posterior portion of frontoclypeus and with 2 scars behind inner margin of eye. Two rows of scars under eye, the upper row with 4-5 scars, a gap, and 2-3 additional scars behind eye. The lower row has 2-3 scars, the most anterior of which is slightly anterior to the anterior scar in the upper row. Frontoclypeal tubercle, when present, approximating an equilateral triangle in anterior view, slanted backward, but not sharply point- ed, and rarely conspicuous. Thoracic nota yellow to tan, each pronotal sclerite with about 15 moderately robust, blade- like spines along lateral two-thirds of anterior margin and about 8 smaller spines on medial one-third of margin; about 20 shorter spines are present in a poorly defined second row posterio- dorsal to the marginal row. Each prono- tal sclerite with 40 or more erect black setae. Frontoclypeus covered with micro- spicules. Genae, pronotum, and meso- notum mostly covered with microscales. Microspicules present on genae near eye, on pronotum in midlateral area, and on mesonotum on lateral and_ posterior margins. Ventral gills present on abdom- inal segment 1; lateral gills typically present on posterior portion of segment 2 (2p), anterior portion of segment 3 (3a), and on 4a, with gill on 3p present on at least one side in 3 of 14 larvae, resulting in 4 lateral gills per side (no other Neophylax of eastern North Amer- ican has lateral gills on 3p). Dorsal gills ce appendage; mlia, median lobe of inferior appendage; acc, accessory lobe of vaginal sclerite; vl, vulval lobe of vaginal sclerite; vs, vaginal sclerite; Xinv, sclerotized invaginations of tenth? sternum; VIII, eighth sternum. 688 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON present from 2a through 7p, occasionally absent form 2a, 6a, 7a, and 7p; ventral gills present form 2a through 6p, occa- sionally absent from 2a and 6a, occa- sionally present on 7p. In Vineyard et al (2005), the larva keys to couplet 11 (N. mitchelli Carpenter and N. etnieri Vineyard and Wiggins) if the frontoclypeus is interpreted as having a tubercle. N. /ewisae larvae differ from the former in consistently having lateral gills, from the latter in lacking a pale eye stripe and a pale spot under the fronto- clypeal tubercle and from both in that the frontoclypeal tubercle is absent to tiny and slanted posteriad, not erect. Speci- mens lacking an apparent frontocylpeal tubercle will key to N. consimilis Betten (couplet 16). The very similar larva of this species lacks prominent pale muscle scars on the frontoclypeus, has only about 20 black spines on each pronotal sclerite (40 or more in N. /ewisae), and has 3 or 4 rows of spines covering the thickened anterior border of the frontoclypeus (2 rows, both marginal, in N. /ewisae). In N. lewisae the entire frontoclypeus is covered with mi- crospicules/micronodules, while only the posterior half of that sclerite is spiculate/ nodulate in N. comsimilis. The unusual N. consimilis-like larvae (Vineyard et al. 2005: 50-51) from Wellington Mills, Franklin County, Tennessee (ROM 700336, 700337) very likely represent an additional Barrens Plateau locality for N. lewisae. Variation.—The only variation noted in adults is in segment X in the female, with the margin of the posterior emargi- nation straight-sided (V-shaped) in Nash- ville Basin specimens, and slightly convex in Moore County specimens. Only | of 9 larvae examined from Moore County had a visible frontoclypeal tubercle, and only one of those had a lateral gill on segment 3p. The frontoclypeal tubercle is much more prominent and erect in the single Smith County prepupa (UT 7.583) than in other specimens examined. Discussion.—Nashville Basin streams are low gradient, shallow, and with smooth, horizontal Ordovician limestone substrates interspersed with slabrocks and some gravel. Streams on the Barrens Plateau portion of the Highland Rim physiographic province have - slightly higher gradients, and substrates are primarily chert gravel and cobbles in- terspersed with occasional limestone cobbles and small boulders. We have taken this species syntopically with Neophylax acutus Vineyard & Wiggins and N. fuscus Banks. Etymology.—With pride and _ affec- tion, I name this species in honor of my wife, Elizabeth Lewis Etnier, frequent contributor to my zoological endeavors. ACKNOWLEDGMENTS Charles R. Parker provided rearing facilities for spring-collected prepupae that needed to be held throughout the summer. Patricia W. Schefter was very helpful in providing insights into mor- phology and homologies of the female genitalia, and confirming my guess that this species was not the same as one being described from Alabama in Vine- yard et al. (2005) prior to the appearance of that monograph. J. T. Baxter, Eliza- beth LL Etnier, Oliver S: Rhnit vie and Chris E. Skelton participated in the collection efforts. E. EL Etnicrsandie: D. Hulsey formatted the illustrations for publication. J. K. Moulton graciously provided his copy of Vineyard et al. (2005) while mine was in absentia. LITERATURE CITED Vineyard, R. N. and G. B. Wiggins. 1987. Seven new from North America in the caddisfly genus Neophylax (Trichoptera: Lim- species nephilidae). Annals of the Entomological Society of America 80: 62-73. Vineyard, R. N., G. B. Wiggins, H. E. Frania, and P. W. Schefter. 2005. The caddisfly genus Neophy- lax (Trichoptera: Uenoidae). Royal Ontario Museum Contributions in Science 2, vi + 141p. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 689-725 KEYS TO THE GENERA AND SPECIES OF BLOW FLIES (DIPTERA: CALLIPHORIDAE) OF AMERICA NORTH OF MEXICO TERRY WHITWORTH 2533 Inter Avenue, Puyallup, WA 98372, U.S.A. (e-mail: wpctwbug@aol.com) Abstract.—A Key separating Calliphoridae from similar Diptera families in North America is given. Keys to 17 genera and 54 species of North American calliphorids north of Mexico are provided. The distribution and key characters for each species are discussed and 100 illustrations are included. Calliphora montana Shannon is redescribed. Key Words: Calliphoridae, Diptera, genera, keys, North America, species The most recent revision of North American blow flies was by Hall (1948); however, Hall’s keys have proven to be difficult to use because of his heavy reliance on proportional measurements of characters and the fact that he measured no more than five to ten specimens per species. He also chose specimens representing size extremes rather than “‘average” individuals (Sab- rosky et al. 1989). Subsequently, James (1953, 1955) and Hall and Townsend (1977) provided revised keys, which clarified the identification of regionally selected species. James addressed the western species of blow flies, while Hall and Townsend provided keys to blow flies found in Virginia. Shewell (1987) provided a key to the genera of North American calliphorids, but did not key species. Rognes (1991) reviewed Palearc- tic and Holarctic species and recom- mended numerous changes in blow fly taxonomy. Many of his name changes affected taxa found in North America. Downes (1965) reduced the North American genera Angioneura Brauer and Bergenstamm and Opsodexia Town- send to subgenera under the Palearctic genus Melanomya Rondani. Later Downes (1986) revised species he had placed within Me/anomya describing one new species. Shewell (1987) resurrected Angioneura and Opsodexia as genera. Dear’s (1985) revision of the New World Chrysomyini resulted in Paralucilia wheeleri (Hough) being synonymized with Compsomyiops callipes Bigot, and Chloroprocta fuscanipennis (Macquart) with C. idioidea (Robineau-Desvoidy). Sabrosky et al. (1989) revised the genus Protocalliphora Hough in North America and described 15 new species. Subsequently, Whitworth (2002, 2003a) described’ three additional” “species: Rognes (1985) synonymized the North American Protocalliphora hirudo (Shan- non and Dobrosky) and the Palearctic Trypocalliphora lindneri Peus with Try- pocalliphora braueri (Hendel). Sabrosky et al. (1989) agreed with this, but reduced Trypocalliphora Peus to a subgenus of Protocalliphora. Whitworth (2003b) re- evaluated the status of 7rypocalliphora and agreed with Rognes (1985) that it should be a separate genus. Rognes (1991) combined Hall’s (1948) tribes Phormiini and Chrysomyini under 690 the subfamily Chrysomyinae. Rognes (1991) also proposed the following syn- onymies: Boreellus Aldrich and Shannon = Protophormia Townsend (Chrysomyi- nae); Phaenicia Robineau-Desvoidy, Bu- folucilia Townsend, and Francilia Shan- non =~ Lucilia Robineau-Desvoidy (Luciliinae); Acrophaga Brauer and Ber- genstamm, Acronesia Hall, Aldrichina Townsend, and Eucalliphora Townsend = Calliphora Robineau-Desvoidy and Bellardia agilis (Meigen) = B. vulgaris (Robineau-Desvoidy) — (Calliphorinae). He disagreed with Shewell’s (1987) re- vival of the genus Acrophaga which Zumpt (1956) had synonymized with Calliphora. Shewell had included three species in this genus, genarum, stelviana and the Palearctic suwbalpina. Rognes (1991) retained these species in Calli- phora. He followed Shewell (1987) in retaining Angioneura and Opsodexia (Melanomylinae) as genera. Thus Mela- nomya is a Palearctic genus that does not occur in North America. Angioneura 1s a Holarctic genus represented by five species in North America while Opso- dexia is a Nearctic genus with four species in North America. I have adopted all of Rognes’ (1991) changes. Recently, interest in blow flies has increased, along with studies in forensic entomology. Smith (1986) published keys to adult blow flies of Britain while Greenberg and Kunich (2002) provided keys for Oriental, Australian, South American, and Holarctic species. These keys include some species shared with the Nearctic Region, but omit species found only in North America. The lack of any comprehensive species keys for North American blow flies prompted this study. MATERIALS AND METHODS Specimens for this project were ob- tained from various entomological col- lections throughout North America and my personal collection. A complete list PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of my sources for specimens is under acknowledgments. Characters used in my keys are usually visible with the aid of a quality stereo- microscope and a good light. An ocular micrometer will assist in making pro- portional measurements. It should be noted that fiberoptic lights tend to “wash out” colors, such as yellow and orange, so workers using incandescent lights should consider that when interpreting color characters. Some specimens need to be relaxed to reveal certain characters. Flies were relaxed over wet sand in shallow plastic containers with tight- fitting lids. Most specimens were suffi- ciently relaxed after 48 hours in a relax- ing chamber so they could be handled without damage. Those left too long in the relaxing chamber were susceptible to mold or rot and could be ruined. If removed too soon, they were brittle and prone to breakage. Older specimens often had to be relaxed longer before they could be manipulated safely. The shape of the male genitalia proved useful to confirm species when external char- acters were not distinctive. Male cerci and surstyli were drawn into view following techniques described by Hall (1948). I encountered problems using a bent insect pin, as recommended by Hall, because it tended to flex and sometimes would snap off the genitalia, damaging them. I had better results exposing genitalia using half of a pair of fine point tweezers bent to a 45-degree angle. Rognes (1991) has shown that female terminalia have characters useful to identify female specimens to species, however I did not rely on them in the keys. Terminology differences in the calli- phorid literature can be confusing. For North American terminology workers should see the Manual of Nearctic Diptera (McAlpine 1981), while for European terminology they should refer to the Manual of Palearctic Diptera VOLUME 108, NUMBER 3 (Papp and Darvis 1998). I have primarily followed McAlpine (1981) except as noted below. Figs. |-S from Rognes (1991) detail many of the characters used for blow fly identification. The names of some char- acters vary from common North Amer- ican usage. The following are equivalent terms, with North American terms listed first: postpronotal lobe = humeral cal- lus; postpronotal setae = posthumeral setae (inner and outer); posterior pre- sutural supra-alar seta = presutural seta; propleuron = proepisternal depression; reclinate orbital seta = _ lateroclinate orbital seta (all are shown in Figs. 1, 2). Some changes to older terms are as follows with the preferred term listed second: parafrontal = _ fronto-orbital plate; bucca = genal dilation; third antennal segment = first flagellomere (Figs. 3-5); inner and outer forceps = cerci and surstyli respectively (Figs. 9, 10); and hypopleuron = meron (Fig. 2). Some variation will be noted in spelling of the following, with the pre- ferred spelling given second: acrostical = acrostichal, and intraalar = intra-alar. For hyphenated species names such as terrae-novae, a species of both Calliphora and Protophormia, the hyphen is dropped, as a result of a ruling by the International Commission on Zoologi- cal Nomenclature (ICZN 1999, Article 3235, 2.29%): Several useful taxonomic characters are available on the wings (Fig. 6), abdomen (Figs. 7, 8) and_ genitalia (males Figs. 9, 10, females Figs. 11—13). Important characters which are species specific are illustrated separately. The ratio of head to frons widths are used throughout keys, see Figs. 23, 24 for how to measure. The average ratio is followed by the range and the total number of specimens measured. Historically many terms have been used to describe the hairs and fine dusting observed in adult flies. I use the 691 following convention: macrotrichia are larger hairs with nerves and sockets, microtrichia are cuticular extensions or dusting without sockets. Macrotrichia can be described as setae, setulae, hairs, or bristles. I will avoid the terms hairs and bristles and consider larger macro- trichia as setae and smaller macrotrichia as setulae. The term vestiture sometimes is used to describe patterns of macro- trichia. Microtrichia patterns have been called dusting, pubescence, pollinosity, microtomentum, or microtrichia. For purposes of this publication, the term microtomentum is used to describe this condition. The scientific names used herein fol- low Rognes (1991). Where possible, characters used are readily observed with a good microscope and without dissec- tion. The first character listed in a couplet is generally the most distinctive; char- acters listed after may not be as reliable or may be more difficult to distinguish. See Table 1 for a list of species im the order they are addressed and the names used by Hall (1948). The only synonyms given are for Hall’s publication. This is not a complete list of synonyms for each species. Partial keys to adult species of Proto- calliphora are provided to be integrated with existing keys in Sabrosky et al. (1989). The keys include 3 new species I have described (Whitworth 2002, 2003a). I also have added revised illustrations and information to assist in the separa- tion of species of this genus. Species keys are not provided for Melanodexia, Opsodexia, or Angioneura. Both Hall (1948) and James (1955) provided species keys for Melanodexia, but they are difficult to use and in any case the genus needs revision, a_ task which is beyond the scope of this study. Downes (1986) provided keys to species of Opsodexia and Angioneura which are effective (N. Woodley, in litt.), but few specimens were available for this study 692 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON greater ampulla presutural seta lesser ampulla lower calypter notopleuron upper calypter presutural area acrostichal setae stem-vein dorsocentral setae anepisternal setae humeral callus humeral setae anepisternum \ (c= =) (ezee transverse suture posthumeral setae notopleuron presutural seta notopleural seta humeral setae prealar seta. intraalar setae numeral callus mediotergite postsutural area- supraalar setae anterior spiracle anatergite proepisternal la J anepimeron depression a katatergite el haltere (1 posterior spiracle « SAL IVY postalar callus. postalar setae proepisternal setae proepisternum . scutellum prebasal scutellar seta proepimeral seta } F I/ meral setae basal scutellar seta j katepimeron lateral scutellar setae yy 4 metakatepisternum subapical scutellar seta proepimeron p Cc} front coxa discal scutellar seta apical scutellar seta katepisternai setae AR Ch katepisternum metasternal area 2 inner venical seta laterociinate orbital seta ocellar seta inner vertical seta \ prociinate orbital setae Vs Tas outer vertical seta outer vertical seta lateroclinate orbital seta postocular setae fronto-orbytal plate i postocular setae : i Uh) . Proclinate orbital ee v y as A = setae Whe scorn fronto-orbital plate =a intrapostocular cilia frontal seta YE frontal vitta parafacial Scape pedicel first lagellomere genal groove funula supravibrissal setulae paratacial - posigena vibnssa Ptilinal suture a genal setae arista subvibnissal setulae facial ridge pew facial plate genal dilation first flagellomere subvibrissal setulae lower facial margin 4 prementum vibrissa humeral crossvein inner vertical seta occipital setulae outer vertical seta postocular setae stem-vein lateroclinate outs orbital seta prociinate orbital setae ocellar seta A,+CuA, CuA, Figs. 1-6. Calliphora subalpina (Ringdahl, 1931). 1, Dorsal view of thorax. 2, Left lateral view of thorax. 3, Left lateral view of head. 4, Anterior view of head. 5, Dorsal view of head (from Rognes 1991). 6, Trypocalliphora braueri, dorsal view of wing; inset, portion of costa showing setulae on underside (from Rognes 1991). VOLUME 108, NUMBER 3 and species keys are therefore not in- cluded. SEPARATING FAMILIES Most calliphorids are readily distin- guished from other families by their metallic blue, green, or bronze color and the relatively large size of adults. Metallic muscids and tachinids are fre- quently found under Calliphoridae in collections because of these shared char- acters. Metallic muscids are readily separated from calliphorids by the ab- sence of a row of setae on the meron. Metallic tachinids can be distinguished by the prominent subscutellum and bare arista. The non-metallic calliphorid genus Pollenia Robineau-Desvoidy is common in North America. It can be recognized by a row of setae on the meron and an abundance of silky, crinkly hairs on the thorax. Other non-metallic calliphorids include the relatively rare Angioneura, Opsodexia, and Melanodexia which are more or less dull colored. Characters provided in the key should distinguish these genera. Non-metallic calliphorids are often found in collections with similar looking muscids, sarcophagids, and tachinids. Key TO SEPARATE CALLIPHORIDS FROM SIMILAR FAMILIES 1. Meron without row of setae, sometimes scattered weak setulae. .... Muscidae, Anthomyidae, Scathophagidae — Meron with distinct row of setae (Figs. 2, IG) Reese aR RE Scene Pee ne ero See 2. Subscutellum strongly developed; arista often bare (not in Dexiini and some other taxa). Tachinidae — Subscutellum absent or weak; arista usually setose (Fig. 3) [except Angioneura and most Miltogramminae (Sarcophagidae)]. ..... 3 Abdomen, and usually thorax, shining metallic blue, green, cr bronze luster, sometimes with darker vittae (e.g., Cochlio- most Calliphoridae — Abdomen and thorax dull gray, brown, or black, never shining metallic.......... 4 No ios) myia). Nw 693 Thorax with silky, crinkly, yellowish setae along with regular dark setae; proe- pisternal depression bare (Fig. 2)...... Calliphoridae, in part (Pollenia) Thorax without silky, crinkly, yellowish setae along with regular dark setae; proe- pisternal depression setose (Fig. 15) or bare (Melanodexia). Scutum with conspicuous — black stripes on a gray to gold background; dorsum of abdomen checkered dark and light; notopleuron usually with two large and two smaller setae. ..... Sarcophagidae Scutum without black stripes; dorsum of abdomen usually not checkered; noto- pleuron usually with only 2 setae (as in Fig. 1 ) (except Trypocalliphora). Coxopleural streak absent. Calli- phoridae, in part (Melanomyinae, 2 genera, 9 species) Coxopleural streak present (Fig. 16). ... 7 Middle of proepisternal depression bare or with a few sparse setae; posterior thoracic three spiracle small.-= . . Soe seas eaee el = . Calliphoridae in part (Melanodexia, 8 species) Middle of proepisternal depression setose (Fig. 15), posterior thoracic spiracle larger (as in Fig. 16)... .... Rhinophoridae (Bezzimyia), and some Sarcophagidae (Miltogramminae) Key To SUBFAMILIES AND GENERA OF CALLIPHORIDAE Basal section of stem vein setose above (Hicao (ehrysomyimnac eee eee 2 Basal section of stem vein bare above (Biss NA) Oe eo Gees oe ge en a 9 Greater ampulla with stiff erect setae (Fig. 17); dorsum of first and second abdominal tergites black, posterior margins of abdominal tergites 3 and 4_ black (Fig. 18). Chrysomya (2 spp.) Greater ampulla bare or with fine, short setulae (Fig. 2); color pattern not as above. 3 Genal dilation orange with mostly yellow setae; head with predomi- nantly yellow vestiture; posterior margin of yellow or hindicoxassctOScsmeieie nena ween ee 4 Genal dilation usually black with dark setae; head with predominantly black vestiture; posterior margin of hind coxa baneror withinweake setulacisi iene 6 Mesonotum without distinct dark vittae: lower calypter bare (as in Fig. 14); wing clouded along C (Fig. 19); parafacial bare; cerci, surstyli and other characters illustrat- ed in Dear (1985: figs. 7-11); tropical, rare in Texas. Chloroprocta (1 sp.) 694 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sT1 T1+2 abdominal marginal abdominal setae a8 spiracle —-discal abdominal setae epandrium if TST7+8 cercus surstylus bacilliform sclerite in \ & Mitrrrgcenmercee™ spsq rg tg 7 ie humeral callus epipret cerc 11 anterior tegula basicosta costa thoracic spiracle proepisternal posterior spiracle haltere base depression abdomen greater upper ampulla calypter lower calypter 47 ° coxopleural % streak 16 meral setae Figs. 7-17. 7-8, Pollenia rudis. 7, Dorsal view of abdomen. 8, Ventral view of abdomen (from Rognes 1991). 9-10, Lucilia magnicornis, male postabdomen. 9, Posterior view. 10, Left lateral view (from Rognes 1991). 11-13, Calliphora stelviana, female postabdomen. 11, Dorsal view. 12, Ventral view. 13, Left lateral = spiracle; st = view (from Shewell 1987). cere = cercus; epipret = epiproct; hypret = hypoproct; spr VOLUME 108, NUMBER 3 695 Mesonotum with distinct dark vittae; lower abdomen bronze-green; frons to head ratio calypter with upper half setose; wing not 0.22 (0.21—0.25/25) (see Fig. 24 for how to clouded along C (Fig. 6); parafacial with measure). Puparia appear bare, with sparse pale or dark setulae (may be inconspicu- spines; prothoracic fringe absent; larvae are OlIS) Segoe. eee Se ad « SE 5 obligate subcutaneous parasites of nestling Setae on posterior margin of hind coxa TSE. ces sents hs a Trypocalliphora (1 sp.) pale: palp filiform (Fig. 20); calypter whit- ish; Bend in M closer to wing margin than dm-cu crossvein (as in Fig. 42). Three species, one widespread (C. macellaria). . . Se adopeleok ayssee ee fia toe a 6 Cochliomyia (4 spp.) Setae on posterior margin of hind coxa long and dark; palp clavate (as in Fig. 3); calypter brown; bend in M closer to dm-cu crossvein or about equal distance between crossvein and wing margin (as in Fig. 44). Found primarily in the southwestern Compsomyiops (callipes)(1 spp.) Presutural acrostichal seta weak or absent; upper calypter with black setae; body Two notopleural setae (as in Fig. 1); calyp- ter usually whitish, if brown, other char- acters vary; male frons usually broader, fronto-orbital plates well separated, frons at narrowest 0.06 or more head width (0.06— 0.16, one species 0.34), surstylus, cercus, and aedeagus variable. Female, with laterocli- nate orbital setae present (Figs. 3—5); thorax bluish (female Protocalliphora aenea and some P. interrupta have an aeneous thorax); frons to head ratio 0.24 or more (0.25—0.35). Puparia usually heavily spined; distinct prothoracic fringe (250—800 ut in diameter); normally an ectoparasite, sometimes metallic, shining very dark blue green found in nestling nares, ears, or at the Without microtmehia.,.. sav... 2 wes aeaeu « base of feather sheaths of nesting 4 ae ean Routt on: ROE Protophormia (2 spp.) birds® =e eee ehOLocdlliphnoral(2&8ispp») Presutural acrostichal seta moderate to 9. Middle of proepisternal depression bare, or strong; upper calypter bare or with pale if setose, then body dull black, subshining, setae (Fig. 21); body metallic with whitish Nowmetallicibiucionercentr stem 10 MUCKOLOMIEM CUM sare Cees Ge ee 7 — Miiddle of proepisternal depression setose Two postsutural intra-alar setae; anterior (Fig. 15); body shining metallic blue green, thoracic spiracle with bright orange setae; or bronze, sheen sometimes dulled by anterior acrostichal seta moderate; scutum (PMC ONS MWIIN, oocno scons oasodsee 13 convex centrally. Scavenger species, not 10. Gena usually half height of eye or more PALASItIGweerwaeee otk coe Phormia (1 sp.) (Fig. 25); coxopleural streak present (as in Three or four postsutural intra-alar setae; Fig. 16); parafacial setose to lower eye anterior thoracic spiracle usually with white margin; facial carina usually — present to dark brown setae; anterior acrostichal (mice 6) sollentna cin ieee eee aes 11 seta strong; scutum often more or less — Gena about one fourth height of eye flattened centrally. Obligate parasites of (Fig. 27); coxopleural streak absent; paraf- Mes ulmMmomoIMNASepem merce ce sha seca oe 8 acial bare on lower half or more; facial One or more accessory notopleural setae carina absent. Melanomyinae ........ 12 between the usual anterior and posterior 11. Thorax with long, crinkly yellowish setae; notopleural seta (Fig. 22 ); calypter yellow- preapical posterodorsal seta on hind tibia ish to brown; frons of male narrow, at absentsonweake jee eee Pollenia (6 spp.) narrowest 0.05 (0.04—0.07/25) head width — Thorax without long, crinkly yellowish (see Fig. 23 for how to measure); fronto- setae; preapical posterodorsal seta on hind orbital plates touching, or nearly so; tibia present, almost as long as preapical surstylus, cercus, and aedeagus distinctive dorsal seta (Fig. 28). .. Melanodexia (8 spp.) (See Figs. in Sabrosky et al. 1989: 272, 273). 12. Arista plumose (as in Fig. 27). Apical Female lateroclinate orbital setae absent (see Figs. 3-5 for location); thorax & scutellansetacslonyo(Bicm29) sae any een RE nT NS, is, Caos eke Opsodexia (4 spp.) sternite; tg = tergite.14, Lucilia sericata, dorsal view of wing base showing suprasquamal ridge (spsq rg) (from Shewell 1987). 15, Diagrammatic left lateral view of anterior portion of thorax, showing setose proepisternal depression. 16, Lucilia coeruleiviridis, left lateral view of posterior thoracic spiracle and coxopleural streak. 17, Chrysomya rufifacies, left lateral view of wing base showing setose greater ampulla. 696 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON S BpeS Son FETT NSNS) ; OS \y humeral callus anterior notopleural accessory notopleural posterior notopleural 22 —— inner vertical seta lateroclinate orbital seta outer vertical seta HW VOLUME 108, NUMBER 3 — Arista with fine pubescence. Apical scutellar setae usually short (Fig. 30). ER PAE cs t5. 1 SN A Nn, es Angioneura (5 spp.) 13. Thorax and abdomen shining green, blue, or bronze. ridge with conspicuous cluster of setae near Suprasquamal the base of scutellum (Fig. 14); lower calypter bare above (Fig. 14). Lucilii- TASS) seh fees, tar ale aon Se ec Lucilia (11 spp.) — Thorax dull, microtomentose; abdomen usually metallic blue with whitish micro- tomentum. Suprasquamal ridge bare or with inconspicuous fine setae (Fig. 31); lower calypter setose above (Fig. 31). Cal- liphorinae 14. Bend of M obtuse (as in Fig. 32), curvature of apical section even; first flagellomere, at most, twice the length of pedicel; costa usually setulose below only to junction with subcosta, as in inset Fig. 6; abdomen blue or olive green; known only from northeastern North America in the Nearctic Region. Jae OG Ed One oR ES Bellardia (2 spp.) — Bend of M acute or right angled (Fig. 42), curvature of apical section greatest just beyond bend: first flagellomere more than twice length of pedicel; costa usually setulose below to junction with R1; abdo- men bluish. 15. Upper and lower calypter white........ 16 — Upper and lower calypter light to dark brown, margin may be white.......... avo aly oes eae Calliphora, in part (11 spp.) 16. Presutural intra-alar seta absent; abdomen shining, no microtomentum visible when viewed posteriorly. Cynomya (2 spp.) — Presutural intra-alar seta present (Fig. 1), abdomen microtomentose when viewed POSteKIOnl Ae eee = aecdens Ss ee 17 17. Orange basicosta; abdomen elongate, lon- ger than length of dorsum of thorax; abdomen with light microtomentum when viewed from rear. California to Washing- ton, Colorado to Alberta, usually at higher CISVALLONS Sey arlene eet ae Cyanus (1 sp.) — Black basicosta; abdomen no longer than dorsum of thorax; abdomen with heavy microtomentum when viewed from rear. 697 Northern Canada, Alaska or high elevation only. Calliphora, in part (2 spp.) CALLIPHORINAE This subfamily includes Bellardia, Cal- liphora, Cyanus, and Cynomya. It can be recognized by the following characters: stem vein bare above; lower calypter setose above; proepisternal depression setose; thorax dull, microtomentose; abdomen more or less shining blue; suprasquamal ridge bare or with only a few inconspicuous setae. Bellardia Robineau-Desvoidy, 1863 This Palearctic genus is a recent 1m- migrant to North America and known only from the northeastern U.S. It was very rare in my search of collections. The genus can be identified by the obtuse bend in vein M (Fig. 32). The species are believed to be earthworm parasites. They are the only North American blow flies which are viviparous (Shewell, 1987). Shewell noted that the terminalia of females are very short; he provided two illustrations (figs. 38,39) and he labeled them Bellardia agilis (Meigen) which is a synonym of B. vulgaris (Robineau- Desvoidy). Key TO SPECIES OF BELLARDIA 1. Upper parafacial with dark brown spots which do not disappear when viewed from above: lower calypter evenly darkened, light tan; male genitalia tiny, cercus longer than surstylus as in Rognes (1991: figs. 79, SO) Fa Gmeallltive eos oslo. bees 6 ue aes — Upper parafacial without spots; lower calypter white, margins yellowish or light tan; male genitalia larger, cercus shorter than surstylus as in Rognes (1991: figs. 68, 69); a larger fly. vulgaris bayeri Figs. 18-24. 18, Chrysomya megacephala, dorsal view of abdominal, tergites. 19, Chloroprocta idioidea, dorsal view of wing (from Shewell 1987). 20, Cochliomyia macellaria, left lateral view of head (from Shewell 1987). plp = palp; lbl = labellum. 21, Phormia regina, dorsal view of right wing base (from Shewell 1987). 22, Trypocalliphora braueri, dorsal view of prothorax. 23, Measuring male head to frons ratio: fw = frons width at narrowest; hw = head width at widest. 24, Measuring female head to frons ratio: fw = frons width at narrowest; hw = head width at narrowest. 698 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Species in order they are discussed and comparison of names used in the current paper with names used by Hall (1948). Table 1. Continued. Whitworth (2006) Hall (1948) Page Number Eel Page Number Whitworth (2006) Hall (1948) CALLIPHORINAE Bellardia Not included 698 bayeri Bellardia Not included 698 vulgaris Calliphora Acronesia alaskensis 700 alaskensis Calliphora Acronesia aldrichia 701 aldrichia Calliphora Calliphora 701 coloradensis coloradensis Calliphora Acronesia collini, 701 genarum A. popoffana Calliphora Aldrichina grahami 703 grahami Calliphora Eucalliphora arta, 703 latifrons E. lilaea Calliphora Calliphora livida 703 livida Calliphora Calliphora mortica 703 loewi Calliphora Acronesia montana 703 montana Calliphora Acronesia abina, 704 stelviana A. anana Calliphora Calliphora vicina 707 vicina Calliphora Calliphora 707 terraenovae terrae-novae Calliphora Calliphora vomitoria 707 vomitoria Cyanus Cyanus elongata 707 elongata Cynomya Cynomyopsis 709 cadaverina cadaverina Cynomya Cynomya mortuorum, 709 mortuorum C. hirta CHRYSOMYINAE Chloroprocta Chloroprocta 709 idioidea idioidea C. fuscanipennis Chrysomya Not included 709 megacephala Chrysomya Not included 710 rufifacies Cochliomyia Callitroga aldrichi 710 aldrichi Cochliomyia Callitroga americana 710 hominivorax Cochliomyia Callitroga macellaria G2 macellaria Cochliomyia Callitroga minima WNP munima Compsomyiops — Paralucilia wheeleri 712 callipes Phormia regina | Phormia regina a2 Protocalliphora — Apaulina 712 Protophormia Boreellus atriceps ING atriceps Protophormia Protophormia WAT terraenovae terrae-novae Trypocalliphora — Apaulina hirudo 718 braueri LUCILIINAE Lucilia cluvia Phaenicia cluvia 720 Lucilia Phaenicia 720 coeruleiviridis caeruleiviridis Lucilia cuprina— Phaenicia pallescens 720 Lucilia elongata Bufolucilia elongata 720 Lucilia eximia Phaenicia eximia 720 Lucilia illustris — Lucilia illustris TI Lucilia Francilia alaskensis 721) magnicornis Lucilia mexicana Phaenicia mexicana 721 Lucilia sericata Phaenicia sericata 721 Lucilia silvarum Bufolucilia silvarum 721 Lucilia thatuna — Phaenicia thatuna 721 POLLENIINAE Melanodexia Melanodexia 721 Melanodexiopsis Pollenia Not included 722 angustigena Pollenia Not included 122 griseotomentosa Pollenia labialis Not included 123 Pollenia Not included 723 pediculata Pollenia rudis Pollenia rudis 723 Pollenia Not included 723 vagabunda MELANOMYINAE Angioneura Not included 723 Opsodexia Not included 123 Bellardia bayeri (Jacentkovsky, 1937). I examined specimens from Strafford Co., New Hampshire and Middlesex Co., Massachusetts. This species has dark calypters, and if generic characters are miussed, it will tend to key 710 Calliphora terraenovae. Bellardia vulgaris (Robineau-Desvoidy, 1830). I examined a single specimen collected from Moorestown, New Jersey. Shewell (1987) reported it only from VOLUME 108, NUMBER 3 New Jersey. This ‘species has pale calypters. Specimens key to Calliphora genarum or C. stelviana if generic char- acters are missed. Calliphora Robineau-Desvoidy, 1830 This species, genus now _ has 13) INearctic since Rognes (1991) suggested the following generic names are syno- nyms: Acronesia, Acrophaga, Aldrichina, and Eucalliphora. Species have the stem vein bare, a dull, microtomentose thorax, and bluish abdomen with whitish micro- tomentum. Key TO SPECIES OF CALLIPHORA 1. Presutural intra-alar seta absent; anterior thoracic spiracle with orange setae; abdo- men blue or dark green with white micro- tomentum; male epandrium large, cerci curve sharply under abdomen with horn- like prominences at each base (Fig. 33). Western, Alaska to California and Color- — Presutural intra-alar seta present (Fig. 1); anterior thoracic spiracle usually with brown setae; abdomen usually metallic bluish with or without white microtomen- tum; cerci and surstyli not as above (as in Nw Calypter wholly white; frons of male broad, at narrowest, usually 0.15—0.21 head width; primarily northern Canada and Alaska or hichvelevationseesier = aris area | Glee — Calypter brown, margin often white; frons of male narrower, at narrowest, usually 0.14 head width or less (C. /atifrons averages 0.24, C. coloradensis averages 0.15); usually not restricted to northern or highvelevationyancasse-6 ai sree oe 3. Arista with short setae above, very short below (Fig. 35); parafacial with dark chest- nut or black ground color; broad undusted stripe between presutural acrostichals usu- ally extending past transverse suture; third abdominal tergite with long median mar- ginal setae, usually more than half the length of those on the fourth tergite; tip of surstylus rounded (see Rognes_ 1991: fig.150). Alaska, northern Canada, to @Quebecrandulcabradorase yee — Arista with long setae above and below (Fig. 36); parafacial orange in ground color on lower half or more; undusted stripe faint, usually narrower and stopping at the grahami eS) genarum 699 transverse suture; third abdominal tergite with shorter median marginal setae, always less than half the length of those on fourth tergite; tip of surstylus pointed (see Rognes 1991: fig. 173). Alaska, Quebec, high elevations in) Colorado = 4.446 4. stelviana Facial ridge with row of short, stout, supravibrissal setae, ascending from the vibrissae to a point almost halfway to antennal base (Fig. 37a); a second set of strong divergent ocellar setae about 2/3 the length of the anterior ocellars, surrounded by only a few sparse setae (Fig. 37a). Male genitalia shorter, with a chisel-shaped point (Fig. 38). Frons of male broad, at narrow- est, almost 2 width of parafacial at lunule, frons 0.24(0.22—0.26)/12 head width; female frons 0.37(0.36—0.39)/8 head width. Primarily western, east to Colorado AnGe\WWASCONSIN ae Ene een Laine latifrons Facial ridge with row of slender supravi- brissal setae (Fig. 37b); second set of ocellar setae weak or absent, if stronger (females of some species), surrounded by dense fine setae; male genitalia usually longer and more slender, as in Fig. 34a; frons of male narrower, at narrowest, equal to or less than width of parafacial at lunule, frons 0.15 head width or less; frons of fenlalesvanialal Came ranean iene nee 5 Three postsutural intra-alar setae...... 6 Two postsutural intra-alar setae (as in ie 1), if Genal dilation reddish; frons of male broad, broader than width of parafacial at lunule, 0.14 (0.12—0.16)/11 times head width: lower portion of surstylus and fifth abdominal segment with dense wavy setae (Figs. 39a, b). Alaska to Ontario and Indiana, south to Mexico. coloradensis Genal dilation, when fully colored, black; frons of male much narrower, less than half width of parafacial at lunule, 0.06 (0.05— 0.07)/7 times head width; lower portion of surstylus and fifth abdominal segment with sparse, straighter setae (Figs. 40a, b). Widespread. livida Basicosta yellow to orange; genal dilation with reddish ground color on anterior half or more; frons of male, at narrowest, 0.075 (0.07—-0.08)/4 head width. Wide- SPreadiey7s ek. Ret wena ae eee vicina Basicosta dark brown or black; genal dilation, when fully colored, usually black (except front half often reddish in C. terraenovae); frons of male, at narrowest, usually less than 0.07 head width....... 8 Postgena and lower posterior corner of genal dilation and back of head with long 700 10. 1] PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON yellow-orange setae, sometimes extending forward along edge of subgena and lower genal dilation (Fig. 41); vestiture of the occiput below postocular setae primarily pale setae; genal groove reddish or orange. Frons of male, at narrowest, 0.044 (0.04— 0.05)/10 head width; long, slender, curved surstyli (Figs. 34a, b); frons of female at narrowest 0.34 (0.31—0.35)/10 head width. Widespreadteen otha 5st meal vomitoria Postgena and genal dilation with mostly dark or black setae, back of head and rear edge of postgena may have yellow setae; vestiture of the occiput below postocular setae with three or more rows of black setae; genal groove usually black or dark brown (except C. terraenovae); other char- ACLENS Waly teats Ste Ruts SUNAERe OENaeE Bend in M much closer to wing margin than length of M from cross vein dm-cu to bend (as in Fig. 42 ); usually 4 (3—5) strong lateral scutellar setae besides apical pair (Fig. 43); male frons, at narrowest, 0.04— 0.07 head width; surstylus curves anteriorly Onusystralchie (igses438 459) hs eer = eee Bend in M about equal distance between wing margin and dm-cu cross vein (varies from slightly closer to wing margin to slightly closer to cross vein) (Fig. 44 shows the latter condition); usually 2 or 3 (occasionally a fourth on one side) strong lateral scutellar setae besides apical pair (Fig. 45); male frons, at narrowest, 0.06— 0.14 head width; surstylus curves posteri- Onlya@bigw4 Ga). 2 ee ees hal Ss oe Genal groove usually reddish or orange; anterior 1/3—1/2 of genal dilation usually reddish when viewed from above; first flagellomere often orange along lower in- side edge; parafacial golden or silvery tan when viewed from above; surstylus long and slender with gentle anterior curve (Fig. 47). Widespread from Alaska _ to Newfoundland, south to southern Califor- nia, and Texas; usually at higher elevations 10 im the Wiest.) G20. een we eae terraenovae Genal groove usually black or dark brown (rarely reddish or reddish brown); anterior portion of genal dilation entirely black or dark brown; first flagellomere uniformly gray; parafacial silvery black; surstylus not as above (Fig. 48). ... 6 Wrath. Katie .Surstylus long, straight, parallel-sided ta- pering to an obtuse point (Fig. 48a); cerci appearing slender when viewed posteriorly (Fig. 48b): frons of male, at narrowest, 0.043(0.035—0.055)/12 head width; fronto- orbital plates touching at narrowest; fifth abdominal tergite of female with posterior incision 1/3—1/2 length of segment (Fig. 49), lateral profile of tergite often tent-like. Alaska, British Columbia, Yukon Terri- LORS cas. he Fee oad ne loewi — Surstylus curved anteriorly at tips (Fig. 50a); cerci pear shaped when viewed posteriorly (Fig. 50b); frons of male wider, at narrowest, 0.06 (0.05—0.07)/ 16 head width, fronto-orbital plates not touching; fifth abdominal tergite of female with short incision, or at most only a slight indentation (Figs. 5la, b), profile not tent- like. Widespread from Alaska to Oregon and east to North Carolina, pri- marily at high elevations in southern locationSs yeu eee alaskensis 12. Two or 3 lateral scutellar setae in addition to apical pair (Fig. 45). When a third seta is present, in the prebasal position (see Fig. 1), it is usually weak. Bend in M vein usually farther from wing margin and closer to crossvein dm-cu (as in Fig. 44). Sometimes the bend is equal distance between wing margin and crossvein. Male frons, at narrowest, 0.11 (0.08—0.14)/15 head width. Cercus of male usually shorter (Fig. 46b). Western south from Alaska through British Columbia to California and*@olorado: Mase ae = ener aldrichia — Three or sometimes 4 lateral scutellar setae in addition to the apical pair, the seta in the prebasal position usually stronger (as in Fig. 43). Bend in M vein usually slightly closer to wing margin or equal distance between wing margin and dm-cu crossvein (similar to Fig. 42, but the bend is shown much closer to the wing margin). Male frons at narrowest, 0.07 (0.06—0.08)/18 head width. Cercus of male usually longer (Fig. 52b). Primarily east of the Rockies, ranging southeast from northwestern Ca- nada (where it overlaps with C. aldrichia) east through the Canadian provinces to Ontariovand babradore - eee montana Calliphora alaskensis Shannon, 1923. This species is widespread but rare, found only at high elevations in the southern portions of its range. Hall (1948) listed specimens from Alaska, Wyoming and Colorado. I also found this species from 7 locations in Canada (3 in British Columbia and 4 in Quebec), from 10 locations in the US. (2 in Oregon, 2 in Utah, 41n Colorado and, surprisingly, one each from mountains in Tennessee and North Carolina). This VOLUME 108, NUMBER 3 species is normally rare, but I examined about 25 females of this species that were bait trapped in the vicinity of Vancouver, British Columbia. These flies were at- tracted to both beef and chicken liver in second growth timber (K. Needham, in litt.). In 7 of 50 specimens examined (6 males, 1 female), the genal groove was reddish to reddish brown which would place them with C. terraenovae. Genitalia will separate males, but females with reddish genal grooves will be difficult to separate. Calliphora aldrichia Shannon, 1923. Hall (1948) gave records from British Columbia and Quebec, Alaska, Colora- do, Wyoming, Montana, and Washing- ton. (I believe the Quebec record was likely C. Montana). 1 also examined specimens from western Alberta, Cali- fornia, and Oregon. This species is morphologically very close to C. mon- tana, the two appear to be sibling species. Males share distinctive surstyli which curve posteriorly (Fig. 46a), unlike other Calliphora. The most obvious difference between the two species is that males of C. aldrichia have, on average, a much wider frons (0.11 of head width at narrowest) than C. montana (0.07 of head width, at narrowest). The term lateral scutellar setae is used in the key in a broad sense to include all stronger setae on the margin of the scutellum other than the apical pair (as in Figs. 43, 45). Other authors assign separate names to these setae, as in Fig. 1. The two species appear to have formed as a result of geographical isolation. Calliphora aldrichia is found west of the Rocky Mountains from Alaska to Cali- fornia and Colorado, while C. montana is found primarily east of the Rockies from Northwest Territories and Alberta east to Labrador. Their ranges overlap in northern British Columbia, and southern Yukon where specimens with intermedi- ate characters were found. Since both species are associated with mountains or 701 northern latitudes the mechanism of isolation is unclear. It appears that C. aldrichia is associated with higher eleva- tions while C. montana is found through lower elevations to the east. The area Where the species converge is lower elevation at the northern edge of the Rockies. Separating females of these species, based on morphology, will be problematic. Outside the zone where populations converge, distribution ap- pears to be the best way to separate females. As more specimens become available, species distinctions should be reevaluated. Calliphora_ coloradensis Hough, 1899. This species is generally rare, but ap- peared to be locally abundant in areas around Flagstaff, Arizona, and Uvalde, Texas. I also examined specimens from California, New Mexico, Oregon, Wyoming, and South Dakota. Hall (1948) reported its range from Mexico north to Alaska, and east to Ontario and Indiana. Most specimens that I examined were from the southwestern U.S. This species has 3 postsutural intra-alar setae, a character it shares only with C. /ivida and some C. J/atifrons. However, it has a reddish genal dilation which separates it from C. /ivida. The character is good in fully sclerotized specimens, but can be confusing in teneral individuals, which are fairly common in this species. Calliphora genarum Zetterstedt, 1838. This species and Calliphora_ stelviana (Brewer and Bergenstamm, 1891) would key to Acrophaga in Shewell (1987). I saw few specimens of this species. Hall (1948) gave its range as Alaska and northern Canada, Manitoba, and Lab- rador. I examined specimens from Yu- kon, Northwest Territories, and Mani- toba. It shares white calypters with C. stelviana, which separates both species from other Calliphora. The differences in seta length on the arista are used to separate these two species from each other, but the fact that the setae are often 702 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Obtuse Angle 32 EPS . Ni PESOS, 2S = NN epandrium “ays : eS | calyp “gd L AB idema 67a ) aS “Ta cemaaueanene sh. of ES gia ‘, 67b a 68a 711 64 MZ PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON outside row of frontal setae, versus pale setulae in C. macellaria. Some specimens have pale setulae mixed with dark in the lower frontal plate. If any dark setulae are present, the specimen is C. homini- vorax. Cochliomyia macellaria — (Fabricius, 1775). This is the most common Cochlio- myia in North America, from the south- ern U.S. to southern Canada. In good specimens this species can be readily identified by the presence of pale setulae outside the row of frontal setae, and pronounced silvery microtomentum on the lateral areas of the fifth tergite. These characters may be difficult to see in old or damaged specimens. For females, the yellowish basicosta is distinctive. The number of proclinate orbital setae 1s variable, in a group of 16 females, 11 had two on each side while 5 had only one on each side. Male genitalia are illustrated in Hall (1948: figs.18 A, B). Cochliomyia minima Shannon, 1926. Dear (1985) identified two females from the Florida Keys, one was from Key West and one was from Stock Island. I examined many Cochliomyia from the Keys and never found this species. Dear (1985) also listed this species from Cuba, the Dominican Republic, Jamaica, Puerto Rico, and the Virgin Islands. Male specimens are readily separated from the similar Cochliomyia aldrichi by the broader frons and distinctive genita- lia. Characters for females are reliable for good specimens, but they are easily damaged and problematic in poor speci- mens. The pattern of microtomentum on the fifth tergite is sometimes readily visible but is somewhat subjective in many specimens. The color of setulae on the occiput can be difficult to in- terpret. Some C. aldrichi have only a few dark setulae to separate them from C. minima with all pale setulae. Compsomyiops Townsend, 1918 A single species. Compsomyiops callipes (Bigot, 1877) is found primarily in the southwestern U.S. I examined specimens from California, Arizona, New Mexico and Texas. It can be separated from Cochliomyia by the clavate palps, long dark setae on the hind coxa, and dark calypter. It is a large bluish fly. Male genitalia are illustrated in Hall (1948: figs. 19A—D); female ovipostitor illustrated in Dear (1985: figs. 47, 48). Phormia Robineau-Desvoidy, 1830 A single species. Phormia regina (Meigen, 1826) is very common throughout North America. It is a shining metallic blue or green fly with bright orange setae around the anterior thoracic spiracle. Characters are illustrated by Rognes (1991: figs. 247-258). Protocalliphora Hough, 1899 Protocalliphora is a large genus with 28 species known in North America. It 1s most diverse in temperate regions of the Intermountain West, less common far- ther south. Sixteen species are found only in the West, 6 only in the East, while 6 are widespread in both areas. This genus has been found in 46 of the lower 48 states and Alaska, but it has not been recorded from Florida or Louisiana. It is uncommon in collections, but common in the nests of many altricial birds. Figs. 69-73. 69, Lucilia mexicana male, cerci and surstyli; posterior view. 70, L. eximia male, cerci and surstyli; a) left lateral view; b) posterior view. 71, L. thatuna female, antenna, left lateral view. 72, L. thatuna male, cerci and surstyli; a) left lateral view; b) posterior view. 73, L. sericata, posterior view of head showing setae below inner vertical setae, left side; L. sericata, L. cuprina. VOLUME 108, NUMBER 3 70b 72b L. sericata L. cuprina 13 714 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Species of this genus are bird nest parasites whose larvae suck the blood of nestling birds. Characters include 3 or 4 postsutural intra-alar setae, 2 noto- pleural setae, strong anterior acrosti- chals, scutum usually flattened on center, puparium usually heavily spined, with a strong prothoracic fringe. This genus is closest to Trypocalliphora. The keys to species relying on adult and puparial Protocalliphora in Sabrosky et al. (1989) work well for reared series with matched males, females, and pu- paria for the 26 North American species known at the time of publication (77ry- pocalliphora braueri was included under Protocalliphora). The key to males per- mits the identification of lone males in good condition and the key to females permits the identification of about 15 species of lone females in good condi- tion. For males, the shape of the surstyli is a critical character and some of the sketches provided in Sabrosky et al. (1989) are misleading. I have redrawn the surstyli for P. beameri Sabrosky, Bennett, and Whitworth, 1989; P. bicolor Sabrosky, Bennett, and Whitworth, 1989; P. hirundo Shannon and Do- broscky, 1924; P. interrupta Sabrosky, Bennett, and Whitworth, 1989; P. me- tallica Townsend, 1919, and P. parorum Sabrosky, Bennett, and Whitworth, 1989 (Figs. 54-59) to better reflect distinctions for each species. Lone females are often difficult to identify because they have few distinctive characters. Perhaps a de- tailed study of female genitalia will produce some distinguishing characters in the future, but preliminary examina- tions have not provided any good characters. Since the publication of Sabrosky et al. (1989), I have identified three addi- tional North American species of Proto- calliphora (Whitworth 2002, 2003a). The former publication describes two new species (P. bennetti Whitworth and P. rugosa Whitworth), the latter splits P. sialia into an eastern and western com- ponent. Protocalliphora sialia Shannon and Dobroscky is the form found in the Midwest and East while P. occidentalis Whitworth is the western form. I have provided a key to assist with identification of these new species and it can be integrated with the adult and puparial key in Sabrosky et al. (1989) starting at couplet 7, p. 77. The unifying character in this group is the digitate surstyli in males. Lone adult females will be difficult to key, but the key is useful to separate females of species in mixed infestations in bird nests. Common mixes in the west included P. bennetti, P. occidentalis, P. rugosa, and occasionally P. hirundo. \n the east P. sialia and P. bennetti are commonly found in the same nest. Adults in this genus are difficult to collect, though their empty puparia are relatively easy to find in old bird nests. I developed a revised key (Whitworth 2003b) for the puparia of 27 North American species. To date I have exam- ined over 8000 bird nests, about half of which were infested with one or more of 27 of the 28 known species of this genus. One species, P. sapphira Hall, has not been collected from a nest and is known primarily from a single distinctive male. Three females matched to the male may not be the same species. I have examined many Protocalliphora from the same area in Alaska where P. sapphira was collect- ed and have found nothing resembling the male holotype. Until recently, P. beameri had never been collected from a bird nest. However, in 2004 I received an adult P. beameri and matched pupar- ium from a black -throated gray warbler nest. The specimens were provided by Piotr Jablonski, who found the nest in the Chiricahua Mountains in Arizona. The puparium matches those of a pre- viously unidentified species I examined from a barn swallow nest near Ft. Davis, west Texas. VOLUME 108, NUMBER 3 gis humeral setae ~ setulae head os 75 inner posthumeral extra humeral setae humeral setae outer posthumeral seta notopleuron presutural seta ‘ presutural intra-alar 80 79a 79b Figs. 74-81. 74, Lucilia cuprina, setae at rear of humeral callus and notopleuron. 75, L. sericata, setae at rear of humeral callus and notopleuron. 76, Pollenia vagabunda, dorsal view of left side of prothorax. 77, P. vagabunda male, cerci and surstyli, posterior view. 78, P. rudis, dorsal view of left side of prothorax. 79, P. griseotomentosa male, cerci and surstyli; a) left lateral view; b) posterior view. 80, P. pediculata, ventral view of junction of humeral crossvein and subcosta showing bundle of pale setulae. 81, P. griseotomentosa, dorsal view of left side of thorax. 716 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON d av pv Vv Orientation of Setae on Legs 82b a anterior ad —_anterodorsal av anteroventral d dorsal p posterior pd _ posterodorsal py _ posteroventral 83 tergite sternite 84 Pollenia rudis male, cerci and surstyli, a) left lateral view; b) posterior view. 83, Figs. 82-84. 82, Diagrammatic sketch of seta orientation on legs: a = anterior; ad = anterodorsal; av = anteroventral; d = dorsal; p = posterior; pd = posterodorsal; py = posteroventral; v = ventral. 84, Ventral view of vestiture on abdomen; P. angustigena, left side, P. rudis, right side. Key TO MALE PROTOCALLIPHORA WITH DIGITATE SURSTYLI, AND WHITE CALYPTERS, WITH NOTES ON FEMALES AND PUPARIA swallow and phoebe nests. Usually in birds UNO OPM MVS, 56 050 a g0coGacoo sue P. cuprina, P. halli, P. hesperia, P. hesperioides Male and female with postalar wall and tympanic pit bare or with a few pale setae 1. Male surstylus digitate (as in Fig. 56), not see Sabrosky et al. (1989: fig. 3b) for appreciably curved. - 4.0 42.5.) 4s 2 location; fore tibia usually with one poste- Male surstylus distinctly curved (Figs. 54, KEY GLE I rcts epoucuoun (leone Gln avowed! Hc 4 55, 57-59), usually slender, or short and Male and female with postalar wall and broad as in Sabrosky et al. (1989: figs. 7 tympanic pit with a conspicuous tuft of OV sey acta 15 species of Protocalliphora black setae; fore tibia with two posterior nN Calypter white in both sexes, primarily parasites of birds which nest in cavities. . 3 Calypter brown, except calypter white in female P. cuprina and P. halli. Female P. cuprina have fifth tergite cupreous, female P. halli are found almost exclusively in barn setae. Found almost exclusively in bank swallow nests. Male and female preocellar area polished just anterior to median ocellus, P. rognesi with polished area varies from small to large and irregular (see Sabrosky et al.-1989: figs. 1,2a VOLUME 108, NUMBER 3 for location); frons of female, at narrowest. averages 0.25(0.22—0.28) head width. Dor- sum of puparium with cuticular ridges faint or absent, or if pronounced, found only east of a line from Alaska to Kentucky. Male and female with preocellar area dull colored, microtomentose (rarely a small polished area); frons of female, at narrow- est, averages 0.28(0.26—0.31) head width. Dorsum of puparium with pronounced cuticular ridges (see Whitworth 2003b for an explanation of puparial characters). Found primarily in the west, one species, P. hirundo, may be found in the east primarily in bank and cliff swallow nests. Male and female with parafacial relatively broad, width at lunule obviously much wider than width of first flagellomere; male frons wider, at narrowest, averaging 0.075— 0.10 head width. Puparium with hyperstig- matal spines longer, averaging 25 u or more; posterior ventral spine bands not TREGIUIGEG! (10): IERIE, © ors tho tae oie che ioc toeare Male and female with parafacial narrow, equal to or barely wider than first flagello- mere; male frons narrower, at narrowest, averaging 0.06(0.05—0.07) head width. Pu- parium with hyperstigmatal spines short, averaging 12.5 u in length; posterior ven- tral spine bands reduced to rear... . Male frons, at narrowest, 0.075(0.065—0.08) head width; about equal to width of first flagellomere; female with large, triangular polished area encompassing ocellar triangle which tapers to a point in preocellar area when viewed from below. Puparium with shorter prothoracic fringe averaging 350 u; dorsal cuticular folds faint. Alaska to northern Idaho, east to northern Minne- sota and southeast to Virginia.... . Male frons broader, at narrowest, 0.10(0.09—0.12) head width; frons of male clearly wider than first flagellomere; female with smaller irregular polished preocellar area, not encompassing ocellar triangle, or if extending upward, not uniformly shining when viewed from below. Puparium with exceptionally long prothoracic fringe, 500 ut or more in diameter; dorsal cuticular folds pronounced. East of a line from Alaska through Saskatchewan and Minne- SOLastONKeentuckyAreeiee ere nels n an Basicosta orange to reddish brown .... . Basicosta black or dark brown. Upper portion of fronto-orbital plate in male significantly narrowed as in Whit- worth (2002: fig. 7). Dorsal cuticular ridges of puparium narrower, less than 50 tt wide, Nn shannonti bennetti sialia 8 occidentalis WANG ridges abundant and close together. Pri- marily in tree, and violet-green swallows, and purple martins. rugosa — Upper portion of fronto-orbital plate in male not significantly narrowed as in Whitworth (2202: fig. 8). Dorsal cuticular ridges of puparium broader, 50 u or more, ridges usually widely spaced. Primarily in cliff and bank swallows. hirundo Protophormia Townsend, 1908 This genus is represented by only two species in North America. Both have a flattened scutum, like Protocalliphora, but the anterior acrostichals are weak or absent. KEY TO SPECIES OF PROTOPHORMIA 1. Lower part of face strongly protruding (Fig. 60); arista almost bare below (Fig. 60); antenna entirely black; two pairs of marginal scutellar setae in addition to the apical pair; anterior spiracle much enlarged (Fig. 61), almost as large as humeral callus in lateral view; cell r4+5 closed, or nearly so at wing margin (Fig. 62); eye small, about two-thirds of head height (Fig. 60). Rare, found only north of 80°N. — Lower part of face not strongly protrud- ing (as in Fig. 3); arista plumose (as in Fig. 3); tip of pedicel and basal part of first flagellomere reddish; 3-4 pairs of marginal scutellar setae in addition to the apical pair; anterior spiracle smaller, much smaller than humeral callus (as in Fig. 2); cell r4+5 open at wing margin; eye larger, three-fourths of head height. Com- mon in the northern U.S., Canada and Alaska. terraenovae atriceps Protophormia atriceps (Zetterstedt, 1845). This is a rare species found north of 80°N in North America (Rognes 1991). It can be recognized by its pro- truding lower face (Fig. 60) and large anterior spiracle (Fig. 61). Various char- acters are illustrated in Rognes (1991: figs: 31,313; 3:115—326): Protophormia terraenovae (Robineau- Desvoidy, 1830). This species is common throughout the northern U.S., Canada, and Alaska. I examined specimens from Washington to Ohio and Alaska to 718 California. The face is not protruding and it has a smaller anterior spiracle. Various characters are illustrated in Rogsnes (1991: fies, 310,312,314, 327— 337). Trypocalliphora Peus 1960 Rognes (1985) considered Trypocalli- phora a valid genus, while Sabrosky et al. (1989) considered it a subgenus of Pro- tocalliphora. As a result of my studies of puparia (Whitworth 2003b), I concluded that Trypocalliphora deserves generic status. It is represented by a single Holarctic species. Trypocalliphora braueri (Hendel, 1901) Widespread, but uncommon throughout most of the U.S., Canada, and Alaska; relatively common in the Northwest (Whitworth, 2003b). Closest to Proto- calliphora, this species has one or more accessory notopleural setae (Fig. 22). Larvae are obligate subcutaneous para- sites of nesting birds. Puparia have very few spines and lack a prothoracic fringe. Various characters are illustrated in Rognes (1991: figs. 338-349). LUCILIINAE This subfamily includes one genus, Lucilia Robineau-Desvoidy, 1830. The genera Phaenicia, Bufolucilia, and Fran- cilia were synonymized with Lucilia by Rognes (1991). It can be recognized by its shining, green, blue or bronze thorax and abdomen, suprasquamal ridge with a cluster of setae, and bare lower calypter. The genus _ includes 11 species in North America. When measuring the head to frons ratios in females, note that the frons is not narrowest at the vertex as in most female calliphorids. Lucilia Robineau-Desvoidy, 1830. Key TO LuciLiAa SPECIES |. Subcostal sclerite on venter of wing with wiry black setulae (Fig. 63); basicosta tan, Nw Ww dark brown or black; palp orange; surstylus and cercus of male as in Fig. 64; ocellar triangle of female large, reaching at least halfway to lunule. Widespread in_ the northern U.S. and Canada. Subcostal sclerite on venter of wing with pubescence only (Fig. 65); basicosta orange or black; palp orange or black; surstylus and cercus of male not as above; ocellar triangle of female small, not reaching haltiwayatorliniules see -foeasns eee Palp black or brown; length of head at level of lunule more than half head height (see Fig. 66 for how to measure); third abdom- inal tergite with | or 2 pairs of long, erect median marginal setae (see Fig. 7 for location); basicosta dark brown or black. Palp orange to yellow, not darkened apically; length of head at level of lunule less than half head height (except in L. thatuna and some L. sericata); third ab- dominal tergite with marginal setae not especially strong or erect (except male L. thatuna); basicosta usually yellow or orange (L. mexicana and L. eximia have brown basicostas):| bs, 9. Se eee eee Three postsutural intra-alar setae with anterior one weak; presutural intra-alar setae absent; arista with short setae, usually much shorter than width of first flagellomere as in Rognes (1991: fig. 411); first flagellomere long, more than _ half eye length in_ profile; male cercus parallel-sided, tip of surstylus straight (Figs. 9, 10). Northern, Alaska to Labra- Two postsutural intra-alar setae; presutural intra-alar setae present; arista normal, with setae longer than width of first flagello- mere; first flagellomere shorter, less than half eye length in profile; male cercus Y- shaped when viewed from rear, tip of surstylus with sharp bend to rear (Figs. 67, 68). Widespread. Two postsutural acrostichal setae, occa- sionally 3 on one side; male frons, at narrowest, 0.13/4 (0.12—0.14) head width; surstylus triangular in lateral view; cerci with short inverted V shape in posterior view (Fig. 67b); fifth sternite of male prominent, as long as fifth tergite; female frons, at narrowest, 0.35/5 (0.33— 0.36) head width. Western only, rare in California, Colorado, Oregon, and Washinstons2swk oo es eee Three postsutural acrostichal setae; male frons, at narrowest, 0.07/6 (0.07—0.09) head width; surstylus slender in lateral view; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON illustris magnicornis elongata VOLUME 108, NUMBER 3 cercl longer inverted V in posterior view (Fig. 68b). Fifth sternite of male shorter than fifth tergite. Female frons, at narrow- est, 0.32/5 (0.32-0.34) head width. Wide- spread, common. Two postsutural acrostichal setae; abdo- men usually uniformly metallic or micro- WOMMMAM Ss cs Go oe OOS o one he cS Three fostsutural acrostichal setae; abdo- men with apparent mesal division in which one half is microtomentose, the other half is shining (except not in L. thatuna). Basicosta usually black or dark brown, sometimes lighter ground color, but always with dusky shading; mostly southern dis- (Hier SU KOVEN, AE Ae tlle! Ons hore takees Gee cietonal boserens Basicosta yellow or orange; primarily eastern. Two or more complete rows of black postocular setae (Figs. 3-5 for location): genal dilation and _ parafacial mostly black, with black vestiture; frontal vitta wider in male, frontal plates separated, frons, at narrowest, 0.055 (0.05—0.06)/8 head width; distal end of cercus Y-shaped when viewed from rear, surstylus with sparse wavy setae (Fig. 69); female frons broader, 0.28(0.26—0.30)/6 head width, at narrowest. Primarily southwestern U.S. into Mexico. One complete row of black postocular setae; genal dilation and parafacial mostly tan to orange, with vestiture reddish to light brown; frontal vitta in male very narrow, frontal plates touching, or nearly so, frons, at narrowest, 0.035 (0.03—0.04)/10 head width; distal end of cercus almost parallel when viewed from rear, surstylus with dense wavy setae (Fig. 70); female frons narrower 0.25 (0.24—0.28)/9 head width, at narrowest. Subtropical, occasion- ally found in Texas and Florida. Frons of male with frontal plates almost touching, frons width, at narrowest, much less than breadth of first flagellomere, frons 0.023/8 (0.015—0.030) head width; male with one lateroclinate orbital seta slightly anterior to median ocellus (see Figs. 3—5 for seta location); female with black setulae outside row of frontal setae on frontal plate; fifth abdominal tergite highly po- lished, tinged with red or purple in both sexes; mature specimens usually larger, 8.0— 9.5mm in length. Maryland south to Florida, north to Michigan and Wisconsin, most common in the southeast, less abun- dant west of the Mississippi River, but westward to California...... silvarum mexicana eximia coeruleiviridis 10. Frons of male with frontal plates well separated, frons width, at narrowest, more than width of first flagellomere, frons 0.11/ 7 (0O.10—0.12) head width: male with later- oclinate orbital seta opposite median ocel- lus, or seta absent; female with pale setulae outside row of frontal setae on frontal plate; fifth abdominal tergite generally not more polished than other tergites, usually without reddish or purple cast; mature specimens usually smaller, 8.0 mm length or less. Florida north to North Carolina and west to southern Mississippi. ... . First flagellomere broader than width of parafacial at level of lunule, often cupped inward, inner margin often reddish, espe- cially in female (Fig. 71), sometimes not so cupped in male, still broader than parafa- cial; frons of male, at narrowest, with frontal plates almost touching, frons 0.044 (0.04—0.05)/9 of head width: male surstylus and cercus as in Fig. 72; frons of female, at narrowest, 0.30/12 (0.27-0.32). Rare, known only from California, Colo- rado, Idaho, Oregon, Utah, and Washing- LUO) CURRY Rr oe Dest Sever operas pty a of nels First flagellomere narrower than width of parafacial at level of lunule, usually not cupped or with reddish margin (similar to Fig. 36); frons of male, at narrowest, with frontal plates widely separated, frons 0.12— 0.21 head width; frons of female, at narrowest, 0.35—0.40 of head width. Central occipital area with single seta below inner vertical seta (Fig. 73, right side); metasternum bare; abdomen dull coppery; humeral callus with 2 or 3 small setulae along posterior margin; noto- pleuron with only 2 or 3 small setulae on posterior border (Fig. 74); frons of male broader, at narrowest obviously much broader than width of the parafacial at level of the lunule, 0.20/7 (0.19—0.21) of head width; frons of female at narrowest 0.39/5(0.38—0.40) head width. Southern, Virginia west through Missouri to Califor- Hideey.. eee: See Central occipital area with group of 2-5 setae below inner vertical seta (Fig. 73, left side); metasternum setose; abdomen usual- ly bright green, occasionally shining cop- pery; humeral callus with 6—8 small setulae along posterior margin; notopleuron usu- ally with 5 or more setulae on rear border (Fig. 75); frons of male narrower, at narrowest about equal to width of parafa- cial at level of lunule, 0.13/6(0.12—0.14) of head width; frons of female at narrowest IN? cluvia thatuna 10 cuprina 720 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0.37/8(0.35—0.40) head width. Wide- Spo red Clidlah eae) noes ecae enna eee sericata Lucilia cluvia (Walker, 1849). This species is found primarily in the South- east; I examined specimens from Arkan- sas to Florida to South Carolina. It is uncommon and very close to L. coer- uleiviridis in appearance, but L. cluvia males can be distinguished by their much broader frons. Females of L. cluvia are difficult to separate with confidence from those of L. coeruleiviridis, the primary distinction being the color of the fine setulae outside the row of frontal bris- tles. This character is often variable, damaged, or difficult to see. The differ- ence in the shininess of the fifth tergite is subjective and variable, but can be useful with practice. Based on the material I examined L. cluvia tends to be smaller than L. coeruleiviridis. Excluding 3 obvi- ously undersized specimens, 20 speci- mens of the former were 7.5—8 mm in length. For L. coeruleiviridis, excluding 5 undersized specimens, 51 ranged from 7.75—9.5 mm in length. Better characters are needed for females distinctions, perhaps a study of ovipositors would reveal useful characters to separate the two species. Lucilia coeruleiviridis Macquart, 1855. This species is in the southeastern U.S., but it may be found in the Northeast and Midwest. It is uncommon in the West, and is generally much more commonly encountered than L. c/uvia. | examined specimens from California to Florida, from Nebraska and Wisconsin to Penn- sylvania and most states south. I did not find it in the Northwest. Lucilia cuprina Wiedemann, 1826. This species is uncommon throughout the south, from Virginia to Florida west to Missouri and Texas and California. It is usually recognized by its dull coppery sheen, but color alone is not reliable. Some L. sericata are quite coppery though usually more shining. The wider frons in L. cuprina readily separates males of each species. A single seta below the inner vertical seta (Fig. 73, right side) versus 2—5 setae in L. sericata (Fig. 73, left side) will distinguish speci- mens of both sexes. This character sometimes varies, or can be hard to see due to the condition of the specimen. The presence or absence of setae on the metasternum (absent in L. cuprina) 1s also useful, but often is difficult to see. Lucilia elongata Shannon, 1924. This species is rarely found in collections. I examined specimens from California, including San Mateo County, Mendo- cino County, Tehama County, and Yolo County; Washington, Pierce County; Oregon, Washington County, and Kla- math County. James (1955) recorded this species from Orcas Island, Washington, as well as California and possibly Colo- rado. This species is close to L. si/varum, but it normally has only 2 postsutural acrostichal setae, while L. si/varum has 3. The frons of males, at narrowest, is much broader, averaging 0.13 of head width in L. elongata versus 0.07 in L. silvarum. A few females and one male were seen with 2 setae on one side and 3 on the other. J. O’Hara (in litt.) reports the Canadian National Collection has 15 specimens of L. elongata, 6 from various areas in British Columbia have 2. postsutural acrostichals each side, 9 from Whatcom County, Washington, include several males with 3 postsutural acrostichals on one side. Lucilia eximia (Wiedemann, 1819). This species is rare, found occasionally only in Texas and Florida. Hall (1948) noted that this is a common fly in market places of Central America. This is one of only 2 species with yellow palps and a dark basicosta. It has only one row of black postocular setae, and an orange genal dilation. By contrast, its close relative, L. mexicana, has two complete rows of postocular setae and a dark genal dilation. The postocular seta char- VOLUME 108, NUMBER 3 acter can be confusing, the row of black postocular setae may be incomplete in L. eximida. Lucilia illustris (Meigen, 1826). This species is widespread and common in the northern U.S. and Canada. I examined specimens from as far south as southern California and Arizona in the West, but in the Midwest I did not find it south of Missouri to Indiana, while on the east coast I did not find it south of South Carolina. Various characters are illustrat- ed in Rognes (1991: figs. 371, 411-422). Lucilia magnicornis (Siebke, 1863). This species 1s uncommon in the far north arom!) Alaska ‘to Labrador. I examined specimens from Alaska, Northwest Territories, and northern Manitoba. This species has brown palps, 3 postsutural intra-alar setae, and the length of the first flagellomere is more than half the eye length. Various char- acters are illustrated in Rognes (1991: figs. 371. 411-422). Lucilia mexicana Macquart, 1843. This species is common in the south- western U.S. I examined specimens from California to Texas, Utah and Oklahoma. Hall (1948) stated that this species extends as far south as Brazil. It has a brown basicosta, like L. eximia, but two complete rows of postocular setae and a dark genal dilation. Its range overlaps with that of L. eximia in Texas. Lucilia sericata (Meigen, 1826). This species is one of the most common Lucilia, and is widespread in the U.S. and southern Canada. It is one of 3 species with 3 postsutural setae. It can be separated from L. cuprina by the pres- ence of 2—S setae on the central occipital area below the inner vertical setae. Specimens tend to be green, but some are so coppery that they can be confused with L. cuprina. It also has a setose metasternum, which is often hidden and very difficult to see. This species can be separated from L. thatuna by the width 72M of the first flagellomere and the much broader frons of the male. Various characters are illustrated in Rognes (1991: figs. 375, 455=465). Lucilia silvarum (Meigen, 1826). I examined specimens from Washington to California in the West and Maine, south to South Carolina and Louisiana in the East. A common, widespread species, Hall (1948) also recorded it from southern Canada. Specimens have 3 postacrostichal setae and black palps, the male frons is much broader than in the similar L. e/ongata. Various charac- ters are illustrated in Rognes (1991: 376, 466-476). Lucilia thatuna Shannon, 1926. This is an uncommon species; I examined speci- mens from 10 counties in California, most in the northern coastal areas; also Pullman, Washington; Baker Co. Ore- gon; and Cache Co. Utah. James (1955) recorded it from many localities in California, and also Montana, Idaho, and Colorado. The presence of three postacrostichal setae and the first flagel- lomere being broader than the parafa- cials separate it from L. cuprina and L. sericata. Specimens are often bluish which separates them from the green or coppery L. sericata or the coppery L. cuprina. Males are distinctive as their frons is much narrower than those of L. cuprina and L. sericata. POLLENIINAE Species in this subfamily are dull colored unlike most calliphorids and there are two genera, Me/anodexia and Pollenia. Melanodexia Williston, 1893 This genus is uncommon in the West; few specimens were encountered in this study and no attempt was made to sort them to individual species. Both Hall (1948) and James (1955) studied this genus, but species distinctions are diffi- (22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cult and the genus needs further study. Hall listed three species under this genus and five more under the name Melano- dexiopsis, a synonym of Melanodexia. Pollenia Robineau-Desvoidy, 1830 This genus is widespread in North America. It was thought to be represent- ed by a single species, P. rudis, until recently (Rognes 1997). Six species are now recognized in North America. Spe- cies of this genus are dull colored calliphorids with distinctive, crinkly yel- low setae on the thorax. The key herein was adapted from the one developed by Knut Rognes for Greenberg (1998). Key TO SPECIES OF POLLENIA 1. Thorax with dark median undusted vitta between presutural acrostichal setae, usu- ally extending forward to extreme anterior slope of thorax and back to the rear of the scutum; usually 2 or more extra setae in front of the regular row of 3 (2-4) humeral setae (Fig. 76); 2 inner posthumeral setae (Fig. 76); male cercus broad and flattened when viewed from rear (Fig. 77). Northeastern and northwestern U.S. and P. vagabunda No median undusted vitta; no extra hu- meral setae in front of the regular setae (Fig. 78); 1 inner posthumeral seta (Fig. 78); cercus not as above (Fig. 79b). 2. Lappets of posterior thoracic spiracle dark brown; facial carina reduced and indistinct; basicosta usually dark brown to black. Northern U.S. and southeastern Canadas Merce see a see see P. labialis — Lappets of posterior thoracic spiracle pale yellow to orange; facial carina usually distinct (Fig. 26) (except P. griseotomen- tosa); basicosta yellowish, orange, or light (aT abe aektir cic Ren REL A ra iit oel it uae ese Node at junction of humeral crossvein and subcosta of wing with a bundle of pale setulae below (Fig. 80); palpus dark brown or black. Widespread in northern portions Oth eRUES ens sen re P. pediculata — Node without setulae; palpus usually ligh- ter brown or orange (except some P. rudis). 4 4. Outer posthumeral seta absent (Fig. 81): femur of mid- and hind leg with mostly black vestiture on posteroventral surface (see Fig. 83 for orientation); facial carina absent or much reduced; male surstyli NO eS) distinctly curved and slender (Fig. 79a); frons of male exceptionally narrow, at narrowest, 0.032 (0.025—0.04)/3 head width. Rare, northeastern and northwestern U.S. ands Wisconsin re P. griseotomentosa — Outer posthumeral seta usually present (Fig. 78); if absent (some P. angustigena) femur of mid- and hind leg with mostly yellow vestiture on posteroventral surface; facial carina distinct (Fig. 27); male surstyli less curved and broader (Fig. 82); frons of male slightly too much broader......... 5 5. Tibia of midleg with one anterodorsal seta (see Fig. 83 for orientation); mid- and hind femora with yellow or orange posteroven- tral vestiture; ventral abdominal vestiture in males normal, not particularly fine, dense, or erect (Fig. 84, left side); male frons narrower, frons 0.038 (0.03—0.04)/6 head width. Locally common in Washing- ton, California, Wisconsin, and northeast- emp Ues: P. angustigena Tibia of midleg usually with 2-3 antero- dorsal setae; mid- and hind femora with black or dark posteroventral vestiture; ventral abdominal vestiture in males fine, erect, and dense (Fig. 84, right side); male frons broader, frons 0.06 (0.055— 0.065)/7 head width. Widespread in North America. P. rudis Pollenia angustigena Wainwright, 1940. Until this study, this species was known only from northeastern North America. I have examined specimens from Califor- nia to Washington, Idaho to Wisconsin, Ohio to New Jersey and south to Virginia. It is similar to P. rudis, but males are usually distinctive. Females are difficult to distinguish since the only good character known is the number of anterodorsal setae on the midtibia. If legs are missing or the setae damaged, then identification of females is difficult. Various characters are illustrat- ed in Rognes (1991: figs. 562,579,594— 603). Pollenia griseotomentosa (Jacentko- vsky, 1944). Rognes (1991) listed this species from Ontario, Canada. I have seen specimens from Maine, Washing- ton, and Wisconsin. It is the only North American Pollenia lacking an _ outer posthumeral seta. Various characters VOLUME 108, NUMBER 3 are illustrated in Rognes (1991: figs. 563, 604-611). Pollenia labialis Robineau-Desvoidy, 1863. Rognes (1991) recorded this spe- cies from Ontario, and Greenberg (1998) listed it from Indiana. I found it from Michigan, Maine, New Hampshire, Ore- gon, and Washington. The lappets of the posterior spiracle are dark brown, which distinguishes it from other species. Dis- colored specimens of other species can be confused with it, although the reduced facial carina separates it from most similar species. Various characters are illustrated in Rognes (1991: figs. 565, 622-628). Pollenia pediculata Macquart, 1834. I examined specimens of this species from Washington to Wisconsin and New York and south to North Carolina. I also found it in Utah, Oregon, and California. Rognes (1991) recorded it from New Mexico. I did not find it in the southeastern U.S. This is the second most common Pollenia I found, next to PY Kudise Olt Vis readily * identified: “by a distinctive bundle of setae on the venter of the wing, at the junction of the humeral crossvein and subcosta. Various characters are illustrated in Rosnmesn(l99l> fies. 7557) 559, 581, 583, 640-650). Pollenia rudis (Fabricus, 1794). This species 1s widespread in North America and was once thought to be the only Pollenia species present. Pollenia speci- mens in most collections are identified as this species, but I have found that half yor simore sare: other species, ‘itis similar to P. angustigena, but males have a broader frons and a denser vestiture on the venter of the abdomen. Female characters are limited to setae on the mid tibia. Various characters are illus- trated in Rognes (1991: figs. 582, 651— 661). Pollenia vagabunda (Meigen, 1826). Rognes (1991) listed this species from British Columbia, Nova Scotia, and 723 Prince Edward Island, and Greenberg (1998) listed it from New York. I also examined specimens from Massachu- setts, New Hampshire, and Washington. In specimens in good condition, a dark median stripe between the presutural acrostichal setae is a distinctive charac- ter. Accessory humeral setae and 2 inner posthumeral setae will further confirm its identity. Various characters are illustrat- ed in Rognes (1991: figs. 569, 662-669). In one sample of 12 Pollenia collected from a home in Tacoma, Washington on April 1, 2005, 8 were P. vagabunda, 3 were P. angustigena, and one was P. rudis. MELANOMYINAE Downes (1986) synonymized Angio- neura and Opsodexia under Melanomya. Shewell (1987) concluded these should be separate genera, an opinion with which Roenes (991) concurred: |Species, oF both genera are rarely encountered in collections, so keys to species are not provided here. The keys in Downes (1986) are useful in making accurate identifications (N. Woodley, in litt.). Species of both genera are dull colored and nondescript. Their biology is poorly known, but Downes (1986) suspects all might be snail parasites. Angioneura Brauer and Bergenstamm, 1893 This genus includes 5 species which have relict populations primarily in the East and Midwest. Opsodexia Townsend, 1915 This genus includes 4 species which apparently have habits and distributions similar to Angioneura. ACKNOWLEDGMENTS This study was made possible with the financial support of my firm, Whit- worth Pest Solutions, Inc. I thank my 724 employees for understanding my late arrivals and early departures from work to pursue this study in my lab in the basement of my home. I especially appreciate the help of my general man- ager, Belinda Bowman, whose diligence has given me the long, undisturbed blocks of time needed to complete this study. Thanks also to my wife Faye, who has accepted my obsession with blow flies and their intrusion into her home. I am especially indebted to James O’Hara of the Canadian National Col- lection, Agriculture and Agri-Food Ca- nada, Ottawa, Ontario, and Knut Rognes of the University of Stavanger, Stavanger, Norway who provided de- tailed answers to my many questions and helped inspire me to complete this study. This work would not have been possible without the cooperation of many museum curators who sent me materials for study. Special thanks to Rich Zack and Will Hanson, curators at Washington State University and Utah State University respectively, who sent me many specimens and were always willing to help. Rich also acted as a liaison to enable me to get specimens that aren’t normally loaned to private individuals. Other curators who sent materials include; from the University of Califor- nia, Berkeley, Cheryl Barr; University of California, Davis, Steve Heydon; Uni- versity of California, Riverside, Doug Yanega; California Academy of Sci- ences, Keve Ribardo; Natural History Museum of Los Angeles, Brian Brown; Florida State Collection of Arthropods, Gary Steck; University of Idaho, Frank Merickel; University of Missouri, Kris Simpson; Montana State University, Richard Hurley; University of New Hampshire, Don Chandler; New Mexico State University, David Richman; Ore- gon State University, Darlene Judd; National Museum of Natural History, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Smithsonian Institution, Norm Wood- ley; Spencer Museum, University of British Columbia, Karen Needham; Uni- versity of Wisconsin, Madison, Steven Krauth. Others who sent materials in- clude Eric Eaton, private collector, Neil Haskell, St. Josephs College, Rensselear, Indiana, and Jeff Wells, West Virginia University, thanks to all. Thanks also to all who reviewed this manuscript, already mentioned are Knut Rognes, James O’Hara, Rich Zack, Norm Woodley, Gary Steck, and Neil Haskell. Other reviewers include James Wallman of the University of Wollon- gong, New South Wales, Australia, Greg Dahlem of Northern Kentucky Univer- sity, Highland Heights, Kentucky, Gail Anderson, Simon Fraser University, Burnaby, British Columbia, and Bruce Cooper, Canadian National Collection. Also, thanks to Dawn Nelson, scien- tific illustrator, who helped me produce quality illustrations to make the keys more understandable. Finally, my Figs. 1-10 from Rognes (1991) are re- produced with permission of E.J.Brill/ Scandinavian Science Press and the author, Knut Rognes. Figs. 11-13 are from McAlpine (1981), Figs. 14, 19-21, 25, 27-32, 42-43, 60, 61, 63, and 65 are from Shewell (1987); all are reproduced with permission of the Minister of Public Works and Government Services Ca- nada, 2004. LITERATURE CITED Catts, P. E. and G. R. Mullen. 2002. Myiasis (Muscoidea, Oestroidea), pp. 318-347. In Mul- len, G. R. and L. A. Durden, eds. Medical and Veterinary Entomology. Academic press, New York. 597 pp. Dear, J. P. 1985. A revision of the New World Chrysomyini (Diptera:Calliphoridae). Revista Brasileira de Zoologia 3(3): 100-169. Downes, W. L. 1965. Tribe Melanomyini, pp. 932— 933. In Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson, eds. A Catalog of the Diptera of North America North of Mexico. Agricultural Handbook 276: IV 1,696 pp. Washington D.C. VOLUME 108, NUMBER 3 1986. The Nearctic Melanomya and relatives (Diptera: Calliphoridae), a problem in calypterate classification. Bulletin of the New York State Museum 460, 35 pp. Greenberg, B. 1998. Reproductive states of some overwintering domestic flies (Diptera: Musci- dae and Calliphoridae) with forensic implica- tions. Arthropod Biology 91(b): 818—820. Greenberg, B. and J. Kunich, eds. (2002). Ento- mology and the law, flies as forensic indicators. Cambridge University Press, 306 pp. Hall, D. G. 1948. The Blowflies of North America. Thomas Say Foundation, Lafayette, Indiana. 477 pp, 51 plates. . 1965. Family Calliphoridae, pp. 922-932. In Stone, A., C. W. Sabrosky, W. W. Wirth, R.H. Foote, and J. R. Coulson, eds. A Catalog of the Diptera of North America North of Mexico. Agricultural Handbook 276, IV + 1,696 pp. Washington D.C. Hall, R. D. and L. H. Townsend. 1977. The blowflies of Virginia: no 11. Virginia Poly- technic Institute and State University, Re- search Division Bulletin 123, vi + 48 pp. International Commission on Zoological Nomen- clature, International Code of Zoological Nomenclature, Fourth Edition. The Interna- tional Trust for Zoological Nomenclature, London, 306 pp. James, M. T. 1953. Notes on the distribution, systematic position, and variation of some Calliphoridae, with particular reference to the species of western North America. Proceedings of the Entomological Society of Washington 55: 143-148. . 1955. The blowflies of California (Diptera: Calliphoridae). Bulletin of the California In- sect Survey 4(1), 34 pp. McAlpine, J. F. 1981. Chapter 2, Morphology and Terminology—Adults, pp. 9-63. In McAlpine, J. F., B: V. Peterson, G. E. Shewell, Hi. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds. Manual of Nearctic Diptera, Vol. 1. Agricul- ture Canada. Monograph 27:i—v1 1—674. Papp, L. and B. Darvis, eds. (1998) Manual of Palearctic Diptera, Vol. 3, Higher Brachycera. Science Herald, Budapest 849 pp. Rognes, K. 1985. Revision of the bird-parasitic blowfly genus Trypocalliphora, Peus, 1960 (Diptera: Calliphoridae). Entomologica Scan- dinavica 15: 371-382. . 1991. Blowflies (Diptera, Calliphoridae) of Fennoscandia and Denmark. E.J. Brill/Scan- dinavian Science Ltd., 272 pp. 125 Sabroskys C= We Go Ee Bennett, and ile i: Whitworth. 1989. Bird blowflies (Protocalli- phora) in North America (Diptera: Calliphor- idae), with notes on Palearctic species. Smith- sonian Institute Press, Washington D.C. , 312 pp. Shahid, S. A., R. D. Hall, N. H. Haskell, and R. W. Merritt. 2000. Chrysomya — rufifacies (Macquart)(Diptera: Calliphoridae) — estab- lished in the vicinity of Knoxville, Tennessee, USA. Journal of Forensic Sciences 45(4): 896-897. Shannon, R. C. 1926. Synopsis of the American Calliphoridae (Diptera). Proceedings of the Entomological Society of Washington 28: 115-119. 1923. Genera of Nearctic Callipho- ridae, blowflies, with revision of the Calliphor- ini. Insecutor Inscitiae Menstruus 11: 101— iG). Shewell, G. E. 1987. Calliphoridae, pp. 1,133—1,145. In McAlpine, J- Fs B. V2 Peterson) ‘G2 E- Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds. Manual of Nearctic Diptera, Vol. 2. Agriculture Canada Monograph 28: i-vi 675-1332. Smith, K. G. V. 1986. A manual of forensic entomology. British Museum (Natural Histo- ry), London, 205 pp. Momberlins J Ke We kK. Reeves and Dy EC: Sheppard. 2001. First record of Chrysomya megacephala (Dipera: Calliphoridae) in Geor- gia, U.S.A. Florida Entomologist 84(2): 300-301. Whitworth, T. L. 2002. Two new species of North American Protocalliphora (Diptera: Calliphori- dae) from bird nests. Proceedings of the Entomological Society of Washington 104: 801-811. . 2003a. A new species of North American Protocalliphora (Diptera: Calliphoridae) from bird nests. Proceedings of the Entomological Society of Washington 105(3): 664-673. . 2003b. A key to the puparia of 27 species of North American Protocalliphora Hough (Diptera: Calliphoridae) from bird nests and two new puparial descriptions. Proceedings of the Entomological Society of Washington 105(4): 995—1,033. Zumpt, F. 1965. Myiasis in Man and Animals in the Old World. A Textbook for Physicians, Veterinarians and Zoologists. Butterworth and Company, London, England, 267 pp. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 726-729 Nore Four New Host Records for Amblyomma geoemydae (Cantor) (Acari: Ixodida: Ixodidae) from Captive Tortoises and Freshwater Turtles (Reptilia: Testudines) in the Turtle Conservation Center, Cuc Phuong National Park, Vietnam There are 23 native species of tortoises and freshwater turtles in Vietnam, and all of them are threatened with extirpation due to insatiable Chinese market demand and widespread, accelerating habitat de- struction (Hendrie 2000). Responding to this crisis, in 1998 Fauna and Flora International, in cooperation with the Vietnamese Ministry of Agriculture and Rural Development, formally established the Turtle Conservation Center (TCC) at Cuc Phuong National Park, in the north- ern part of the country (20°14-20°24’N, 105°29-105°44'E). The TCC is charged with promoting awareness of threats to Vietnam’s native turtles, training wildlife protection officers, and establishing a con- servation breeding program for critically endangered Species. Currently, the TEC encompasses ~ 2,000 m2, with more than 50 outdoor enclosures for terrestrial spe- cies, and several dozen aquatic and semi- aquatic enclosures, including six “‘free- ranging’ enclosures with functioning streams and constructed wetlands. Enclo- sures measure from | m2 to > 84 m2, the larger ones having been designed to represent near-natural environments, with an abundance of leaf litter, shelter sites, and native vegetation (Calle and Hendrie 2003, Hendrie 2003, Calle et al. 2005). Between March 2003 and March 2004, we were able to obtain 75 tick collections, comprising 1,515 specimens, from 5 of the 17 captive tortoise and turtle species then held at the TCC (all seized from illegal traffickers): the Indochinese box turtle, Cuora galbinifrons (Bourret); the three- keeled box turtle, C. mouhotii (Gray); the yellow-headed temple turtle, Hieremys annandalii (Boulenger); the elongated or yellow tortoise, /ndotestudo elongata (Blyth); and the impressed tortoise, Man- ouria impressa (Gunther). The vast major- ity of tick specimens (98 2, 93 2, 1,031 nymphs, 286 larvae) removed from these hosts were Amblyomma geoemydae (Can- tor). In addition, 4 ¢ and 3 & of Amblyomma supinoi Neumann were taken from 1 ¢ and 1 2, respectively, of Z elongata, the principal host of this un- commonly collected tick (Robbins and Platt 2001). All ticks were preserved in 70-80% ethanol and shipped to RGR for identification and analysis, prior to de- position in the Division of Entomology, Peabody Museum of Natural History, Yale University, New Haven, Connecticut. Published records of A. geoemydae exist for the yellow-margined box turtle, Cuora flavomarginata (Gray); C. mouho- tii; the Asian leaf turtle, Cyclemys dentata (Gray); the black-breasted leaf turtle, Geoemyda spengleri (Gmelin); the spiny turtle, Heosemys spinosa (Gray); the Travancore tortoise, Indotestudo tra- vancorica (Boulenger); the Asian brown tortoise, Manouria emys (Schlegel and Miller); the Japanese turtle, Mauremys japonica (TYemminck and Schlegel); and the Indian black turtle, Me/lanochelys trijuga (Schweigger) (Simmons and Bur- ridge. 2000) Thus; four yot sthes ive testudines examined for ticks at the TCC provided new host records for A. geoemydae, and the 27 tick collections made from these hosts are listed in Table 1. Significantly, our two new host tortoises—J. elongata and M. impressa— appear in Appendix II (species not necessarily threatened with extinction but that may become so unless trade is VOLUME 108, NUMBER 3 Table 1. C207 Collections of Amblyomma geoemydae from previously unreported hosts at the Turtle Conservation Center, Cuc Phuong National Park, Vietnam, 12 March 2003-17 March 2004 (Cg = Cuora galbinifrons, Ha = Hieremys annandalii, le = Indotestudo elongata, Mi = Manouria impressa; N = nymph, ee — slave): Tortoise/Turtle No./Sex Tick Totals Tick Attachment Sites Date Case? i 2 no data 15 April 2003 Ha4, ¢ ) 2 TUN carapace, neck, tail 21 March 2003 Iel5, & 1 2 carapace 10 November 2003 Ie32, 2 1 2 rear leg 12 March 2003 Ie33, § 1 2 carapace 20 October 2003 1e34, 2 1 2 carapace 9 April 2003 [e93, @ i carapace 20 October 2003 Ie170, 2 1 rear leg 5 June 2003 1e268. ¢ list & carapace, tail 30 March 2003 1e305. & Bg carapace 20 October 2003 1e314. o il & carapace 13 March 2003 1e365. 2 Ayam les carapace, plastron 15 October 2003 1e416, & Dard sah 8 carapace 15 October 2003 1e424. 2 5 & carapace, plastron 15 October 2003 1e425. & DE carapace 20 October 2003 1e428. ¢ 12 plastron 4 October 2003 1e459. & ite plastron 15 October 2003 1e589, 3 lied: carapace 30 March 2003 1e643, & il & carapace 11 April 2003 1e649. & i carapace 20 October 2003 Ie651, rg carapace 20 October 2003 Ie661, 2 IN front leg 20 August 2003 1e662, ¢ 3N front leg, neck 20 August 2003 1e663, ° 2 E22 iN carapace, plastron, rear leg 20 August 2003 1e664, ¢ 6 2. 27N front leg, neck, plastron, rear leg 20 August 2003 legge. 12 rear leg 17 March 2004 Mi25. ° 4422.60N,16L carapace, front leg, neck, rear leg 17 March 2003 strictly regulated) of the Convention on International Trade in Endangered Spe- cies of Wild Fauna and Flora (van Dijk et al. 2000), a signal that opportunities to investigate similar host-parasite relation- ships may be fading. Amblyomma geoemydae 1s chiefly a tick of southern and southeastern Asia, most records having come from peninsular and insular Malaysia (Sabah, Sarawak), Indonesia (Java, Kalimantan Borneo, Sumatra), the Philippines, Singapore, Thailand, and Vietnam (Tanskul et al. 1983, Kolonin 1995, Petney and Keirans 1995, Voltzit and Keirans 2002). In eastern Asia, this species is known from southern Japan (Kyushu and the Ryu- kyu Islands) and Taiwan (Yamaguti et al. 1971, Robbins 2005). An unpublished collection from Sri Lanka (U.S. National Tick Collection No. RML 106681) and recent records from the Western Ghats of India (states of Karnataka and Kerala) suggest that A. geoemydae may occur in all countries bordering the Bay of Bengal (Vijaya 1983, Frazier and Keirans 1990). Among the most distinc- tive amblyommines in Asia (the female foveae are unusually large and_ the nymphal scutum is conspicuously or- 728 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Non-testudine hosts of Amblyomma the U.S. National Tick Collection. geoemydae specimens in Host Location Tick Stage(s) Reptiles Python reticulatus (Schneider) Malaysia: Johor 2 IN IL Varanus dumerilii (Schlegel) Malaysia: Johor N Varanus salvator (Laurent) Malaysia: Johor, Perak, Sabah, Selangor Gis FIN Thailand: Tak Birds Anthracoceros malayanus (Raffles) Malaysia: Pahang N Centropus sinensis (Stephens) Malaysia: Pahang N Ceyx erithacus (L.) Malaysia: Negeri Sembilan N Dupetor flavicollis (Latham) Malaysia: Pahang iL. Lacedo pulchella (Horsfield) Thailand: Nakhon Si Thammarat N Napothera macrodactyla (Strickland) Malaysia: Negeri Sembilan N Pitta brachyura (L.) Malaysia: Pahang INS Pitta moluccensis (Statius Muller) Malaysia: Pahang L Pitta sordida (Statius Miller) Malaysia: Negeri Sembilan, Pahang N, L Platylophus galericulatus (Cuvier) Malaysia: Selangor N Rhizothera longirostris (Temminck) Malaysia: Kelantan N Stachyris poliocephala (Temminck) Malaysia: Negeri Sembilan, Selangor N Zoothera marginata Blyth Thailand: Chanthaburi N Mammals Arctogalidia trivirgata (Gray) Malaysia: Perak 2 Hemigalus derbyanus (Gray) Malaysia: Terengganu N Homo sapiens L. Indonesia: Sumatra # Malaysia: Sarawak Manis crassicaudata Gray Sri Lanka 2 Pardofelis marmorata Martin Thailand: Nakhon Ratchasima N Sus scrofa L. Malaysia: Johor, Kedah, Kelantan, ose SN Pahang, Perak, Selangor nate), A. geoemydae principally parasi- tizes testudines, though other reptiles and, occasionally, mammals may serve as hosts of all stages, while birds are sometimes infested by immatures (Hoog- straal and Aeschlimann 1982). Non- testudine hosts of specimens in the U.S. National Tick Collection are summa- rized in Table 2. Despite the diversity of apparently suitable host species, no disease agents have been associated with bites of this tick. We anticipate that ongoing studies at the TCC will yield additional new host records for A. geoemydae, as well as data on tick population dynamics. We thank the officers of Cuc Phuong National Park and the staff of the TCC for their assistance in making and ship- ping our tick collections. Thanks also to the Wildlife Conservation Society’s Field Veterinary Program for invaluable logis- tical support. Dr. James E. Keirans, former Curator of the U.S. National Tick Collection, Institute of Arthropodology and Parasitology, Georgia Southern Uni- versity, Statesboro, kindly supplied all records of A. geoemydae in his care. This study was supported by a grant from the Conservation Endowment Fund of the American Zoo and Aquarium Associa- tion and by the Wildlife Conservation Society’s Health Sciences Division. LITERATURE CITED Calle, P. P. and D. Hendrie. 2003. Development of a chelonian health care program at the Turtle Conservation Center, Cuc Phuong National Park, Vietnam. Proceedings of the Association VOLUME 108, NUMBER 3 of Zoo Veterinary Technicians, 23™4 Annual Meeting, New York, New York, 24-28 August 2008 psa: Calle; BP. P., B: &. Raphael, J. L. Behler, W. Holmstrom, D. McAloose, M. D. M. Busch, J. S. Spratt, T. M. Norton, D. B. Hendrie, and B. D. Phong. 2005. Development of a chelonian health care program for the Turtle Conserva- tion Center, Cuc Phuong National Park, Vietnam. Proceedings of the Association of Reptilian and Amphibian Veterinarians, 12th Annual Conference, Tucson, Arizona, 9—14 April 2005, pp. 17-18. Frazier, J. G. and J. E. Keirans. 1990. Ticks (Acari: Ixodidae) collected on chelonians (Reptilia) from India and Burma. Journal of the Bombay Natural History Society 87: 247-249. Hendrie, D. B. 2000. Status and conservation of tortoises and freshwater turtles in Vietnam, pp. 63-73. In van Dijk, P. P., B. L. Stuart, and A. G. J. Rhodin, eds. Asian Turtle Trade: Proceedings of a Workshop on Conservation and Trade of Freshwater Turtles and Tortoises in Asia. Chelonian Research Monographs 2. Chelonian Research Foundation, Lunenburg, Massachusetts. 2003. Turtle Conservation Indochina. Newsletter of the Turtle Conservation and Ecology Program [sic] (TCEP). Accessed on the Worldwide Web at URL , September 2005. Hoogstraal, H. and A. Aeschlimann. 1982. Tick- host specificity. Mitteilungen der Schweizer- ischen Entomologischen Gesellschaft/Bulletin de la Société Entomologique Suisse 55: 5—32. Kolonin, G. V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. Journal of Medical Entomology 32: 276—282. Petney, T. N. and J. E. Keirans. 1995. Ticks of the genera. Amblyomma and Hyalomma_ from South-east Asia. Tropical Biomedicine 12: 45—S6. Robbins, R. G. 2005. The ticks (Acari: [xodida: Argasidae, Ixodidae) of Taiwan: A synonymic checklist. Proceedings of the Entomological Society of Washington 107: 245-253. Robbins, R. G. and S. G. Platt. 2001. First report of Amblyomma supinoi Neumann (Acari: Ix- odida: Ixodidae) from the Arakan forest turtle, Geoemyda depressa Anderson (Reptilia: Testu- dines: Emydidae), with additional records of this tick from the Union of Myanmar. Proceedings of the Entomological Society of Washington 103: 1023-1024. Simmons, L.-A. and M. J. Burridge. 2000. In- troduction of the exotic ticks Amblyomma humerale Koch and Amblyomma geoemydae 2S) (Cantor) (Acari: Ixodidae) into the United States on imported reptiles. International Journal of Acarology 26: 239-242. Tanskul, P. (L.), H. E. Stark, and I. Inlao. 1983. A checklist of ticks of Thailand (Acari: Metas- tigmata: Ixodoidea). Journal of Medical Ento- mology 20: 330-341. van Dijk, P. P., B. L. Stuart, and A. G. J. Rhodin, eds. 2000. Asian Turtle Trade: Proceedings of a Workshop on Conservation and Trade of Freshwater Turtles and Tortoises in Asia. Chelonian Research Monographs 2. Chelonian Research Foundation, Lunenburg, Massachu- setts, 164 pp. Vijaya, J. 1983. First report of Amblyomma geoemydae from India. Hamadryad 8: 13. Voltzit, O. V. and J. E. Keirans. 2002. A review of Asian Amblyomma species (Acari, Ixodida, Ixodidae). Acarina 10: 95-136. Yamaguti, N., V. J. Tipton, H. L. Keegan, and S. Toshioka. 1971. Ticks of Japan, Korea, and the Ryukyu Islands. Brigham Young Univer- sity Science Bulletin, Biological Series 15: i-iv, 1-226. Richard G. Robbins, Bui Dang Phong, Tim McCormack, John L. Behler, Hendrik A. Zwartepoorte, Douglas B. Hendrie, and Paul P. Calle. (RGR) Armed Forces Pest Management Board, Walter Reed Army Medical Center, Washington, DC 20307- 5001, U.S.A. (e-mail: richard.robbins@ osd.mil); (BDP) Turtle Conservation Cen- ter, Cuc Phuong National Park, Vietnam (e-mail: phongtcccucphuong@gmail.com); (TM) Turtle Conservation Center, Cuc Phuong National Park, Vietnam (e-mail: the_agent_badger@yahoo.co.uk); (JLB) Department of Herpetology, Wildlife Conservation Society and IUCN/ISSC Tortoise and Freshwater Specialist Group, 2300 Southern Boulevard, Bronx, NY 10460-1099, U.S.A. (e-mail: jbehler@ wes.org); (HAZ) Rotterdam Zoo and Turtle Survival Alliance-Europe, Rotterdam, The Netherlands (e-mail: h.zwartepoorte@ rotterdamzoo.nl); (DBH) Wildlife Conser- vation Society and Cleveland Metroparks Zoo, 3900 Wildlife Way, Cleveland, OH 44109-3187, U.S.A. (e-mail: dhendrie@ fpt.vn); (PPC) Wildlife Health Sciences, Wildlife Conservation Society, 2300 Southern Boulevard, Bronx, NY 10460-1099, U.S.A. (e-mail: pcalle@wes.org ) PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 730-731 NOTE A New Record of the Flea Tarsopsylla octodecimdentata coloradensis (Baker, 1895) (Siphonaptera: Ceratophyllidae) in Utah Tarsopsylla Wagner, 1927, is a holarctic genus for which the nominate species [Tarsopsylla octodecimdentata octodecim- dentata (Kolenati, 1863)] occurs from Europe to extreme eastern Asia. Nearctic populations are considered a subspecies [Tarsopsylla octodecimdentata coloraden- sis (Baker, 1895)] of the nominate species and are found from Alaska to New Mexico (Haddow et al. 1983). Baker (1895) originally described 7. 0. color- adensis from a single male (holotype: Pulex coloradensis USNM No. 104605) collected on Fremont’s Chickaree [= Tamiasciurus —hudsonicus (Erxleben)], George Town, 39°43'N 105°42’W, [Clear Creeks County], Colorado, |. Bruner: Holland (1985) provided a detailed ac- count of records throughout Alaska and western Canada. In addition to the original description, only five records have been documented in montane re- gions of the western United States. These include; 1) one female collected from the nest of Glaucomys sabrinus (Shaw), Powwatka Ridge, Wallowa County, Ore- gon, 17.XII.1977, M. Huff. (Lewis et al. 1988, Whitaker et al. 1983) (collection data obtained from R. E. Lewis, pers. Comm.); 2) two females collected from Martes americana (Turton), Larimer County, Colorado, X.1977 (Eads et al. 1979): 3),one female collected. from 7: hudsonicus, San Pedro Mt., 2670-2733 m, Sandoval County, New Mexico, V.1969— 1971 (Haas et al. 1973); 4) 13 males and 12 females collected from four T. hudso- nicus nests, 6.X.1953 (Morlan 1955): and 5) two males collected from G. sabrinus, Deer Park, Boise, Idaho, 15—18.XII.1939, W.H. Marshall. The latter specimens were originally described as Opisodasys Jellisoni 1. Fox, 1941 (USNM No. 54249) and Holland (1949) subsequently synon- ymized O. jellisoni with that of T. o. coloradensis. This flea is rarely collected and Morlan’s (1955) collection of such a large series of adults from four nests would support the premise that this is a true nest flea collected only during the cooler months of the year. It undoubted- ly occurs in many montane localities in the western United States in conjunction with the red squirrel, 7: hudsonicus, and the northern flying squirrel, G. sabrinus. These squirrels frequent each others nest- ing tree cavities and both are occasionally prey to the pine marten, M. americana. Therefore, this flea may be found on all three of these host species but the true host is probably 7. hudsonicus. This first report of 7: 0. coloradensis in Utah is represented by a single female collected from G. sabrinus, nr. Marsh Lake and China Lake (Smith’s Fork drainage, north slope of Uinta Moun- tains) (40°57’N- 110°24’W), Summit County, Utah, 13.X1.2001. Holland (1985) pointed out the variation that may occur in the female seventh sternite. The caudal margin of the seventh sternite of the Utah female has a broadly squared dorsal lobe subtended by a broad mod- erately deep sinus and a ventral lobe. This specimen is retained in the author’s personal flea collection. Tipton and Saunders (1971) reported 115 species of fleas in Utah. Two are considered invalid species [Eumolpianus eumolpi americanus (Hubbard, 1950) (see Lewis and Jameson 2002), and Meringis jJewetti Hubbard, 1940 (see Smit 1953)] and the presence of a third, Stenistomera hubbardi Egoscue, 1968, can not be VOLUME 108, NUMBER 3 validated in Utah. Two additional species were subsequently documented in Utah by Kucera (1995) [Euhoplopsyllus glacia- lis lynx (Baker, 1904)] and Hastriter (1997) (Tunga monositus Barnes and Radovsky, 1969). With the addition of T. o. coloradensis, the total number of recognized species (including subspecies) in Utah is 115 with 53 genera represented. Acknowledgment.—I am grateful to Duke S. Rogers, Department of Integra- tive Biology, Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah, for graciously contributing this new record for study. LITERATURE CITED Baker, C. F. 1895. Preliminary studies in Sipho- naptera, III. Canadian Entomologist 27: 108-111. Eads, R. B., E.G. Campos, and A. M. Barnes. 1979. New records for several flea (Siphonaptera) species in the United States, with observations on species parasitizing carnivores in the Rocky Mountain region. Proceedings of the Entomo- logical Society of Washington 81: 38—42. Fox, I. 1941. A new Opisodasys from Idaho (Siphonaptera: Dolichopsyllidae). Entomologi- cal News 52: 45-47. Haas, G. E., R. P. Martin, M. Swickard, and B. E. Miller. 1973. Siphonaptera-mammal relation- ships in northcentral New Mexico. Journal of Medical Entomology 10: 281—289. Haddow, J. R., Traub, and M. Rothschild. 1983. Distribution of ceratophyllid fleas and notes on their hosts, pp. 42-163. In Traub, R., M. Rothschild, and J. F. Haddow, eds. The Rothschild Collection of Fleas. The Cerato- phyllidae: Key to the genera and _ host relationships. University Press, Cambridge. 288 pp. 731 Hastriter, M. W. 1994. Establishment of the tungid flea, Tunga monositus (Siphonaptera: Pulici- dae), in the United States. Great Basin Naturalist 57: 281—282. Holland, G. P. 1985. The fleas of Canada, Alaska, and Greenland (Siphonaptera). Memoirs of the Entomological Society of Canada, No. 130, 631 pp. Kucera, J. R. 1995. Additional records of fleas (Siphonaptera) from Utah. Great Basin Natu- ralist 55: 92-94. Lewis, R. E. and E. W. Jameson, Jr. 2002. A review of the flea genus Ewmnolpianus Smit, 1953 with a discussion of its geographic distribution and host associations (Siphonaptera: Certaophylli- dae: Ceratophyllinae). Journal of Vector Ecol- ogy 27: 235-249. Lewis, R. E., J. H. Lewis, and C. Maser. 1988. Fleas of the Pacific Northwest. Oregon State University Press, Corvallis. 296 pp. Morlan, H. B. 1955. Mammal fleas of Santa Fe County, New Mexico. Texas Reports on Biology and Medicine 13: 93-125. Smit, F. G. A. M. 1953. Monstrosities in Sipho- naptera. IV. More cases of castration. Ento- mologische Berichten 14: 393-400. Tipton, V. J. and R. C. Saunders. 1971. A list of arthropods of medical importance which occur in Utah with a review of arthropod-borne diseases endemic in the state. Brigham Young University Science Bulletin, Biological Series 15, 1-31. Whitaker, J. O., Jr., E. A. Lyons, M. A. Smith, and C. Maser. 1983. Nest inhabitants and ectopar- asites of Northern Flying Squirrels, Glaucomys sabrinus (Shaw), from northeastern Oregon. Northwest Science 57: 291—295. Michael W. Hastriter, Monte L. Bean Life Science Museum, Brigham Young University, 290 MLBM, P.O. Box 20200, Provo, UT 84602-0200, U.S.A. (e-mail: mwhastri ter@sprintmail. com ) PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 732-733 NOTE Types and Voucher Specimens of New World Aulacidae (Hymenoptera) in the Naturhistoriska Riksmuseet, Stockholm, Sweden Kieffer (1904) and Roman (1917) de- scribed several New World species of Aulacidae, the types of which are de- posited in the Naturhistoriska Riksmu- seet, Stockholm, Sweden (NHRS). Ad- ditionally, Roman (1917) recorded other species from South America based on specimens located in the same museum. Some of these specimens are not clearly labeled. I had the opportunity to study these specimens and here record them and designate several lectotypes that I feel are necessary. Aulacinus amazonicus Roman 1917: 18. This was described from *“‘1 female.” The holotype is labeled “‘Mana-os,” **Amazon Roman,” ““Type”’ (small white label), and Roman’s determination label. I have added a red holotype label. Smith (2005) redescribed and illustrated this Species, hey icorrect, yecombination) 1s Aulacus amazonicus (Roman) (Smith 2001). Pristaulacus zonatipennis Roman 1917: 18. This was described from “‘1 female.” The holotype is labeled “Amazon Ro- man,” ““Mana-os,”’ ““Type”’ (small white label), and Roman’s determination label. I have added a red holotype label. It was redescribed and illustrated by Smith (2005). The species is correctly placed in Pristaulacus. Pristaulacus haemorrhoidellus (West- wood). Roman (1917: 18) recorded two females from Rio Autaz, Cururuzinho, Brazil. Two females labeled ““Rio Autaz”’ and ““Amazon Roman” are at NHRS, one with Roman’s determination label. Both are certainly the specimens Roman recorded, and I have labeled both speci- mens. Both specimens were correctly identified. Pristaulacus bicornutus (Schletterer). Roman (1917: 18) recorded one female from Rio Autaz, Cururuzinho, Brazil, and one female from Manaos, Rio Taruma, Brazil. Both specimens on which Roman’s records are based are at NHRS, one labeled ‘“‘Mana—os,”’ ““Ama- zon Roman” and the other ““Rio Autaz,”’ *“Amazon Roman.” One bears Roman’s determination label. The species is a syn- onym of Pristaulacus ruficeps (West- wood) (Smith 2005), and I have labeled both specimens as such. Pristaulacus flavipes Kieffer 1904: 559. Kieffer described this species from ‘3 Exemplare,” all males, from “Illinois: Belfrage.” Three specimens at NHRS bear the labels “‘Illinois” and “‘Belfrage,”’ but only one has an additional label “type” and handwritten label *‘Pristau- lacus flavipes.” I designate the specimen with the type label as lectotype, to preserve stability of the name. The other two specimens are labeled paralecto- types. Pristaulacus flavipes was correctly placed as a synonym of Pristaulacus flavicrurus (Bradley) by Townes (1950). Aulacus erythrogaster Kieffer 1904: 561. This was described from ‘‘Nevada (3 2,1 ¢).” There are two 2 and two males labeled ““Nevada”’ at NHRS, but only one 2 and one ¢@ have a small “type” label. Perhaps Kieffer recorded the wrong sex for one of the specimens. I consider the female and male with the type labels as syntypes and choose the female with the following labels as lectotype to preserve stability of the name: ““Nevada,” “type” “‘Aulacus ery- throgaster [handwritten].’’ The male with the type label is a paralectotype. The other male and female, without “‘type”’ VOLUME 108, NUMBER 3 labels were undoubtedly part of the type series, but since the sexes do not match and they lack a type label, I do not consider them paralectotypes. All speci- mens are clearly labeled. Aulacus ery- throgaster was correctly placed as a syn- onym of Pristaulacus rufitarsis (Cresson) by Townes (1950). Aulacinus costulatus Kieffer 1904: 561. The female was described and the locality given is “‘Brasilien.”” Kieffer did not state the number of specimens examined. One female at NHRS bears the labels. Brasil,” “F. S [illegible, probably the collector], “Type,” and ‘“Aulacinus costulatus [handwritten].” I designate this specimen as lectotype to preserve stability of the name. This species was correctly placed as Aulacus costulatus by Smith (2001). There are four other specimens identical to the lectotype at NRHS from “‘S:ta Cathar- ina, Nova Teutonia, Brasil, 1.1938, Fr. Plaumann.” Acknowledgments.—I thank Fredrik Ronquist, formerly at the Naturhistor- iska Riksmuseet, Stockholm, for arran- ging loan of the specimens. Ronald A. Ochoa and Thomas J. Henry, Systematic 733 Entomology Laboratory, USDA, Belts- ville. MD, and Washington, DC, re- spectively, reviewed and offered sugges- tions for improvement of the manu- script. LITERATURE CITED Kieffer, J. J. 1904. Beschreibung neuer Proctotry- piden und Evaniiden. Arkiv f6ér Zoologi 1: 525-562. Roman, A. 1917. Schlupfwespen aus Amazonien. Arkiv for Zoologi 11: 1—24. Smith, D. R. 2001. World catalog of the family Aulacidae (Hymenoptera). Contributions on Entomology, International 4(3): 263-319. ——. 2005. Aulacidae (Hymenoptera) of north- ern South America, emphasizing Colombia. Transactions of the American Entomological Society 131: 217-253. Townes, H. 1950. The Nearctic species of Gaster- uptiidae (Hymenoptera). Proceedings of the United States National Museum 100: 85—145. David R. Smith, Systematic Entomo- logy Laboratory, PSI, Agricultural Re- search Service, U.S. Department of Agriculture, clo National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 168, Washington, DC 20012-7012, U.S.A. (e-mail: dsmith@ sel. barc.usda. gov ) PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 734-735 NOoTtTE Remarks on Parvaverrucosa annulata (= Verrucosa annulata Poinar and Brown 2005) (Hemiptera: Sternorrhyncha: Aphidoidea) In a recent publication (Poinar and Brown 2005), a fossil aphid in Lower Cretaceous amber from Myanmar (Burma) was described in the genus Verrucosa Poinar and Brown. It was brought to our attention that this name is preoccupied by Verrucosa MacCook (1888), a genus of orb weaving spiders in the family Araneidae. As a replacement name for Verrucosa Poinar and Brown and the family Verrucosidae Poinar and Brown, we propose the genus and family name Parvaverrucosa and Parvaverruco- sidae, respectively. “Parva” is from the Latin ‘“‘parvus’ for little (feminine): ‘“Verrucosus’ 1s Latin for “‘full of warts” (feminine) in reference to the size of the warts comprising the wing armature. In the original description of Parva- verrucosa annulata, we did not observe the hind wings. However in a recently acquired specimen (B-He-13C deposited in the Poinar amber collection at Oregon State University) originating from the same locality as the holotype, the right hind wing is reduced to a stub or hamulohalter (Figs. 1—2). In re-examin- ing the holotype, a similar structure was noted (Fig. 3). These hamulohalters are covered with warts, smaller than those on the forewings, and contain a pair of modified hamuli protruding near their tips. These hamuli are very similar to those on the hind wings of the western wheat aphid (Diuraphis tritici (Gillette) (Ni et al. 2002). The other two described aphids from Burmese amber, Burmita- phis prolatum Poinar and Brown (2005) and Caulinus burmitis Poinar and Brown (2005), do not have hamuli on their hamulohalters. The presence of hamuli on P. annulata supports our view that the hamulohalter is a greatly reduced hind wing and that this condition is a second- ary adaptation. It is interesting that these hamuli are so well developed in compari- son to the reduced hind wings. Whether they still attach to the claval fold on the forewing during flight is unknown. LITERATURE CITED MacCook, H. C. 1888. Necessity for Revising the Nomenclature of American Spiders. Proceed- ings of the Academy of Natural Sciences, Philadelphia, 1888: 74-79. Ni, X., G. D. Johnson, and S. S. Quisenberry. 2002. Comparison of hind wings hamuli from five species of cereal aphids (Hemiptera: Aphididae). Annals of the Entomological Society of America 95: 109-114. Poinar, Jr., G. O. and A. E. Brown. 2005. New Aphidoidea (Hemiptera: Sternorrhyncha) in Burmese amber. Proceedings of the Entomo- logical Society of Washington 107: 835-845. George Poinar, Jr. and Alex E. Brown, (GP) Department of Zoology, Oregon State University, Corvallis, OR 97331, U.S.A. (e-mail: poinarg@science.oregonstate. edu ); (AEB) 629 Euclid Avenue, Berkeley, CA 94708, U.S.A. (e-mail: alex.brown3(@comcast. net ) VOLUME 108, NUMBER 3 135 Figs. 1-3. Parvaverrucosa annulata. 1, Dorsal view of specimen B-He-13C. Arrows show rostrum. Bar = 514 um. 2, Hamuli (arrow) on hamulohalter of specimen B-He-13C. Bar = 36 um. 3, Hamuli (arrow) on hamulohalter of holotype. Bar = 42 um. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 736-737 NOTE First Record of Aedes (Ochlerotatus) grossbecki (Dyar and Knab) (Diptera: Culicidae) in Alabama, U.S.A. Aedes ( Ochlerotatus) grossbecki (Dyar and Knab) (Savage and Strickman 2004) is described as a rare eastern/northeast- ern species reported from a few scattered localities in the Southeast (King et al. 1939, Carpenter et al. 1946, King et al. 1960). Its distribution has expanded to reach from Texas northeastward to New Hampshire, with the conspicuous excep- tion of Alabama, Georgia, and Florida (Darsie and Ward 2005). The current recognized distribution of Ae. grossbecki in the United States is Arkansas, Dela- ware, Illinois, Kentucky, Louisiana, Maryland, Mississippi, Missouri, New Jersey, New York, North Carolina, Ohio, South Carolina, Tennessee, Ver- mont, Virginia, Connecticut, Indiana, Pennsylvania, and Wisconsin (Rayburn et al. 2004, Darsie and Ward 2005). Larvae of the species are typically found in early spring pools and are common throughout the post oak flats along the Mississippi River (Carpenter and La- @asserl1 955. Kane et aly 1960). Mo: date; information regarding adult habits of Ae. grossbecki is limited, although it 1s reported to be a persistent biter (Car- penter et al. 1946, Carpenter and La- Casse 1955, King et al. 1960). It is also speculated that the species is univoltine (Headlee 1945, Carpenter et al. 1946, King et al. 1960), thereby limiting their nuisance potential. We report the first collections of Ae. grossbecki from Ala- bama, leaving only a southeastern gap in Georgia and Florida where the species has not been recorded. Alabama collection records are: Red- stone Arsenal, 19 April 2005, CDC light trap baited with CQOs,, Collector: Michael S. King, 1 2; Redstone Arsenal, 3 May 2005, CDC light trap baited with CO>, Collector: Michael S. King, 1 ¢. Mosquito surveillance is conducted at Redstone Arsenal, Alabama, as part of the United States Army Center for Health Promotion and Preventive Medi- cine-South (USACHPPM-S) West Nile virus surveillance program. Collections are made using CO, baited CDC light traps hanging five to six feet high. In accordance with the surveillance pro- gram at Redstone Arsenal, the traps are placed in the late afternoon and retrieved early the following morning. Mosquitoes are killed by freezing at 0°C for 30 min- utes, placed in microcentrifuge tubes labeled with collection data and then shipped to USACHPPM-South after preliminary identification by the pre- ventive medicine staff at Redstone Arse- nal (Darsie and Ward 1981). On 26 April and 4 May 2005, two separate submissions from Redstone Ar- senal were received at USACHPPM- South for identification and assay proces- sing. Each of these submissions contained one Ae. grossbecki specimen, the identifi- cation of which was confirmed by the Entomological Sciences Division (ESD) staff (Carpenter and LaCasse 1955) and further verified by Leopoldo M. Rueda as Ae. grossbecki (Dyar and Knab) (Carpen- ter and LaCasse 1955, Darsie and Ward 2005). Specimens have been deposited in the Walter Reed Biosystematics Unit mosquito collection at the Smithsonian Institution in Washington, DC. The two specimens were collected from trap sites 7 (34°40.40’N, 86°37.53'W) and 11 (34°42.01'’N, 86°38.19’W) on Redstone Arsenal. Both locations are within 100 feet of a creek and one is VOLUME 108, NUMBER 3 near a construction site. The construc- tion site may have possessed temporary vernal pools, which provided oviposition sites for this woodland pool breeder (Carpenter and WaCasse 1955;9Kung et al. 1960). Both traps sites are in close proximity to human activity, one near a parking lot used for permanent party barracks and the other behind a set of Youth Services batting cages. There are wooded areas near both traps, which although they are limited in size and density, could have provided a breeding site for the species from where they flew some distance to the traps. Based on the accepted distribution of Ae. grossbecki, its discovery in Alabama is not surprising. According to records from the 2004 USACHPPM-South arboviral surveillance program, twenty-one Ae. grossbecki specimens were submitted from Redstone Arsenal in April and May of that year. However, these specimens were not submitted for confirmatory identifica- tion or deposition. Based on the 2004 and 2005 records, the species most commonly associated with Ae. grossbecki at Redstone Arsenal were Aedes vexans (Meigen) and Anopheles punctipennis (Say). Many thanks to Leopoldo M. Rueda for his confirmatory identification, LTC Mustapha Debboun for his literary guid- ance, and COL(R) Daniel Strickman and LTC(R) John Gingrich for reviewing the manuscript and providing suggestions and advice. The opinions and assertions presented herein are those of the authors and not representative of official views of the Department of Defense. LITERATURE CITED Carpenter, S. J. and W. J. LaCasse. 1955. Mos- quitoes of North America (North of Mexico). University of California Press, Berkeley and Los Angeles, California, 360 pp. Carpenter, S. J.. W. W. Middlekauff, and R. W. Chamberlain. 1946. The Mosquitoes of the Southern United States East of Oklahoma and Texas. The University Press, Notre Dame, Indiana, 292 pp. for 737 Darsie, R. F., Jr and R. A. Ward. 1981. Identifica- tion and geographical distribution of the mos- quitoes of North America, north of Mexico. Mosquito Systematics Supplement |: 1-313. 2005. Identification and Geographical Distribution of the Mosquitoes of North America, North of Mexico. University Press of Florida, Gainesville, Florida, 383 pp. Headlee, T. J. 1945. The Mosquitoes of New Jersey and their Control. Rutgers University Press, New Brunswick, New Jersey, 229 pp. King, W. V., G. H. Bradley, and T. E. McNeel. 1939. The Mosquitoes of the Southeastern States. Miscellaneous Publication No. 336. U.S. De- partment of Agriculture, Washington DC, 91 pp. King, W. V., G. H. Bradley, C. N. Smith, and W. C. McDuffie. 1960. A Handbook of the Mosquitoes of the Southeastern United States. Agricultural Handbook No. 173, U.S. Depart- ment of Agriculture, Washington DC, 188 pp. Rayburn, W. H., Jr, B. M. Parker, J. E. Andrews, R. F. Collins, and B. A. Harrison. 2004. Three new mosquito records for North Carolina. Journal of the American Mosquito Control Association 20(4): 451-453. Savage, H. M. and D. Strickman. 2004. The genus and subgenus categories within Culicidae and placement of Ochlerotatus as a subgenus of Aedes. Journal of the American Mosquito Control Association 20(2): 208-214. Jennifer Caci, Michael S. King, Jeri Humphries, Kristin Cobb, Stephen Garvin, Dennis D. Kuhr, Jimmy Wedincamp, and Erin Stanwix, (JC) U.S. Army Command and General Staff College, Intermediate Ley- el Education- Army Operational Warfight- er Course, Fort Leavenworth, KS 66027, U.S.A. (e-mail: Jennifer.caci@us.army.mil); (MSK) Department of Preventive Medicine, Fox Army Community Hospital, Redstone Arsenal, AL 35808, U.S.A.; (JH) Colorado State University, Department of Technical Communications, Fort Collins, CO 80523, U.S.A.; (KC, SG, DDK) U.S. Army Center Health Promotion and Preventive Medicine-South, Fort McPherson, GA 30330-1075, U.S.A.; (JW) MathlScience Division, 131 College Circle, East Georgia College, Swainsboro, GA 30401, U.S.A.; (ES) Reedy Creek Improvement District, 2192 South Service Lane, Lake Buena Vista, Florida, 32830, U.S.A. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, p. 738 Note New Synonym and Western Range Extension for Heterocloeon anoka (Daggy) (Ephemeroptera: Baetidae) Pseudocloeon anoka Daggy, Ps. ed- mundsi Jensen, and Ps. myrsum Burks were placed into synonymy with Baetis punctiventris (McDunnough) (now in Plauditus Lugo-Ortiz and McCafferty) by McCafferty and Waltz (1990). In the revision of the genus Heterocloeon McDunnough by McCafferty et al. (2005), Ps. anoka (now in Heterocloeon) was shown to be a valid and distinctive species, Ps. myrsum was confirmed to be a variant of P. punctiventris (McCafferty et al. 2005, Figs. 20-21), and no action was taken with respect to Ps. edmundsi because, unlike the above, its types had not been reexamined. The differences between H. anoka and P. punctiventris were unclear until very recently, and this resulted in numerous identifications and records of H. anoka having been in- correctly attributed to P. punctiventris. McCafferty et al. (2005) reviewed newly established distribution records of H. anoka, indicating a primarily eastern and central Canada and USA range overlapping with that of P. punctiventris to a considerable extent. Because of the historical confusion between H. anoka and P. punctiventris, it became necessary to reassess the synonymy of Ps. edmundsi to determine if it is perhaps equivalent to H. anoka instead of P. punctiventris. Jensens’s (1969) description of Ps. edmundsi was of no aid in resolving this issue, but fortunately the types of Ps. edmundsi, consisting of a male adult holotype and male adult paratype taken at the Snake River on Highway 20-26 eight miles northwest of Parma in Canyon County, Idaho, were found in the Purdue Entomological Research Collection, hav- ing been donated by George Edmunds in the 1990s. Color patterns of the type specimens indicated that Ps. edmundsi is synonymous with H. anoka, not P. punctiventris (see McCafferty et al. 2005), and therefore Ps. edmundsi is placed as a subjective junior synonym of H. anoka, new synonym, and removed from synonymy with P. punctiventris. The Idaho record of H. anoka represents a considerable western extension of its range in the USA, although the species has been reported from nearly as far west in Canada by Webb and McCafferty (2003). Based on anecdotal evidence, the lotic habitat of H. anoka 1s of a higher gradient than that of P. punctiventris. LITERATURE CITED Jensen, S. L. 1969. New species of Pseuwdocloeon from Idaho. Pan-Pacific Entomologist 45: 14-15. Mic Cafferty, We and.) R= ss Waltz OU: Revisionary synopsis of the Baetidae (Ephe- meroptera) of North and Middle America. Transactions of the American Entomological Society 116: 769-799. McCafferty, W. P., R. D. Waltz, J. M. Webb, and L. M. Jacobus. 2005. Revision of Heterocloeon McDunnough (Ephemeroptera: Baetidae). Journal of Insect Science. 5(35). Webb, J. M. and W. P. McCafferty. 2003. New records of mayflies (Ephemeroptera) from Alberta. Entomological News 114: 230-232. W. P. McCafferty, Department of Entomology, Purdue University, West Lafayette, IN 47907, U.S.A. (e-mail: mcecaffer@purdue. edu ) PROC. ENTOMOL. SOC. WASH. 108(3), 2006, p. 739 NOTE New Records for Leptopteromyia americana Hardy (Diptera:Asilidae) in Alabama Leptopteromyia americana — Hardy (1947) (Diptera: Asilidae) is a small (6— 6.6 mm long) predatory fly that feeds on Aleyrodidae (Hardy 1947) and possibly other small insects. This species was known from Texas and ranged from Hidalgo to Brownsville and southern Cameron County (Martin 1971). Presum- ably L. americana also ranges into north- ern Mexico. The seasonality of L. amer- icana is poorly documented, with collection dates for the type series of 28 December 1945 and 29 June 1938 being the sole reported collections (Hardy 1947). We report collections of adult L. americana from Malaise traps from Mo- bile River Delta in Alabama, which is outside the known range for this fly. Adults of L. americana were collected from Alabama, Baldwin County, North Delta, 31.13 5N 87.859°W,. 20 Oct. 2004, Coll. J.W. McCreadie; Little Briar Creek, 30.84°N 87.04°W, 2 and 20 Nov. 2000, Coll. J.W. McCreadie; Raft River, 30.77°N, 87.95°W, 20 Nov. 2000, Coll. J.W. McCreadie; Byrnes Lake, 30.79°N Sega. 27 Nov. 2001, Coll, J.W. McCreadie; Ft. Morgan Rd., 30.24°N, 88.86°W, 15 Dec. 2004, Coll. E. Benton; Bon Secour, 30.30°N, 88.74°W, 15 Dec 2004, E. Benton; and Mobile County, Camp Scoutshire, 31.05°N, 88.18°W, 20 Jun. 2004, Coll. E. Benton; County Rd 11, 30.42°N 88.35°W, 14 Dec. 2004, Coll. E. Benton; Camp Sid Edmonds, 30.99°N, 87.79°W, 15 Dec. 2004, E. Benton. Leptopteromyia americana probably ranges along the Gulf Coast from Mexico to Florida. Flies were predomi- nantly trapped in the late fall and early winter from October to December. How- ever the collection in June indicates that adults are also active in the summer. All specimens were identified by A.G. Scar- brough, and vouchers are deposited in the in the National Museum of Natural History, Smithsonian Institution, Wash- ington, DC. We thank Elizabeth Benton and Peter Adler for their help with collecting specimens. The findings and conclusions in this report are those of the authors and do not necessarily represent the views of the funding agency. LITERATURE CITED Hardy, D. E. 1947. The genus Leptopteromyia (Asilidae-Diptera). Journal of the Kansas Entomological Society 20: 72-75. Martin, C. H. 1971. A review of the genus Leptopteromyia in the Western Hemisphere (Diptera: Leptogastridae). Pan-Pacific Ento- mologist 47: 265-271. Will K. Reeves, Aubrey G. Scar- brough, and John W. McCreadie, (WKR) Centers for Disease Control and Prevention, 1600 Clifton Rd. NE, Mailstop G-13, Atlanta, GA 30333, U.S.A. (e-mail: cui8; cacde. gov); (AGS) Visiting Professor, Department of Entomology, University of Arizona, Tucson, AZ “85721, USA; (JWC) Department of Biology, University of South Alabama, Mobile, AL 36688, U_S. A. PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 740-742 Book REVIEW Arthropod Collection and Identification, Laboratory and Field Techniques. Timothy J. Gibb and Christian Y. Oseto. Academic Press, Oxford. 2006. 336 pp., 400+figures. ISBN 0-12-369545-7. U.S. $34.95. American entomologists who are in- volved in the collection and identifica- tion of insects and their allies are mostly familiar with the USDA Miscellaneous Publication 1443, “Insects and Mites: Techniques for Collection and Preserva- tion, by Steyskal, Murphy, and Hoover, published in 1986. The present volume, “Arthropod Collection and Identifica- tion’ continues the valuable tradition of presenting current information on the subject. The authors have produced a worthy volume that should become a standard addition to libraries to those interested in entomology, from novice to professional, throughout the varied dis- ciplines of the science. When the volume was first presented to me for review, I was bemused by the images of insects (and a whip scorpion) on the cover. All seven insect species are foreign to the United States, and include two species of birdwing butterflies. Because this volume concerns the U.S. fauna, one would expect the use of equally attractive species from the Nearctic. Many species come easily to mind: regal moth, great crested grasshopper, Phanaeus scarab, rattlebox moth, handsome twig cricket, numerous species of tiger and longhorn beetles. It is easily surmised that the cover images were selected by the pub- lishers, who were at least consistent in selecting foreign species. This soft-covered volume is comfort- able to hold, and the construction allows it to remain open on a flat surface, which is practical when using keys for identifi- cation. It is divided into two main parts: 1) Basic Tools and General Techniques, and 2) Classification of Insects and Mites. The remainder is composed of a glossary, references section, appendix and index. These additions have tripled the size of Steyskal et al. (1986) from 103 to 311 pages. The reference (bibliogra- phy) section has also increased, from 469 to 578 citations. Illustrations of the various collecting traps and methods of preparation in Part One are excellent, and create instant reminders of the efforts necessary for comprehensive sur- veys of entomological faunas of an area. Most of the text throughout the volume is slightly revised from Steyskal et al. (1986), but the authors have included, as Appendix HI, a current Directory of State Extension Staff, an essential in- gredient for many entomological disci- plines, especially with agricultural sub- jects. This Directory reflects current mailing addresses, FAX/telephone num- bers, and internet addresses. The glossa- ry is also new from previous versions, and a welcome addition. Certainly, there has been an effort to provide modern terms and technology, e.g., a discussion of collecting insects for molecular re- search. Especially helpful is the revised format for bibliographic reading sugges- tions for specific subjects, mainly found in Part One. Part Iwo. is “composed basically of a key to the orders of insects, followed by a summary of information for each order. The authors have en- hanced this text, and ~inchidedseine derivation of the ordinal names. As orders, Protura and Diplura have been added to Steyskal et al. (1986) (in which they were omitted “due to their slight importance to agriculture.”’) Illustrations have been added to the keys, and are helpful to understand terminology and structures. VOLUME 108, NUMBER 3 Although the authors have produced an excellent and important volume, I would offer the following suggestions for consideration in future revisions. On page 101, the North American entomo- fauna is stated at over 80,000 species. This figure should be more than 90,000, and is usually noted as ““North America, North of Mexico.”” Numbers of species per order should also be updated. As an example, the number of species of Diptera “in North America” retain the Steyskal et al. (1986) figure of 18,000, but is now some 20,000. I have detected only four problems related to illustra- tions: the syrphid fly (page 189) is actually a bee fly, the horse fly and the house fly (pages 189 and 190) are the same image, the aphid (page 178) has only one pair of wings, and the ant (page 147), intended to show the petiole, does not show well the petiole as separate from the remaining portion of the abdomen. I found no mention that wasps should not be left long in killing jars due to their color changes from yellow to red, nor the hint to blow open the wings of microlepidoptera before spreading their wings. An alternative and more practical method of preparing small Diptera has become popular throughout many insti- tutions, and should be reviewed for future revisions: a square block of polyporous (or similar material) is pinned at appropriate height on a regular insect pin, with a minuten nadeln (small pin) forced into the block. This allows the collector to prepare flies in the field with the simple move of pushing the point of the ““minuten” into the right side of the specimen, beneath the wing. In labelling, many entomologists are now using three-letter abbreviations for months in labels, to avoid any confusion. Several important volumes that aid in identification of insects were not includ- ed in the bibliography. Examples are: American Beetles by Arnett et al. (2001, 2002), Manual of Nearctic Diptera by 741 McAlpine et al. (1981, 1987), How to Know the True Bugs by Slater & Bar- anowski (1978), How to Know the Grass- hoppers by Helfer (1972), Bee Genera of North and Central America by Michener, et al. (1994), and Moths of North America North of Mexico (Sphingoidea), by Hodges (1971). A few minor typo- graphical errors were found, and should be corrected for future revisions. In- cluded are: Antartica (for Antarctica) on page 100, triungulan (for triungulin) on page 181, Rhipiphoridae (for Ripi- phoridae) on page 181, angelwing (for anglewing) on page 186, tachnid (for tachinid) fly on page 189, and tephiid (for tiphiid) on page 195. This volume is a worthy addition to the entomological literature, and will certainly be widely chosen for personal and institutional bookshelves. The wealth of accurate, up-to-date informa- tion included will serve science well, and surely influence all readers into the direction of understanding and respect- ing nature. LITERATURE CITED Arnett, R. H., Jr. and M. C. Thomas. 2001. American Beetles. Vol. | Archostemata, Myx- ophaga, Adephaga, Polyphaga: Staphylinifor- mia). CRC Press, Boca Raton, FL, 443 pp. , P. E. Skelley, and J. H. Frank. 2002. American Beetles, Volume 2 (Polyphaga: Scarabaeoidea through Curculionoidea). CRC Press, Boca Raton, Florida, 861 pp. Helfer, J. R. 1972. How to Know The Grass- hoppers, Cockroaches, and Their Allies. Wm. C. Brown Co., Dubuque, Iowa, 353 pp. Hodges, R. W. 1971. The Moths of North America (Sphingoidea). Curwen Press, London, 170 pp. (the first volume in a long series with many authors). McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds. 1981. Manual of Nearctic Diptera, Vol.1. Monograph No. 27. Research Branch, Agricul- ture Canada, 674 pp. . eds. 1987. Manual of Nearctic Diptera, Vol. 2. Monograph No. 28. Research Branch, Agriculture Canada, 1332 pp. Michener, C. D., R. J. McGinley, and B. N. Danforth. 1994. The Bee Genera of North and 742 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Central America (Hymenoptera: Apoidea). Smithsonian Press, Washington, D.C. & Lon- don, 209 pp. Slater, J. A. and R. M. Baranowski. 1978. How to Know the True Bugs. Wm. C. Brown Co., Dubuque, IA, 256 pp. Steyskal, G. C., W. L. Murphy, and E. M. Hoover. 1986. Insects and mites: Techniques for Collec- tion and Preservation. United States Depart- ment of Agriculture Miscellaneous Publication. No. 1443, 103 pp. Gary F. Hevel, Department of Entomol- ogy, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 187, Washington, DC 20013-7012, U.S.A. (e-mail: hevelg@si.edu) PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 743-746 Book REVIEW Field Guide to Grasshoppers, Katydids, and Crickets of the United States. John L. Capinera, Ralph D. Scott, and Thomas J. Walker. Comstock Publishing Associ- ates, Cornell University Press, Ithaca and London. 2004. 249 pp. $45. This is the first comprehensive field guide to the grasshoppers, katydids, and crickets of the United States since Jacques Helfer’s 1987 edition of How to Know the Grasshoppers, Crickets, Cock- roaches and their Allies. Unlike that book, which also included earwigs, termites, walkingsticks, cockroaches, and man- tids, this new field guide covers only the Orthoptera sensu stricto, the saltator- ial forms (primarily grasshoppers, katy- dids, crickets, mole crickets, and grylla- cridoid families). Two of its authors, John L. Capinera (a leading authority of U.S. grasshoppers) and Thomas J. Walk- er (the leading U.S. specialist on cricket and katydid acoustical behavior and systematics) are both from the Universi- ty of Florida, Gainesville, and bring with them a wealth of knowledge of the Orthoptera from that region. Ralph D. Scott, a biologist and illustrator from grasshopper country in Montana, pro- vides one of the important aspects to the value of this book as an excellent field guide with his color illustrations of nearly 220 species. This is a beautifully produced book. The 48 color plates are so well rendered that, with a cursory knowledge of their subfamilial position, many species can be distinguished solely from the illustra- tions. The first portion of the book covers aspects of the biology of this interesting diverse order, including mor- phology, life history, feeding habits, ecology, and acoustical behavior. It goes on to present information on collecting, rearing, and photography. The remain- der of the book covers the seven major families: Acrididae, Tetrigidae, Tridac- tylidae, Tettigontidae, Gryllidae, Gryllo- talpidae, and Gryllacrididae. Situated in the central portion of the book are the 48 color plates with species information detailed in the text of the book, Whe field guide includes (1) highly detailed, full-color drawings of many_ species, including sexually dimorphic differences and color phases, when appropriate; (2) information on distributional limits, habitat preferences, ecology, behavior, and life cycles; and (3) excellent, shaded distribution maps. Although not as comprehensive in scope as Helfer’s book (which covers 643 species, compared with 348 species), this field guide presents more up-to- date information about the species that are covered. For example, regarding the grasshoppers, it pulls details from Daniel Otte’s North American Grass- hoppers, Volume 1, 1981 (Gomphocerinae and Acridinae), and Volume 2, 1983 (Oedipodinae), such as new synonymies and the most up-to-date geographical ranges. And with respect to crickets and katydids, much of what is currently known about these groups is derived directly from the excellent research of T. J. Walker. These groups were partic- ularly in need of updating, since Helfer’s (1987) treatment of them was limited primarily to the decticine (now tettigoniine) katydids of the western states. The book is not without its own set of problems, however. In searching out something to find fault with, I was initially hard pressed, and I could un- cover only a few items which deserve mention. One of the shortcomings in the grasshopper section of the field guide is the reluctance of the authors to use the 744 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON most up-to-date placement of genera into the seven subfamilies currently recognized for the United States fauna. If they had done so, they would have recognized Gomphocerinae (in which they correctly covered 23 genera and 44 species), Acridinae (again, which they correctly covered, with the single genus and species, Metaleptea brevicorne), Oe- dipodinae (another correctly annotated subfamily, in which they covered 25 genera and 72 species). But then they combine the remaining 87 species into the subfamily Cyrta- canthacridinae. With little additional effort, they could have separated these species—according to D. Otte’s Orthop- tera Species File, volume 4 (1995)—into four currently recognized subfamilies of spur-throated grasshoppers, as fol- lows: Leptysminae [in their text they presented four genera (Leptysma, Para- pomala, Prorocorypha, and Stenacris), Podisminae [four genera presented (Bra- dynotes, Paroxya, Phaetaliotes, and Poe- cilotettix)|], Cyrtacanthacridinae sensu stricto (the genus Schistocerca), and Melanoplinae sensu stricto (all remaining genera). Since these genera are distribut- ed alphabetically, it is difficult to make sense of their true relationships. For example, the leptysmine grasshoppers, all of which are called ‘“‘toothpick grass- hoppers,” are distributed on pages 71, 72, 114, and 147, and on Plates 3 and 30. They would be better understood, if all were together, both in the text and on the same plate. In addition, most researchers in North American orthopterology today recog- nize the family Romaleidae (the field guide incorporates information on five U.S. genera), but the authors place them as a subfamily [not used in the literature since J.A.G. Rehn and H.J. Grant, Jr. [Monograph of the Orthoptera of North America (North of Mexico). Mono- graphs of the Academy of Natural Sciences of Philadelphia. 1971]. Since Capinera, Scott, and Walker did present the katydids in seven subfamilies, with each subfamily being characterized with one or two short paragraphs, it seems reasonable that the same treat- ment could have been applied to the grasshoppers. To incorporate this in- formation would have extended our taxonomic awareness of this group to the persons who will undoubtedly be using this otherwise very useful text for many years to come. A second shortcoming that caught my attention involved problems with scale on most or all of the color plates. Taken individually, the figures of in- dividual species are all excellent, and indeed they are one of the major assets of the field guide. But one is unable to compare size of the related species figured on each plate, because the scale of each figure differs. To demonstrate this for myself, I randomly selected four plates (Plates: 2} 22,35; ands), measured each species figured on each plate and measured the total lengths of representative actual specimens of each species from our National Collec- tion of Orthoptera. Selecting the total length of the smallest actual species and dividing that value into the total length of each of the other species on the plate, I obtained actual size ratios. The same procedure was applied to the images on each plate. The results showed that size cannot be compared and that figures are useful only for their gestalt value. For example, Sti/pno- chlora couloniana is the largest species on Plate 36 and should be 3.7 times larger than the smallest species, Platylyra californica. However, it 1s figured as being only 1.9 times larger. Similarly, Montezumina modesta is figured as being 0.7 times as large as Stilpnochlora but in reality it is only 0.4 times as large. While plates depicting grasshoppers demonstrate fewer problems with scale than the plates depicting katydids, the VOLUME 108, NUMBER 3 inconsistencies are nevertheless distract- ing. Yet another shortcoming relates to the unequal treatment of certain orthop- teran groups. Only three species of Rhaphidophorinae (Gryllacrididae) are presented (two of them quite rare and geographically highly restricted). Nu- merous other species which could have been represented were not, although they are either quite common (Ceuthophilus maculatus, C. brevipes, Udeopsylla robus- ta) or are nuisance pests (Ceuthophilus utahensis, Tachycines asynamorus). The same problem exists for the tetrigids. Helfer (1987) reviewed 26 species; this field guide reviews only two _ species. Persons interested in using this book as a field guide to identify these groups would not be well served. Finally, I had some reservations with the authors’ treatment of common names. It prompted me to review the rules for common names and to review common names used in the literature for the species covered in this volume. I compared the names used by WS. Blatchley (Orthoptera of Northeastern America, 1922) and Helfer (1987) with those used by Capinera, Scott, and Walker. While they correctly used com- mon names previously applied on 184 species by Helfer and Blatchley, and introduced 87 new common names for species having no prior common name, they arbitrarily (and probably inappro- priately) changed common names for 31 species. For example, Aulacara elliotti (Helfer’s ‘“‘Elliott’s grasshopper’) was reassigned the common name “big-head- ed grasshopper,” but Helfer correctly applied this common name to the species Eupigodes megacephala. Gryllodes sigilla- tus, the ““decorated cricket,’ is changed in this volume to the “tropical house cricket,” and Aflanticus testaceus, the ““short-legged shield-back” katydid is changed to the ‘“Protean shieldback”’ katydid. 745 Numerous other common name changes have greater merit, in that they are more reflective of some aspect of the group. For example, the three species of the katydid genus /nscudderia previously had common names that did not reflect their close relationship to each other and to their common ecological niche: J. strigata was the “‘striped katydid” (now the “Guinea cypress katydid”’), 1. taxodii was the “bald cypress katydid” (now the ““western cypress katydid”’), and J. walk- eri was the ““Walker’s katydid” (now the “eastern cypress katydid’’). One change that was totally for the better: Boopedon nubilum, formerly the “black males grasshopper” [because only males of this species are black in color] to the ““ebony grasshopper.” Another distraction with the authors’ use of common names is the frequent misuse of the placement of the hyphen. Whereas word conjunctions such as ‘‘red-legged”’, and ‘“‘clear-winged” are employed correctly, others such as ‘“‘spurthroated” and “coneheaded” are ignored; these latter words should cor- rectly be ‘“‘spur-throated” and “‘cone- headed.” The field guide’s important contribu- tion is in its updating of biological information and providing color images of most species. The authors have made good use of contributions of Otte’s 2- volume book series and the results of years of Walker’s research on the ecolo- gy and acoustical behavior of tettigoniids and gryllids of southeastern United States. Overall, I highly recommend this book as a “must have’ for persons interested in grasshoppers, crickets, and katydids of North America. Enthusiasts of Or- thoptera with an interest in collecting these interesting insects will find this volume the important first step in rapid field identification. It also takes the avid collector into the realm of identifying these insects by the sounds 746 _ PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON they produce. Spectrograms and descrip- tions of calling songs will help the orthopterist to evaluate the species in a region, even if they are unable to locate actual specimens for their collec- tions. David A. Nickle, Systematic Entomol- ogy Laboratory, PSI, Agricultural Re- search Service, U.S. Department of Agriculture, Bldg. 005, BARC-West, Beltsville, MD 20705, U.S.A. (e-mail: dnickle@sel.barc.usda. gov ) PROC. ENTOMOL. SOC. WASH. 108(3), 2006, pp. 747-748 INSTRUCTIONS FOR AUTHORS General policy.—Articles for publica- tion are accepted from members and non- members. However, non-members pay a higher publication charge and cannot apply for a waiver of page charges. Only one author of an article needs to be a member for member rates. Articles must be in English, but an abstract in French, German, Spanish, Portuguese, Russian, and some other languages is acceptable. Such an abstract should be placed imme- diately after the English abstract and may be a translation of that abstract. Manuscripts of 15 printed pages or less are encouraged, but longer manuscripts are acceptable and considered. Publication charges for pages over 15 are at a higher rate and are never waived. About two and one-half double-spaced typewritten pages equal one printed page. Manuscripts are peer reviewed before acceptance. Final ac- ceptance is the responsibility of the Editor. 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SMITHSON Pare 3 9088 90053 2508 (Continued from front cover) McKAMEY, STUART H.—Review of the Neotropical leafhopper genus Chlorogonalia (Hemiptera: Cicadellidae: Cicadellinae), with notes on the genus Caldwelliola .......... 611 McKAMEY, STUART H.—New combinations in sharpshooter leafhoppers (Hemiptera: Cicadeliidae: Cicadellinwe yin yoy UNE mime Nel Vee Ame a: "MON aN eS Ae RE READ 672 MORON, MIGUEL ANGEL and JOSE ROBERTO SALVADORI—The third-stage larva and pupa of Demodema brevitarsis (Blanchard) (Coleoptera: Scarabaeidae: Melolonthinae) from SOlitherm ys Taz pyr 5 18 Ss eas es Re eer eller ails velit ok runny etd ok Coe dani, a Grol sen MRE Ntisar, SP Oe Open ae 511 PARDO-LOCARNO, LUIS CARLOS and MIGUEL ANGEL MORON—Description of the third-instar larva and pupa of Lycomedes hirtipes Arrow (Coleoptera: Scarabaeidae: Dynastinae: Agaocephalini) with notes on its biology and distribution in Colombia....... 661 REINERT, JOHN F.—Tewarius Reinert, a new genus of Aedini (Diptera: Culicidae).......... 639 SHANG, SUQIN, LIN SHEN, YALIN ZHANG, and HOUHUN LI—Taxonomic study on the leafhopper genus Bhatia (Hemiptera: Cicadellidae) from China ...................... 565 SHENG, MAO-LING—A new genus and species of Poemeniini (Hymenoptera: Ichneumonidae) froin Chia aie eee BOT tea ea RN Haas Or Pes 2k AON Ne ee a 651 WHITWORTH, TERRY—Keys to the genera and species of blow flies (Diptera: Calliphoridae) ofAmentcaliorthy Of MExICOES. icy. agate Bal st Ae Sree UelS typ h si tas Lae a ee CIS, oe eed TAA 689 YAMADA, KAZUTAKA, TOMOHIDE YASUNAGA, YUKINOBU NAKATANI, and TOSHIYA HIROWATARI—The minute pirate-bug genus Xylocoris Dufour (Hemiptera: Heteroptera: Anthocoridae) from rice mills inal Wailand.| 4.) )ohch oaieta k Roane Sot tok terare el oes By ene 525 YANG, DING and PATRICK GROOTAERT—Adéditions to the fauna of Drapetis Meigen (Diptera: Empididae)itrom: Chimay 0) Shs as et CRS or, eave eaputad bya RC EN a ee 677 YANG, ZHONG-QI, JOHN S. STRAZANAC, YAN-XIA YAO, and XIAO-YI WANG—A new species of emerald ash borer parasitoid from China belonging to the genus Tetrastichus Haliday: (Hymenoptera Eulophidae) ae 2 fee rok (nid eee ietet ona aiia tees (ede a aN e taNeIat Ones aN Coma ies 550 NOTES CACI, JENNIFER, MICHAEL S. KING, JERI HUMPHRIES, KRISTIN COBB, STEPHEN GARVIN, DENNIS D. KUHR, JIMMY WEDINCAMR, and ERIN STANWIX—First record of Aedes (Ochlerotatus) grossbecki (Dyar and Knab) (Diptera: Culicidae) in Alabama, U.S.A 736 HASTRITER, MICHAEL W.—A new record of the flea Tarsopsylla octodecimdentata colo- radensis (Baker, 1895) (Siphonaptera: Ceratophyllidae) in Utah ...................... 730 McCAFFERTY, W. P—New synonym and western range extension for Heterocloeon anoka (Dagsy) (Ephemeroptera:/Bactidae) is) xiiei oa Sane D os cea ert naa LAD Ue aaa aa ea 738 POINAR, GEORGE, JR. and ALEX E. BROWN—Remarks on Parvaverrucosa annulata (= Verrucosa annulata Poinar and Brown 2005) (Hemiptera: Sternorrhyncha: Aphidoidea) ... 734 REEVES, WILL K., AUBREY G. SCARBROUGH, and JOHN W. McCREADIE—New records for Leptopteromyia americana Hardy (Diptera: Asilidae) in Alabama.................. 739 ROBBINS, RICHARD G., BUI DANG PHONG, TIM McCORMACK, JOHN L. BEHLER, HENDRIK A. ZWARTEPOORTE, DOUGLAS B. HENDRIE, and PAUL P. CALLE—Four new host records for Amblyomma geoemydae (Cantor) (Acari: Ixodida: Ixodidae) from cap- tive tortoises and freshwater turtles (Reptilia: Testudines) in the Turtle Conservation Center, Cuc'Phuong National Park Vietnariays oise0y pute ahaa levee Chey ars veh ame (aan eel ea 726 SMITH, DAVID R.—Types and voucher specimens of New World Aulacidae (Hymenoptera) in the Naturhistoriska Riksmuseet, Stockholm::Sweden = )) 20! sys Gen ahs eee ealebeke 732 BOOK REVIEWS HEVEL, GARY F—4rthropod Collection and Identification, Laboratory and Field Techniques, by, Timothy: JGibbiand Christian) Oseto Po yes 2 crake oto hice mn naire ne het era mae 740 NICKLE, DAVID A.—Field Guide to Grasshoppers, Katydids, and Crickets of the United States, by John L..Capinera; Ralph DY Scott; and: Thomas) Walker: 2 sisi). eevee eee 743 Instructions. for Authors) feo fey s. ce PR iu Bie LSE oe AUR ora Me ARN aC Ae a, TAT VOL. 108 OCTOBER 2006 NO. 4 (ISSN 0013-8797) OL ‘| PROCEEDINGS ENT of the | PUBLISHED QUARTERLY CONTENTS BOOTH, JANIE M. and PENNY J. GULLAN—Synonymy of three pestiferous Matsucoccus scale insects (Hemiptera: Coccoidea: Matsucoccidae) based on morphological and molecular SSUALG SLAVS AA Te OT Sk RNR EGE on Be OA nr oat he ot, nS MN, a Na. Pa Ogee MRO 9 NG 749 BROWN, JOHN W—A new species of Cochylis (Lepidoptera: Tortricidae: Cochylini) from Argentina: A potential biocontrol agent against pompom weed (Asteraceae)............ 899 CONLE, OSKAR V., FRANK H. HENNEMANN, and DANIEL E. PEREZ-GELABERT— Studies on Neotropical Phasmatodea III: A new species of the genus Anisomorpha Gray, 1835 (Phasmatodea: Pseudophasmatidae: Pseudophasmatinae) from Hispaniola ......... 885 CONTRERAS-RAMOS, ATILANO—A > new species of Chloronia Banks (Megaloptera: Cor dalidac) dion Bolivia AnGIRSKU ssa ce 0s) G/T eI NN ROR GN SUNN) AL CDN Nien Een a we 808 CONTRERAS-RAMOS, ATILANO—Protosialis ranchograndis, a new species of alderfly from Venezuela, with a redescription of P brasiliensis Navas (Megaloptera: Sialidae) ......... 977 FRITZLER, CYNTHIA J., CRYSTAL B. MAYLE, JOHN S. STRAZANAC, and LINDA BUTLER—Diversity of Syrphidae (Diptera) in central Appalachian forests............. 964 GAGNE, RAYMOND J. and JEAN ETIENNE—Gephyraulus mangiferae (Felt), n. comb. (Diptera: Cecidomyiidae): A mango pest from India newly recorded from the Western TETSU EL S| PAIR Pea ania RETR GNSS eet teen at See eens Cue rue cae aM ei vest ere ee 930 GRISSELL, E. E.—Two new species of Torymoides Walker (Hymenoptera: Torymidae) from the Amenicamsomtnwest,,withia keyito Nearctic Speeiesta mito jl fo is ays ie miei apenas are eink 765 GRISSELL, E. E. and DAVID R. SMITH—First report of Acroclisoides Girault and Dodd (Hymenoptera: Pteromalidae) in the Western Hemisphere, with description of anew species 923 HENRY, THOMAS J.—Resurrection of the plant bug genus Pappus Distant, with clarification of Mea species (Hemiptera, meteroptera: MiniGae),..12 0 ase ae. eign ls cc Lib eiete eevelag ators 822 HENRY, THOMAS J. and A. G. WHEELER, JR.—Redescription and hosts of Melanorhopala infuscata Parshley, with notes and new distribution records for M. froeschneri Henry and Wikeelertienipterasieteroptera: Linfidae iol). Sandel hie eo eees Sieuaes Uhstee e cue lela eee elle. = 917 HINOJOSA-DIAZ, ISMAEL A., ELISA BONACCORSO, and MICHAEL S. ENGEL—The po- tential distribution of Zorotypus hubbardi Caudell (Zoraptera: Zorotypidae) in North America, asipredictcd by ecolosical mishe modeling sis 2/ bea F ie.v sab pascal s\creme oN, be wlcld's geata he ee 860 HUANG, MIN and YALIN ZHANG—Systematic study of the Oriental genus Vatana Dworak- owska (Hemiptera: Cicadellidae: Typhlocybinae: Typhlocybini)...................-... 945 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2006 STEVEN W. LINGAFELTER, President Jon A. LEwIs, Custodian MICHAEL W. GaTEs, President-Elect MICHAEL G. POGUE, Treasurer STUART H. McKamey, Recording Secretary JOHN W. BROWN AND DavID G. FurTH, Program Chairs HOoLtis B. WILLIAMS, Membership Secretary JASON P. W. HALL, Past President DAvip R. Smitn, Editor Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY WAYNE N. MATHIS Honorary Members WILLIAM E. BICKLEY DAVID R. SMITH RONALD W. HODGES PAUL J. SPANGLER MANYA B. STOETZEL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168. MEETINGS.— Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu- tion, on the first Thursday of each month from October to June, inclusive, at 7:00 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. An- nual 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 ad- dress changes to the Entomological Society of Washington, % Department of Entomology, Smithsonian Institu- tion, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomological 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. 1027-028 OF THIS ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Wash- ington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 12 October 2006 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 108(4), 2006, pp. 749-760 SYNONYMY OF THREE PESTIFEROUS MATSUCOCCUS SCALE INSECTS (HEMIPTERA: COCCOIDEA: MATSUCOCCIDAE) BASED ON MORPHOLOGICAL AND MOLECULAR EVIDENCE JANIE M. BOOTH AND PENNY J. GULLAN Department of Entomology, University of California, 1 Shields Avenue, Davis, CA 95616-8584, U.S.A. (e-mail: pjgullan@ucdavis.edu) Abstract.—The scale insect genus Matsucoccus Cockerell (Coccoidea: Matsucoc- cidae) contains several economically important species that cause damage to pine trees, Pinus species, in the United States and elsewhere in the Holarctic Region. Efforts to reconstruct the phylogeny of the group have provided information on genetic variation within and among species. Here, three species of Matsucoccus are synonymized based on newly acquired molecular data and reassessment of morphological data. Matsucoccus resinosae Bean and Godwin, described from the eastern United States, and Matsucoccus thunbergianae Miller and Park, from South Korea, are considered to be new synonyms of Xylococcus (now Matsucoccus) matsumurae Kuwana. The taxonomic confusion surrounding these names _ is discussed. In addition, we suggest that several other species of Matsucoccus, including M. pini Green, should be investigated as possible synonyms of ™. matsumurae. Key Words: Margarodidae, Matsucoccus, Matsucoccus Cockerell is a morphologi- cally conservative group of scale insects placed either in the scale insect family Margarodidae (Ben-Dov et al. 2005) or in its own family, Matsucoccidae (Koteja 1984, 1986; Foldi 2004). All species feed exclusively on Pinus (Pinaceae) and are mainly Holarctic, with limited records from the Neotropical and Indotropical regions (Ben-Dov 2005, Ben-Dov et al. 2005). This genus has received moderate taxonomic treatment (Ray and Williams 1984, 1991; Gill 1993; Foldi 2004), and 34 extant species are recognized (Ben-Dov 2005). Recent research to reconstruct the phylogeny of Matsucoccus has provided the first hypothesis of species relation- ships based on molecular and morpho- logical data (Booth, Cook and Gullan, red pine scale, taxonomy unpubl. data). One relationship of par- ticular interest jis (thattiofi sthree. pest species, Matsucoccus matsumurae (Ku- wana), the Japanese pine bast scale, M. resinosae Bean and Godwin, the red pine scale, and M. thunbergianae Miller and Park, the black pine bast scale. New evidence supports the synonymy of the three species. Entomologists have specu- lated that M. matsumurae and M. resinosae are synonymous (Ray 1982; McClure 1983b, 1987; Foldi 2004). The recent catalogue of Margarodidae (Ben- Dov 2005) treats these two species separately, but mentions previous work suggesting that M. resinosae may be a junior synonym of M. matsumurae (McClure 1983a, Young et al. 1984, Park et al. 1986). 750 Matsucoccus resinosae infests red pine, Pinus resinosa, on the east coast of the United States. Red pine scale was first recognized in 1946 in Easton, Connecti- cut (Plumb 1950), and its rapid spread and the high tree mortality that it caused suggested that 1t was a new introduction (Bean and Godwin 1955). It has been hypothesized that it was introduced during the 1939 New York World Fair on exotic pines imported as a display, because the same truck that transported the exotic pines from the port was used to transport red pines from Easton to the fairgrounds (Doane 1959). The feeding cyst stage of M. resinosae can damage the needles, causing extensive flagging and needle drop (McClure 1976, Duda 1977). Damage is particularly severe in plantations found south of red pine’s native range (Bean and Godwin 1955), apparently because low winter tempera- tures in the pine’s natural range to the north prevent survival of red pine scale nymphs (Doane 1959; McClure 1983a, b). Damage from the red pine scale has resulted in almost complete removal of once abundant red pine plantations in Connecticut and New York states and it is now difficult to locate intact stands (J. Booth pers. observation, Providence Water Supply Board 2005). The Japanese pine bast scale, M. matsumurae, has similar biology and morphology to M. resinosae (McClure 1976, 1983a). This scale is a pest in Asia, especially on black pine, Pinus thunbergii (Taketani 1972, Cheng and Ming 1979). The species was described originally by Kuwana as Xylococcus matsumurae (Ku- wana 1905, 1907), and later transferred to Matsucoccus by Cockerell as the type species of his new genus (Cockerell 1909). The third species, M. thunbergianae Miller and Park, from South Korea, was described as similar to M. matsumurae and M. resinosae (Miller and Park 1987). if is not possible to distinguish M. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON thunbergianae from M. matsumurae and M. resinosae, based on the adult females or the first-instar nymphs, but ™. thunbergianae is said to differ from the other two species in the size of the adult male, in the number of generations per year, and in overwintering as the second- instar nymph (as the first-instar nymph in the other two species). Matsucoccus thunbergianae is univoltine or has only a partial second generation (Miller and Park 1987), whereas M. matsumurae and M. resinosae are bivoltine or have a partial third generation (McClure 1977, Miller and Park 1987). Matsucoc- cus thunbergianae 1s considered a pest 1n Korea on black pine (Miller and Park 1987, Chung et al. 2000). Several available pieces of biological evidence support the synonymy of these three Matsucoccus species. First, the sex pheromones of MM. matsumurae, M. thunbergianae, and M. resinosae have been shown to be cross attractive in bioassay studies (Young et al. 1984, Park et al. 1986). The primary component of the sex attractant was identified and the pheromone is identical for the three species and has been named “‘matsuone”’ (Lanier et al. 1989, Hibbard et al. 1991). Second, all species occur on hosts found in the same Pinus subsection, subsection Pinus, of the pine tree phylogeny of Gernandt et al. (2005). Matsucoccus matsumurae occurs on seven Pinus spe- cies (McClure 1983a), Matsucoccus thun- bergianae only on P. thunbergii and P. densiflora (Miller and Park 1987), and M. resinosae is found on P. resinosa (Kuwana 1905, Bean and Godwin 1971, Miller and Park 1987), which is the only species of the Pinus subsection Pinus in the United States (Gernandt et al. 2005). Ray (1982) and McClure (McClure 1983b) went further and noted that M. resinosae and M. matsumurae are more specifically found only on members of the Sylvestres group of the subsection Pinus. However, the Sylvestres group 1s VOLUME 108, NUMBER 4 Table I. Til Specimens of adult females examined for morphological analysis. SSS Species Name #- Slides (# Females) Collection Information Depository M. matsumurae 1 (1 non-type) JAPAN: Kanagawa-ken, ex pine tree, 13.v.1919; BME Coll. S. I. Kuwana (mounted from dry material sent by Kuwana to F. B. Herbert) 1 (2 non-type) JAPAN: Nagashima, ex P. densiflora, 10.v.1970; BME Coll. M. Inoure 2 (2 non-type) JAPAN: Nagashima, ex P. densiflora, 4.v.1970; USNM Coll. M. Inoure 3 (4 non-type) JAPAN: Japan: Mie Prefecture, Shimagahara BME Village, ex P. thunbergii; 24.11.2004; Coll. T. Kondo M. pini 3 (5 paralectotypes) ENGLAND: Oxshott, Surrey, ex P. sylvestris, BMNH 31.x.1922; Coll. F.C. Withycombe M. resinosae 3 (holotype and 2 USA: Connecticut, Easton, ex P. resinosa, USNM paratypes) 2.v.1948; Coll. George H. Plumb M. 4 (holotype and 5 SOUTH KOREA: Kohung, Chollanam-do, USNM thunbergianae paratypes) ex P. thunbergiana (now P. thunbergii), collected x1i1.1983, lab reared iv.1984; CollNSAE* Park 1 (2 paratypes) SOUTH KOREA: Kohung, ex P. thunbergii, BME collected xi1.1983, lab reared iv.1984; Coll. Sa@a bank not recognized in the most current Pinus phylogeny (Gernandt et al. 2005). Third, the distribution of M. matsumurae, M. resinosae and M. thunbergianae supports their synonymy. McClure (1983b) point- ed out that the first two species are restricted to a similar northern latitudi- nal limit: 41°50’N in the U.S., 41°30’N in Japan, and 41°30’N in China. Similarly, M. thunbergianae has been reported only from South Korea (Miller and Park 1987), for which the northern boundary lies at about 39°N. Here we present the first molecular data and reassess the morphological evidence to show that specimens de- scribed as M. matsumurae, M. resinosae and M. thunbergianae belong to the same species. We synonymize M. resinosae and M. thunbergianae under the senior syno- nym M. matsumurae and provide a new diagnosis for this species. MATERIALS AND METHODS Morphology.—Morphological charac- ters were evaluated and measured using a Leica compound microscope. Type material was examined for M. resinosae and M. thunbergianae, whereas subse- quent material collected by the original author was studied for M. matsumurae (see below under “‘Type material”; also Table 1). Specimens from each collection were scored for 14 morphological char- acters (Table 3), including those previ- ously recognized by Ray (1982). The diagnosis was prepared based on the specimens listed in Table 1. Specimens examined are housed at the Bohart Museum of Entomology (BME), Uni- versity of California, Davis; The Natural History Museum, London (BMNH); and the Coccoidea Collection of the National Museum of Natural History, Smithso- nian Institution (USNM) in Beltsville, Maryland. Genetic analysis.—Molecular data were acquired for 13 specimens belong- ing to six named Matsucoccus species that the authors field collected or that colleagues donated (Table 2). Specimens were stored in 75% ethanol for slide mounting and 100% ethanol for molecu- lar work. Genomic DNA was extracted PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON AVA FOF SHMSD MsBAIGUNY] “q ayeway yNpe ALEr.9T1I “N, IO. SOOT Hp ‘opureurloyyD ‘Aud nlexy :va1oy yINos IZOGING avunisiaquny) “Py Joe ‘OD MD -pOOT HILT -Speoy 2[qq0D pue urejunoj| DSOUISIAA “q ayeuUlay yNpe M,61co€L ‘N,10 Japug, Jo uonounl ‘ueeuryd :oD preyyouT LO ‘vsa OCOAWEL IDSOUISAA “A Ioog “f TOD “pOOT XS Feary es eidniny DSOUISAA “q ysAO M,ProEl “NLS. apLAdnadvjey ‘apiaaadvjeyT 20D ssoysing AN “WSN STOAWE IDSOUISAA “J BuldsurA “XK TOD ‘SOOT AST *RUIYD UBAIGUNY] ‘q ysAO Ay leSGl NFO JSBOYION SSOUIAOIgG Ulf Ul AJUNOD Suoji,A :vuryD LPOAWL avanuinsypiut wy opuoy “L MSO -pooc Wye 11s4aquny] “q a]euldy yNpe AL€0.9€1 ‘N,9P SOBR][IA VILYRSCUNIYS ‘oINOIJoIg Ap :uedee FIOPING avaniuinsjpiul “Py JOP “Cs pue yoog “f “S[IOD -SOOT HES DUDIUIBAIA “ql ss30 N,£S.9L “N,10 SAUW[IOB] INNS “OPPASIag :OD .ses10ay s9UuLIg CW 9COaWL SNOIYIDS “WW y100g “f MOD -pOOT XE! -peoy peotyioary DPB ‘ ys M,6€.CL N.SS SOYSHOY ISVY “PRsysIATY :OD ALOJNS AN poouWer SnJOIN]DS “W UIaITID) “CM MOD -v00c XV OT DpIsld o[eulo} y[Npe M.8P0CL “N.CSo ‘OL WxO “Cop AMP “9][IATOURI :OD YLOHNS AN tcOaWe SNJOIIVS "W YIOOg “f [OD {POOT' A] {4191U9D S.AOUSIA 189104 119]]NOD ‘dq syduiAu NM,STo9IT (NZS oZE [PUONEN purpaagyD ‘vunse Til 70D osaiq ures VO 6cOAIWL Snsojasiq “W 4100 “f TOD -POOTAT6 Aled psosapuod ‘q S330 NM, COcIZI “N.ZI a1e1g soul osidurgq ‘AaT[@A SSBPID 70D BPRAON WO 9TOAIWL SNSOJASIG “JV opuoy \L pue yioog “f “SOD ‘POOT ANT “PU o8Pray vsosapuod ‘q sydwiAu M,9ScITI “Ns 8To UMOJI[SUIYS ppTST “UMO}JI[SUIYS 70D vIseYS WO €lLOAWe SNSOJASIG “We yycydouow ‘gq syduicu M,9S.811 “N6b.PE AIPM “CF WOD 1007 L AW4ed Jozery sop UE WD OOM INE SNIdAJDID “PW yIoog “f TOD ‘SOOT HEOE +68 AMH JO JJO SIMA Gl o[euloy yNpe M.IToCIT NuCbove ‘sstq UIA SNBUTPY JO AA TU C*] OD redearyx ZY ScOdWN snidAjpon “Py yIoog “fF TOD *sOOTHEOE -A9TIPA TOMAS synp2 “d a]RUldy Npe NM SEoTIT (NU 6ToVE jo q sojiw 9 ‘py uiseg toddoad :oD iedearx ZV LEOUWe sniddjpov “We JSOH{ Snug aBeIS IIT so] RUIpIOOD UONBWIOJUT UOTIDAT[OD 2P9D VWNA auRN saroedg “SISATBUR IP[NSITOU IOJ pasn suoumtdedg "~ FQuLl VOLUME 108, NUMBER 4 using a Qiagen DNeasy® kit (Qiagen Inc., Valencia, California, U.S.A.). DNA was extracted non-destructively in many sam- ples so that the cuticle could be saved for identification. In the remaining cases, a dead adult female found in association with the other life stages used for DNA extraction was preserved and mounted for identification. Identification was per- formed using published keys and descrip- tions (Kuwana 1905, Bean and Godwin 1955, Ray 1982, Miller and Park 1987, Foldi 2004) supported by knowledge of each species’ known distribution and host-plant(s) and, in the case of ™. thunbergianae, by the authorative identi- fication of Seung-Chan Park who was one of the describers of this species (Miller and Park 1987). Standard methods for scale insect DNA analysis were utilized for molecular work (Cook et al. 2002, Downie and Gullan 2004). Targeted DNA sequences (Table 4) were obtained using Polymerase Chain Reaction, gel agarose DNA visualization, and auto- mated DNA sequencing at the UC Davis Division of Biological Sciences DNA Sequencing Facility. Sequences were edi- ted in Sequencher version 4.0.5 (Gene Codes Corp, Ann Arbor, Michigan, U.S.A.) and aligned in Se-Al (Rambaut 1996). PAUP* (Swofford 2003) was used for determining pairwise differences among species of Matsucoccus. As part of a larger phylogenetic project, sequence data from two nuclear ribosomal genes were analyzed (Booth, Cook, and Gullan, unpublished data). The markers exam- ined were the small subunit ribosomal gene (SSU rDNA or 18S) and the D2, D3 and D10 expansion regions of the large subunit ribosomal gene (LSU rDNA or 28S) for a total of approximately 2,070 base pairs. These sequences were aligned and compared to assess the amount of genetic difference between purported spe- cies. Polymorphic sites within individuals were identified by examining electropher- ograms in Sequencher 4.0.5. Electropher- YS) ograms of individuals within species were aligned in Sequencher 4.0.5 and examined to identify variable sites and indels. RESULTS AND DISCUSSION Molecular evidence supports the syn- onymy of M. matsumurae and M. resinosae with the new inclusion of M. thunbergianae. Morphological analysis corroborates this information - no dis- cernible consistent morphological differ- ences are observed. Nucleotide sequence data.—Matsu- coccus matsumurae, M. resinosae, and M. thunbergianae had identical sequences for thel8S and 28S D10 regions. The D2—D3 region of 28S revealed a total of four polymorphisms among these three species. This amount of divergence is similar to that seen in other Matsucoccus species in terms of polymorphic sites or intraspecific divergence, as exemplified by M. acalyptus Herbert, M. bisetosus Morrison, and M. gallicolus Morrison (Table 5). A pairwise difference compar- ison showed a maximum of 0.1% se- quence difference in the D2—D3 region of 28S between M. matsumurae, M. resino- sae, and M. thunbergianae (Table 6). This is less than the 0.8% recorded within M. acalyptus, 0.7% within M. bisetosus and 0.9% within M. gallicolus (Table 6). Maximum parsimony recon- struction revealed no phylogenetic struc- ture among the individuals of M. matsu- murae, M. resinosae, and M. thunber- gianae. However, the clade containing M. matsumurae, M. resinosae and M. thunbergianae had 100 percent bootstrap support and Bayesian posterior proba- bilities value of 100 (Booth, Cook, and Gullan, unpublished data). Morphology.—Many — coccidologists have treated M. matsumurae and M. resinosae as synonyms based on mor- phology and life history data (Herbert 1921,” Morrison’ "1928; “Ray, 1932. McClure 1983b, Kosztarab 1996) but no formal synonymy has been published 754 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Morphological features of adult females examined for cladistic analysis. Morphological Characters Character States Size Body Shape (a) under 3 mm; (b) typically larger than 3 mm (a) elongate-ovoid; (b) ovoid with two parallel lobes; (c) club shaped Dorsum Bilocular tubular duct distribution Cicatrix bands Cicatrix diameter Antennal segmentation Segment 5 of antennae with 1—2 fleshy setae Long trochanter setae Long setae near coxae Long setae midventrally on abdominal segments V—VII Abdominal spiracles Multilocular disk pores Setae in marginal abdominal bands of bilocular ducts (a) on apical part of abdomen; (b) in rows on entire body (a) absent; (b) 1-4 bands; (c) =5 bands (a) <9 um; (b) 9-20 um; (c) =25 um (a) absent; (b) reduced; (c) fully developed (a) 2—3; (b) 4-8: (c) 9 (a) absent; (b) present (a) 1 long setae; (b) 2 long setae; (c) 0 setae (a) absent; (b) present (a) absent; (b) present (a) 3 pairs; (b) >3 pairs (a) absent; (b) present (a) absent; (b) present (see Ben-Dov 2005). Below we formally synonymize these three names. No dis- cernible and consistent morphological differences were observed among speci- mens identified as M. matsumurae, M. resinosae, and M. thunbergianae. A re- view of the morphology of the adult females for a larger cladistic study (Booth, Cook and Gullan, unpublished) shows the three species to be identical based on 14 characters, including pore types, antennal morphology, and setal characters, typically used to distinguish species of Matsucoccus (Table 3). The Table 4. description of M. thunbergianae recog- nizes this similarity (Miller and Park 1987), but notes that the main differences lie in the morphology of the adult male and several biological characteristics, as we explained in the introduction. The morphological differences among the males of the three species are primarily size differences. However, there is sub- stantial overlap among the three species in the size ranges for all morphological features measured by Miller and Park (1987), including: penial sheath length, aedeagus length, length of antennal Genes and associated primers used for molecular analyses. Gene Region Primer Sequence (5’—3’) Primer Name Primer Source 18S 24-585 28S D2-D3 expansion region D10 expansion region CACAATGATAGGAAGAGCC CTGGTTGATCCTGCCAGTAG CCGCGGCTGCTGGCACCAGA GAGAGTTMAASAGTACGTGAAAC TCGGARGGAACCAGCTACTA GAATGGATTAACGAGATTCTCAA 18S—2880 Tautz et al. (1988) 18S—B von Dohlen and Moran (1995) $3660 Dowton and Austin (1998) A335 Whiting et al. (1997) None Modified from Dietrich et al. (2001) None Dietrich et al. (2001) VOLUME 108, NUMBER 4 Table 5. ~) Nn Nn Genetic differences among species for DNA sequences 18S and 28S. rr Number of Polymorphic Sites within Individuals Species M. acalyptus JMBO037 0 JMBO038 JM B040 0 M. bisetosus JMBO13 0 JMB026 JMBO029 0 M. gallicolus JMBO023 0 JMBO024 JMB036 0 M. matsumurae; M. resinosae; M. thunbergianae (matsumurae ) JMBO014 JMB047 (resinosae) JMBO025 JM B030 (thunbergianae) JMBO21 — Do —S — — Number of Polymorphic Sites within Species 5 sites + two indels 6 sites; no indels 1 site: no indels 4 sites: no indels segments II—X, hind femur length, hind tibia length, forewing length, ratio of length of femur/length of tarsus, and length of longest tubular duct on ab- dominal segment VII. Bean and Godwin (1955) differentiat- ed adult females of M. matsumurae and M. resinosae based on life history and a subtle morphological disparity con- cerning the position at which the trachea enters the thoracic spiracles in the in- termediate or cyst instar (Bean and Godwin 1955). Herbert (1921) had re- described M. matsumurae based on both Japanese and American material. How- ever, Herbert based his description partly on American material collected by Mr. J.G. Sanders from the host species P. rigida and P. virginiana. These pines are both known hosts of M. gallicolus, and not hosts of M. resinosae (Ben-Dov 2005). As a consequence of this confu- sion with M. gallicolus, Morrison (1928) even suggested that M. matsumurae may be indigenous to the Atlantic seaboard of the USA. Later, Morrison (1939) recog- nized this error and, in his original description of M. gallicolus, noted: ““This is the insect which was figured and described (in part) by Herbert under the name matsumurae.” Ray (1982) also lists Herbert’s M. matsumurae as a syno- nym of M. gallicolus. This information was not included in the recent catalogue of the Margarodidae (Ben-Dov 2005). Tang and Hao (1995) questioned the species concept of M. matsumurae used by American authors, partly because of the confusion created by Herbert’s (1921) and Morrison’s (1928) mixing up of M. matsumurae and M. gallicolus, and also because they surmised that there may be two species of Matsucoccus in Japan. Tang and Hao (1995) suggested that M. thunbergianae might be synony- mous with Kuwana’s (1905, 1907) con- cept of M. matsumurae, and that M™. resinosae and M. liaoningensis Tang may be synonyms. They based their argument on purported differences in biology and adult body size of the two species pairs, but they were selective in their use of morphological data and their assertions cannot be supported. In addition, certain morphological features in Matsucoccus can vary in response to environmental conditions. For example, Miller and Park (1987) 756 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 6. Pairwise differences table (uncorrected): D2—D3 region of 28S. M. M. M. M. M. M. M. M. acalyptus acalyptus — acalyptus —acalyptus —_acalyptus bisetosus __ bisetosus bisetosus (JMB002) (JMB037) (JMBO038) (JMBO039) (JMB040) (JMBO013) (JMB026) (JMBO029) M. acalyptus (JMBO02) M. acalyptus (JMB037) 0.001 M. acalyptus (JMBO038) 0.001 0.000 M. acalyptus (JMBO039) 0.004 0.003 0.003 M. acalyptus (JMB040) 0.007 0.006 0.006 0.008 M. bisetosus (JMBO13) 0.111 0.109 0.111 0.107 0.112 M. bisetosus (JMBO026) 0.109 0.108 0.109 0.106 0.110 0.000 M. bisetosus (JMBO29) 0.115 0.114 0.116 0.112 0.116 0.007 0.007 M. gallicolus (JMBO023) OMB7 0.137 0.136 0.133 0.136 Osmy, 0.117 0.118 M. gallicolus (JMBO24) 0.129 0.128 0.127 0.124 0.128 0.111 0.111 0.114 M. gallicolus (JMB036) 0.129 0.128 0.127 0.124 0.128 0.111 0.111 0.114 M. matsumurae (JMB047) 0.097 0.095 0.097 0.094 0.098 0.032 0.031 0.037 M. matsumurae (JMBO14) 0.098 0.097 0.098 0.095 0.100 0.034 0.032 0.038 M. resinosae (JMBO25) 0.099 0.097 0.098 0.095 0.100 0.034 0.033 0.038 M. resinosae (JMB030) 0.097 0.095 0.097 0.094 0.098 0.033 0.031 0.037 M. thunbergianae (JMBO021) 0.099 0.098 0.099 0.096 0.100 0.034 0.033 0.039 M. M. M. M. M. M. M. gallicolus gallicolus gallicolus matsumurae — matsumurae resinosae resinosae (JMB023) (JMBO024) (JMB036) (JMB047) (JMBO14) (JMBO025) (JMBO030) M. gallicolus (JMBO24) 0.007 M. gallicolus (JMB036) 0.009 0.000 M. matsumurae (JMB047) 0.115 0.108 0.108 M. matsumurae (JMBO14) 0.117 0.110 0.110 0.000 M. resinosae (JMBO25) 0.115 0.108 0.108 0.001 0.001 M. resinosae (JMBO30) 0.116 0.108 0.108 0.000 0.000 0.000 M. thunbergianae 0.118 0.111 0.110 0.000 0.000 0.001 0.000 (JMBO21) examined the overwintering and summer 1952, Foldi 2004). Boratynski (1952) generations of both M. matsumurae from China and M. resinosae from the U.S.A. and showed that the adult females of the overwintering generation of both species had more multilocular pores and larger cicatrices than the summer populations (Miller and Park 1987). Furthermore, morphological plasticity in body size and additional features has been observed for other species of Matsucoccus (Boratynski 1952, Ben-Dov 1981). A fourth species of Matsucoccus, M. pini (Green), is found on a member of the Pinus subsection, namely Pinus sylvestris (Green 1925). This species is distributed throughout Europe and differs only subtly from M. matsumurae (Boratynski indicates in his published key to the genus that the main difference between M. matsumurae and M. pini is the width of the dorsal cicatrices, the number of peripheral loculi in the multilocular pores, the host tree, and the country of origin. Foldi (2004) states that ™. matsumurae and M. pini differ in the length of the bilocular tubular ducts and the number of multilocular pores at the apex of the abdomen. We examined the paralectotype females of M. pini housed at the BMNH and their morphology falls within the range of variation of ™. matsumurae. Foldi (2004) states that the average body size for M. pini is 2.8 mm long, which is less than we recorded for VOLUME 108, NUMBER 4 the specimens of M. matsumurae that we measured, however, Foldi also provides a greater body size range for M. matsu- murae (2.5—4.5 mm long). No specimens of M. pini were avail- able for molecular analysis, but given the evidence of seasonal plasticity in size of cuticular features in Matsucoccus, the synonymy of M. pini with M. matsu- murae seems likely. Other species that should be examined in this context are the Chinese species M. dahuriensis Hu and Hu described from P. sylvestris var. mongolica (Hu and Hu 1981), M. liao- ningensis ex P. tubulaeformis (Tang 1QIS)S Me. “yunnanensis. Ferris’ *ex € > 300 SE Se20 — ee 6 5 o's 2 2 =a 200 = 5 co Be @ £10 ogi =h6 100 Ree = a 0 0 tee AGN eee eACSen@) INerD MONTHS —a&— Temperature —@— Precipitation Fig. 1. Monthly population levels, in numbers of individuals, for each of the four Aphodiinae species found in cow and horse dung samples in El Llano de las Flores, Oaxaca, January—December 2003; and mean monthly temperature and total monthly precipitation (data taken from the clima- tological station of San Juan de Atepec, Oaxaca, provided by the National Water Commission). Over the course of the study year, the area under study experienced alternating dry and rainy periods. Dry periods occurred from January to May and November to December, corresponding HUY 100 INDIVIDUALS (number) o oO 0 iF MAM Jd J AS ON D —#— O. chevrolati retusus 100 7 2 E a = 4 50 <= = Qa 2 a Zz 0 Jew AM J J AS © N. B —¢— O. viridiobscurus 30 400 > € >O 300 SE eZ csi eg Es $2 200 = = © oO Ss @ 2 10 2 5 =5 100 = i a 0 0 J MAN IM) UP I vA\ Si Oe Neb MONTHS —&— Temperature —@— Precipitation Fig. 2. Monthly abundances of individuals of Onthophagus chevrolati retusus (Scarabaeinae) and Onthotrupes viridiobscurus (Geotrupinae) found in samples of cow and horse dung from El Llano de las Flores, Oaxaca, January—December 2003; and mean monthly temperature and total monthly precipitation (data taken from the climatological station of San Juan de Atepec, Oaxaca, provided by the National Water Commission). to winter and spring. During the study year, mean temperature was 20°C, al- though from March to May mean monthly temperature was almost 23°C. 778 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Number (n) of individuals of each dung beetle species collected in 60 samples of cow dung and 60 samples of horse dung, in El Llano de las Flores, Oaxaca, during the months of the year when the species were active. The affinity of each species with each dung type (as determined by Student’s /-test) is also indicated. Months of Activity Dupe Species (number) Cattle (n) Horse (n) Student’s ¢ test Planolinus vittatus iil 1,055 183 tm—1) = 4.30; P < 0.001 Gonaphodielus opisthius i PIS\E WS) tm—1) = 0.88; P = 0.40 Cephalocyclus durangoensis 2 49 330 ta—1) = —1.03; P = 0.32 Liothorax levatus 3 4 160 tm—1) = —1.25; P = 0.24 Onthophagus chevrolati 6 178 134 too LE2ZTR 10205 Onthotrupes viridiobscurus 5 19 26 fa) = = 083 2a 10105 Total precipitation for the dry season was just 90 mm; in fact, in February and March it did not rain at all. The rainy season from June to October occurred during the hot, humid summer and autumn; during these months, mean temperature was approximately 19°C. Total precipitation for the rainy season was 799 mm, with June and September showing the greatest precipitation (Figs. 1-2). During the rainy season, most of the species studied were active on the sur- face. The highest population levels of P. vittatus, G. opisthius, O. viridiobscurus, O. chevrolati retusus, and C. durangoensis were all seen during this season. The one exception was L. levatus, a species that was active only at the beginning of the dry season, but while the soil was still humid (Figs. 1, 2). Otherwise, during the dry season, only P. vittatus was found, and only in small numbers. O. chevrolati retusus was also found at the beginning of the dry season, but not in April or May, when precipitation is very low and temperature is high (Fig. 2). Species trophic preferences.—The to- tal number of individuals obtained were just those encountered during the months the species were active (Table 2). Apparently, P. vittatus prefers cow dung, and C. durangoensis and L. levatus prefer horse dung. It may be that G. opisthius, O. chevrolati retusus, and O. viridiob- scurus have no preference for either of the two kinds of manure. Nevertheless, it was only for P. vittatus that a statistically significant preference was seen, for cow dune @i_1 = 4.30; P =0.001, 277-9ie 62.19) (Table 2). Species reproductive activity.— Plano- linus vittatus: Female P. vittatus have 5 ovarioles in each ovary. No immature females were seen, that is, females without oocytes in their ovarioles. The ovarioles of maturing females had from one to three oocytes, with basal oocytes mature, with chorion, before oviposition. Mature oocytes before oviposition mea- sured 0.83 + 0.07 mm in length. After Oviposition, immature basal oocytes without chorion were small (Table 3). Males have 7 testis follicles, 2 larger than the other 5; in’ cachsot the 2=testicles Testis follicle length did not vary much with the state of sexual maturity. In contrast, the volume of glandular secre- tions stored in the two glandular reser- voirs, representing the greatest part of the seminal liquid of the spermatophore, varied significantly depending on sexual maturity. Few immature males of these species were seen as well, that is, those without secretions in their glandular reservoirs. Males at the beginning of maturation showed the smallest glandu- lar reservoirs. Mature males before copulation showed the greatest volume of secretions; after copulation, this vol- VOLUME 108, NUMBER 4 779 Table 3. Number of ovarioles per ovary, and basal oocyte length in Aphodiinae females, in different stages of sexual maturity (representing a total of 15 females for each species). Immature females do not have oocytes in their ovarioles, and hence do not appear in the table (*, mature oocytes with chorion; bo, females before oviposition; ao, females after oviposition). oo eee Length of Basal Oocytes (for 15 females/species) ¥ + ds (mm) : Mature Ovarioles per Species Ovary (number) Maturing bo ao Planolinus vittatus 5—5 ORS ==3 0107 10583) ==)0107/ 0.36 + 0.11 Gonaphodiellus opisthius 7—7 8-8 0.02 = 0.01 * 0.80 + 0.06 0.46 + 0.07 Cephalocyclus 5—5 Not observed 05/97-=0106 O22 72-0105 durangoensis Liothorax levatus 5-5 Not observed 10S == 0106 0.34 + 0.14 ume diminished by up to one-third (Table 4). In February and March, all the females and males observed were mature. Females were ready to lay eggs or had laid them. Males were ready to copulate or had recently copulated. From April to June, almost all females and males observed were maturing, with these individuals representing the first genera- tion of the year. In July and August, once again, females were found ready for oviposition or had just oviposited; males were ready to copulate or had just copulated. In September and October, individuals were again in a state of maturing; this group represented the second generation of the year. In No- vember and December, both sexes were mature (Fig. 3). This species is bivoltine, with two generations per year, the first emerging from April through June, and the second from September to October (Fig. 3). The emergence of the two new generations occurred during those periods that expe- rience the highest population levels of this species, in June and October (Fig. 1). Gonaphodielus. opisthius: Female G. opisthius have 7 to 8 ovarioles per ovary. Maturing females have small basal oo- cytes. In mature females, each ovariole has from one to three oocytes. Prior to Oviposition, each ovariole has a_ basal oocyte with chorion, measuring 0.80 + 0.06 mm. After oviposition, basal oo- cytes measure about half the length of mature oocytes (Table 3). Males have 7 testis follicles, all of the same size. Only the glandular reservoirs vary with state of sexual maturity. These reservoirs are very small in immature males, trans- parent and without secretions. At the beginning of males’ maturation process, the volume of these reservoirs is small. In mature males before copulation, the reservoirs hold much more volume, which is reduced after copulation by one-half or more (Table 4). The greatest number of both females and males is seen in June and July. Females are found just prior to or after oviposition, and similarly, males are seen just prior to or after copulation. In September, only immature females and females are found. Beginning in October, females and males are found in both immature and maturing stages (Fig. 3). Gonaphodielus opisthius 1s univoltine, emerges at the point that the rainy season begin, and reproduces to found the one new generation of the year, emerging from September to December. The species is in diapause from January to May. The emergence of mature individuals coincides with the period of highest species abundance, in July (Fic. 1): Cephalocyclus durangoensis: Female C. durangoensis have 5 ovarioles per 780 Table 4. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Number (n) and length of testis follicles per testicle, and volume of glandular reservoir, in males of various Aphodiidae species (10 males sampled per species) in different stages of sexual maturity. Immature males do not have secretions in their glandular reservoirs, and no data are shown on this group (bc, males before copulation; ac, males after copulation). Testis Follicles Length (mm) * + ds Glandular Reservoir Volume (10~3 mm3) ¥ + ds Mature Species Same Size Large and Small Maturing be ac Planolinus vittatus (2) 0.456 + 0.05 (5) 0.183 + 0.03 Tos) 22 Pell 3 58/ ==1658 NB Se 2,710 Gonaphodielus (7) 0.31 + 0.03 1.09 + 0.66 116.1 = 11.3" 65.644 13.14 opisthius Cephalocyclus durangoensis Liothorax levatus (2) 0.440 + 0.04 (2) 0.470 + 0.09 (4) 0.31 + 0.03 (©)1028 2220105 ~) eS) 1) 90 aS I+ aN eS) i) © ~ n 1) I+ aN fen Not observed Not observed PUT z= (AO) ovary. Neither immature or maturing females were found. Mature females had 3 or 4 oocytes per ovariole. Before oviposition, the basal oocytes, and the second, and often the third, oocyte were mature, with chorion. Basal oocytes measured a mean 0.79 + 0.06 mm. Following oviposition, basal oocytes were observed to be small (Table 3). Males have 6 testis follicles in each testicle, with 2 of the follicles larger than the other 4. No immature or maturing males were found. All the males collected were mature, with maxium glandular reservoir volume seen immediately be- fore copulation; after copulation, volume was reduced, by up to more than half from this value (Table 4). The species is univoltine, with the population emerging in the first months of the rainy season, and reproductive activity declining just a little more than a month after the beginning of this active period. The new generation was not seen at the surface, possibly passing through diapause, from January to May and September to December (Fig. 3). The only high population levels seen in this species corresponded to the emergence of mature females and males principally in July, in June only 6 specimens and in August another 2, where find (Fig. 2). Liothorax levatus: Female L. levatus have 5 ovarioles per ovary. Neither immature or maturing females were observed. All the females collected were mature, showing one or two oocytes per ovariole. Mature females before oviposi- tion showed the largest oocytes of all species observed; after oviposition, the following basal oocytes measured far less (Table 3). Males have 7 testis follicles. All males observed were mature. Prior to copulation, males had the most volumi- nous glandular reservoirs; after copula- tion, this volume declined by more than half (Table 4). This species is also univoltine, and only active during three months, after which it appears to enter into diapause for nine months of the year (Fig. 1). All individuals observed were mature fe- males or males (Fig. 3). The reproductive cycles of O. chevro- lati retusus and O. viridiobscurus were not studied. DISCUSSION Trophic preferences.—Several species appeared to prefer either cow or horse dung, while other species showed no preference. Nevertheless, a_ statistically significant preference (for cow dung) was seen only in P. vittatus. Another study based on a larger number of individuals would be needed to obtain sufficient data to analyze whether other species’ appar- ent preferences are real. Recent studies VOLUME 108, NUMBER 4 FEMALES (%) MALES (%) FEMALES (%) MALES (%) 100 50 100 100 100 50 Fig. Planolinus vittatus Ee ae vaeAC Mies JacA eSeOeIN iD J Cephalocyclus durangoensis F WM A Ml al da S ©) IN MONTHS inmature 781 Gonaphodielus opisthius 100 50 100 50 le Ml Ne IM dh al ANS) ©) INI D Liothorax levatus 100 a 0 100 50 EM ASM Ase On Nia MONTHS ae maturing Pal mature Monthly percentages of sexually immature, maturing, and mature females and males, for Aphodiinae species found in El Llano de la Flores, Oaxaca, January—December 2003 (with 10 females and 10 males of each species examined each month). 782 on other dung beetle species indicate that there are marked species preferences for selected foods (Martin-Piera and Lobo 1996, Galante and Cartagena 1999, Finn and Giller 2002, Laurent et al. 2004). Phenology.—The timing of active pe- riods differed from species to species. Most species examined— P.vittatus, G. opisthius, C. durangoensis, O. chevrolati retusus, and O. viridiobscurus—were ac- tive during the rainy period in the hot, humid summer. The exceptions were L. levatus, which was active only at the beginning of the dry period, correspond- ing to winter, and P. vittatus, which was also active only during the dry period. The phenology at G. opisthius at El Llano de las Flores, Oaxaca, is quite similar to that observed in Cutyachapa, Veracruz, in the same species (Cruz et al. 2002). Similarly, the phenology seen in C. durangoensis is very like that observed in Cephalocyclus hogei (Bates 1887), another species of the same genus that has been studied in Cuiyachapa, Vera- cruz (Cruz et al. 2002) and San José de Aguazuelas, Veracruz (Martinez 2005). The phenology of P. vittatus resembles that of the same species in Las Vigas de Ramirez, Veracruz (Martinez, M. L., unpublished observations). In contrast, the phenology of L. /evatus in El Llano de las Flores, Oaxaca, is not like that seen in the same species in San José de Aguazuelas, Veracruz (Martinez 2005). In the Oaxaca location, this species was active in January, November, and De- cember; in the Veracruz location, the species was active from August to December. We do not know what factor(s) cause this difference. Reproductive cycles.—Three of the four Aphodiinae species examined, G-. opisthius, C. durangoensis, and L. levatus, univoltine, like the majority of Aphodiinae species that have been stud- ied. (Martinez, 20014 @ruz vet vale 12002, Martinez 2005). Planolinus vittatus was the only bivoltine species, as are other are PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON European and Asiatic species (Schmidt 1935, Landin 1961, Yoshida and Kata- kura 1985, Verdu and Galante 1995, Gittings and Guiller 1997, Vitner 1998). Onthophagus chevrolati retusus appears to be bivoltine, although this needs to be confirmed with further study of its reproductive cycle. Onthotrupes viridiob- scurus 1s univoltine. Gonaphodielus opisthius, L. levatus, and C. durangoensis females and males emerge mature to copulate and lay eggs. The reproductive period is short for these species, and the diapause long, from five months in the first species to nine months in the other two species. Planolinus vittatus 1s in diapause for just one month. We do not know, for any of these species, in what preimaginal or imaginal stage of development they exist during this period. Diapause has been studied in only a few Aphodiinae species (see Martinez 2001). Onthophagus chey- rolati retusus has a period of activity lasting ten months and a diapause of two months. In Onthotrupes viridiobscurus, seven months of activity are followed by a diapause of five months. Most of the species studied are active during the hottest, most humid period of the year, summer and fall. They enter diapause in the dry, cold period, during winter and spring. The only exceptions are P. vittatus and L. levatus, which are active at the beginning of the dry period. It appears that all these species’ activity periods are regulated principally by soil moisture, which is determined by rain and the warm temperature seen during summer and part of fall; this activity pattern is seen frequently in dung beetle species (Martinez 1992, Martinez and Vazquez 1995, Martinez and Cruz 2002, Cruz et al. 2002, Martinez 2005) and other insects (Engelmann 1970). For dung beetles, photoperiod is probably not a significant determinant, given that these species live the greater part of their lives buried; and seasonal variation in VOLUME 108, NUMBER 4 photoperiod is not as marked in tropical as in temperate zones. This kind of basic research can be highly useful for understanding insect diversity in pasture, how species’ phe- nology or reproductive cycles might be affected by agronomic or veterinary residuals, and the behavior of introduced species. All this information is essential in establishing measures to conserve and manage these ecosystems. ACKNOWLEDGMENTS We are grateful to Marco Dellacasa (University of Pisa, Italy) and Leonardo Delgado for species identification, to Francisco Cabrero for his help with statistical analysis, and to Sergio Ibanez (Instituto de Ecologia A. C. Xalapa, Mexico) for his suggestions on _ the manuscript. To R. Gordon and another anonymous reviewer for their valuable comments and to Ann Covalt for English translation of the manuscript. This re- search was conducted thanks to the support of the Departamento de Biodi- versidad y Ecologia Animal del Instituto de Ecologia A. C. LITERATURE CITED Bryan, R. P. 1973. The effects of dung beetle activity on the number of parasitic gastroin- testinal helminths larvae recovered from pas- ture samples. Australian Journal of Agricul- tural Research 24: 161-168. Chirico, J., S. Wiktelius, and P. J. Waller. 2003. Dung beetle activity and trichostrongylid eggs into infective larve in cattle faeces. Veterinary Parasitology 118(1—2): 157-163. Cruz, R. M., I. Martinez, M., and M. Alvarado. 2002. Population and reproductive features of Aphodius (Trichaphodius) opisthius Bates and Cephalocyclus hogei Bates (Coleoptera, Apho- diidae: Aphodiinae). The Coleopterists Bulle- tin 56(2): 221—235. Dellacasa, M., R. D. Gordon, and G. Dellacasa. 2002. Aphodiinae described or recorded by Bates in Biologia Centrali Americana (Co- leoptera Scarabaeoidea: Aphodiidae). Acta Zoologica Mexicana (n. s.) 86: 155-223 development of 783 Engelmann, F. 1970. The Physiology of Insect Reproduction. Pergamon Press, Oxford. 307 pp. Fincher, G. T. 1975. Effect on dung beetle activity on the number of nematode parasitoid re- quired by grazing cattle. Journal of Parasitol- ogy 61: 759-762. . 1981. The potential value of dung beetles in pasture ecosystems. Journal of Georgia Entomological Society 16: 316-333. Finn, J. and P. S. Giller. 2002. Experimental investigations of colonization by north tem- perate dung beetles of different types of domestic herbivore dung. Applied Soil Ecolo- gy 20: 1-13. Galante, E. and C. Cartagena. 1999. Comparison of Mediterranean dung bettles (Coleoptera: Scarabaeoidea) in cattle and rabbit dung. Environmental Entomology 28: 420-424. Gittings, T. and P. S. Giller. 1997. Life history traits and resource utilisation in an assemblage of north temperate Aphodius dung beetles (Coleoptera: Scarabaeidae). Ecography 20: 55—66. Halffter, G. and W. D. Edmonds. 1982. The nesting behavior of dung beetles (Scarabaei- nae). An ecological and evolutive approach. Publ. 10, Instituto de Ecologia, A.C. México. 176 pp. Hanski, I. and Y. Cambefort. 1991. Dung Beetles Ecology. Princeton Univ. Press. Princenton, New Jersey. 481 pp. Landin, B. O. 1961. Ecological studies on dung- beetles (Col. Scarabaeidae). Opuscula Ento- mologica supplementum 19: 209-221. Laurent, D., G. Epinat, and J. P. Lumaret. 2004. Trophic preferences mediated by olfactory cues in dung beetles colonizating cattle and horse dung. Environmental Entomology 33(2): 370-377. umarcty Jib: Martinez, M. 2005. El impacto de los productos veterinarios sobre insectos coprofagos: Consecuencias sobre la degradacion del estiércol en pastizales. Acta Zoologica Mexicana 21(3): 137-148. Martin-Piera, F. and J. M. Lobo. 1996. A comparative discussion of the trophique pref- erences in dung beetles communities. Miscella- nia Zoologica 19: 13-31. Martinez, M. I. 1992. Données comparatives sur Vactivité reproductrice chez Canthon indiga- ceus chevrolati Harold et Canthon cyanellus cyanellus LeConte (Coleoptera, Scarabaeidae). Annales de la Société entomologique de France (n.s.) 28(4): 397-408. ———. 2001. La biologia reproductiva en Apho- diidae (Coleoptera Scarabaeoidea): Sintesis de datos conocidos. Bolletino della Societa En- tomologica Italiana 133(2): 113-130. and) le 784 . 2002. Técnicas basicas de anatomia micro- scopica y de morfometria para estudiar los insectos. Boletin de la de la Sociedad Entomo- logica Aragonesa 30: 187-195. . 2003. Spermatophore in Aphodiinae (Co- leoptera Scarabaeidae). Proceedings of the Entomological Society of Washington 105(4): 982-989. . 2005. Abundancias poblacionales y ciclos reproductivos de tres especies de escarabajos estercoleros (Coleoptera: Aphodiinae: Apho- diini) del Volcan Cofre de Perote, Veracruz, México. Folia Entomologica Mexicana 44(1): 27-36. Martinez, M. I. and M. Alvarado. 2001. Compor- tamiento de nidificacion en Aphodius ( Tricha- pPhodius) opisthius Bates. Elytron 15: 73-78. Martinez, M. I. and M. Cruz R. 1999. Comparative morphological analysis of testis follicles in dung beetles (Coleoptera Scarabaeidae: Scar- abaeinae, Aphodiinae, Geotrupinae). Proceed- ings of the Entomological Society of Washing- ton 101(4): 804-815. . 2002. Fenologia y ciclos reproductivos en Ataenius apicalis Hinton y Ataenius sculptor Harold (Coleoptera Aphodiidae). Bulletin de la Société entomologique de France 107(2): 177-186. Martinez, M. I. and J. P. Lumaret. 2006. Las practicas agropecuarias y sus consecuencias en la entomofauna y el entorno ambiental. Folia Entomologica Mexicana 45(1): 57-68. Martinez, M. I. and A. Vazquez. 1995. Influencia de algunos factores ambientales sobre la reproducci6n en Canthon cyanellus cyanellus LeConte (Coleoptera Scarabaeidae: Scarabaei- nae). Elytron 9: 5-13. Martinez, M. I., M. Cruz, R., and J. P. Lumaret. 2000. Efecto del diferente manejo de los pastizales y del ganado sobre los escarabajos coprofagos Ataenius apicalis Hinton y Ataenius sculptor Harold (Scrabaeidae: Aphodiinae: Eupariini). Acta Zoologica Mexicana 80: 185-196. Martinez, M. I., C. Deloya, and M. Dellacasa. 200la. Anatomical and functional data on female and male reproductive systems of some dung beetles species of Aphodiinae and Eu- pariinae inabiting México (Coleoptera: Apho- diidae). Proceedings of the Entomological Society of Washington 103(1): 227-248. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Martinez, M. I., J. P. Lumaret, and M. Cruz R. 2001b. Suspected side effects of a herbicide on dung beetle populations (Coleoptera: Scara- baeidae). Comptes Rendus de Académie des Sciences. Paris. Sciences de la Vie 324: 989— 994. Moon, R. D., E. C. Loomis, and J. R. Anderson. 1980. Influence of two species of dung beetles on larvae of face fly. Environmental Entomol- ogy 9: 607-612. Ridsdill-Smith, T. J. and L. Hayles. 1990. Stages of bush fly, Musca vetustissima (Diptera: Musci- dae), killed by scarabeinae dung beetles (Co- leoptera: Scarabaeidae) in favourable cattle dung. Bulletin of Entomological Research 80: 473-478. Ridsdill-Smith, T. J., L. Hayles, and M. J. Palmer. 1987. Mortality of eggs and larvae of the bush fly Musca vetustissima Walker (Diptera: Mus- cidae) caused by scarabeinae dung _ beetles (Coleoptera: Scarabaeidae) in favourable cattle dung. Bulletin of Entomological Research 78: 633-639. Rougon, D., C. Rougon, J. Trichet, and J. Levieux. 1988. Enrichissement en matiere organique d’un sol sahélien aun Niger par les insectes coprophages (Coleoptera, Scarabaeidae) im- plications agronomiques. Revue d’Ecologie et de Biologie du Sol 25: 413-434. Schmidt, G. 1935. Beitrage zur Biologie des Aphodiinae. Stettiner Entomologische Zeitung 96: 343-350. Verdu, J. R. and E. Galante. 1995. Life history and description of the larval stage of Aphodius lusitanicus Erichson, 1848 (Coleoptera: Scar- abaeoidea: Aphodiinae). Acta Zoologica cra- coviensia 38(2): 205—212. Vitner, J. 1998. Reproductive biology of the Central European Aphodius species with large scutellum (Coleoptera: Scarabaeidae: Aphodii- nae). Acta Societatis Zoologicae Bohemicae 62: 277-253. Yokoyama, K., H. Kai, T. Koga, and T. Aibe. 1991. Nitrogen mineralization and microbial- populations in cow dung, dung balls snd underyling soil affected by paracoprid dung beetles. Soil Biology & Biochemistry 23(7): 643-647. Yoshida, N. and H. Katakura. 1985. Life cycles of Aphodius dung beetles (Coleoptera: Scarabaei- dae) in Sapporo Northern Japan. Environ- mental Science, Hokkaido 8(22): 209-229. PROC. ENTOMOL. SOC. WASH. 108(4), 2006, pp. 785-792 EARLY CRETACEOUS PHLEBOTOMINE SAND FLY LARVAE (DIPTERA: PSYCHODIDAE) GEORGE POINAR, JR., R. L. JACOBSON, AND CAROL L. EISENBERGER (GP) Department of Zoology, Oregon State University, Corvallis, OR 97331, U.S.A. (e-mail: poinarg@science.oregonstate.edu); (RLJ) Department of Parasitol- ogy, The Hebrew University—Hadassah Medical School, POB 12272, Jerusalem, 91120, Israel (e-mail: jacobsr@cc.huji.ac.il); (CLE) P.O. Box 173, Tzur Hadassah, 99875, Israel (e-mail: carol@biowabbit.com) Abstract.—Two sand fly larvae (Diptera: Psychodidae) are characterized from Early Cretaceous Burmese amber. Both larvae, which are considered to be post-first instars, possess only a single pair of caudal setae on the terminal (9th) abdominal segment. All known extant post-first instar sand fly larvae possess two pairs of caudal setae except for the Old World Phlebotomus (Larroussius) tobbi Adler and Theodor and New World Brumptomyia Franga and Parrot. The fossil larvae could represent an ancestral line continued today by P. tobbi or a completely separate lineage, and they show that a single pair of caudal setae was an ancient characteristic. A close association of the fossil larvae with the fruiting bodies of a non-gilled coral fungus, Palaeoclavaria burmitis Poinar and Brown (Hymenomycetes: Palaeoclavar- laceae), suggests a source of nourishment for Early Cretaceous sand flies. Key Words: sand fly larvae, Early Cretaceous, caudal setae, food source, fossil fungus, Burmese amber During a survey of hematophagous insects in amber, a piece of Early Cretaceous amber from Myanmar (Burma) was discovered to contain a pair of sand fly larvae together with their possible fungal food source (Figs. 1, 2). These larvae are characterized, and their morphological features compared with those of extant sand fly larvae. This is the first fossil record of sand fly larvae. MATERIALS AND METHODS The amber piece containing the fossil sand fly larvae was received in the summer of 2004 from the Leeward Capital Corp., a Calgary-based company working with a Burmese mining compa- ny in Noitje Bum. The amber with the fossils is semicircular in outline, measur- ing 13 mm along the longest side, 9 mm wide, 4mm deep and weighs 0.4 gm. Observations, drawings and_ photo- graphs were made with a Nikon SMZ- 10 R stereoscopic microscope and Nikon Optiphot compound microscope with magnifications up to 600. The amber piece containing the fossil flies is de- posited in the Poinar amber collection (accession #B-D-24) maintained at Ore- gon State University. Amber from Burma occurs 1n lignitic seams in sandstone-limestone deposits in the Hukawng Valley, southwest of Main- gkhwan in the state of Kachin (26°20’N, 96°36’E). Palynomorphs obtained from the amber beds have been assigned to the 786 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 108, NUMBER 4 Upper Albian (~ 100-105 mya) of the Early Cretaceous (Cruickshank and Ko 2003), however since the amber is sec- ondarily deposited, the age could be older. Nuclear magnetic resonance (NMR) spectra of amber samples taken from the same locality as the fossils indicated an araucarian (Araucariaceae) (possibly Agathis) as a possible source of the amber (J. B. Lambert and Y. Wu, unpublished data, 2002). RESULTS Both sand fly larvae are similar in size and have the basic characters typical of phlebotomine larvae (Quate and Vock- esoth, 198i, Perfiltey 1966), such as a hypognathous head, 3 thoracic and 9 abdominal segments, oval third antennal segments and long, paired caudal setae on the last abdominal segment (Figs. 1— 6). The first two thoracic segments bear small pseudopods (tubercles) on their ventral surfaces, the last thoracic and first 8 abdominal segments bear promi- nent pseudopods and the last abdominal segment has a small pseudopod. The larva nearest the edge of the amber (Neowih)s (hig. 1) ise 2377.mim,..in length and has an approximate head capsule length of 214 um. The larva adjacent to a fungal fruiting body (No. 2) (Fig. 2) is 2.83 mm in length and has an approxi- mate head capsule length of 220 um. Only approximate head capsule lengths could be obtained since both larvae have their heads slightly turned making defi- nite measurements impossible. There is no evidence that the larvae are shrunken or distorted. Si Both larvae are uniformly cream colored and have light head capsules. However, the ventral pseudopods on the thoracic and abdominal segments appear as darker areas. The antennae are located near the front border of the head (Fig. 5). Only the oval-oblong third segment of the antenna is visible, and from this segment projects a minute, blunt- tipped apical spine. Depression lines extend through these terminal seg- ments. The whorled spinules on the setae are quite reduced and many appear to be absent with only the ringlike bases remaining. The mouthparts are ob- scured. Both larvae possess only one pair of caudal setae arising from the dorsum of the terminal (9th) abdominal segment (Figs. 1-6). These setae vary from 0.725 to 0.910 mm in length (the tip of the left caudal seta on larva | was polished away). A pair of shorter spinulose setae (0.16 to 0.21 mm in length) is located on the last abdominal segment. This pair extends outward and slightly downward from a subdorsal position beneath and outside the caudal setae. There are no other noticeable setae on the last ab- dominal segment. The amber piece also contains several fruiting bodies of the coral mushroom, Palaeoclavaria burmitis Poinar and Brown, a member of the non-gilled Aphyllophorales (Hymenomycetes). The posterior portion of larva No.2 is attached to a fungal fruiting body by a mucilaginous-appearing substance. Another fruiting body has what appears to be an isolated piece of fungal tissue adjacent to a hole in its side. Both larvae Figs. 1-4. Early Cretaceous sand fly larvae. 1, Dorsal-lateral view of larva No. 1. Bar = 503 um. 2, Lateral view of larva No. 2. Adjacent to the larva is its likely food source, a fruiting body of the coral fungus, Palaeoclavaria burmitis (arrow). Bar = 608 ttm. 3, Dorsal view of last (9th) abdominal segment showing paired caudal setae (arrows). Bar = 200 tm. 4, Dorsal view of last (9th) abdominal segment showing paired spinulose setae (arrows). Bar = 200 um. 788 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5—6. Early Cretaceous sand fly larva (camera-lucida line drawings). 5, Lateral posterior view of head of larva No. 1 showing antenna and arrangement and types of cephalic setae. Bar = 50 um. 6, Dorsal view of last (9'») abdominal segment showing paired inner caudal setae and paired outer spinulose setae. Bar = 8 um. VOLUME 108, NUMBER 4 have small particles of what could be fungal debris attached to their bodies. Voided fecal pellets occur at the terminus of both larvae. A mite (Acari) is the only other arthropod inclusion in the amber. Mites would be expected in such a habitat and some are ectoparasites of sand flies (Warburg et al. 1991). DISCUSSION The generic identity of the fossil sand fly larvae is unknown, but on the basis of size and morphology, both probably belong to the same stage and same species. Thus far, Palaeomyia burmitis Poinar (2004) is the only sand fly de- scribed from Burmese amber, and while the larvae could belong to this species, they could also be from undescribed species from that amber source. It is difficult to determine the instar of the fossil larvae since Early Cretaceous sand flies are smaller than extant Old World species (the Burmese amber mines are located on the Burma Plate (Mitchell 1993, Cruickshank and Ko 2003) which in the Early Cretaceous, was part of Laurasia, thus the fossil larvae belong to Old World clades). Adult lengths of some Early Creta- ceous sand flies and closely related psychodid flies are 0.887 mm_ and 1.04 mm for the Burmese amber Palaeo- myia burmitis Poinar (2004) and Eophle- botomus connectens (Cockerell) (Duck- house 2000), respectively, and 1.06 mm, 1.07 mm, 0.93 mm, and 1.05 mm for the Lebanese amber Mesophlebotomites hen- nigi Azar et al. (1999), Libanophleboto- mus lutfallahi Azar et al. (1999), Phlebo- tomites brevifilis Hennig (1972), and Eophlebotomus gezei Azar et al. (2003), respectively. The two specimens of Eophile- botomus carentonensis Azar et al. from Early Cretaceous French amber have lengths of 0.93 and 0.97 mm (Azar et al. 2003). The Cretaceous adult lengths (0.89— 1.07 mm) are smaller than those of 789 extant Old World adults, which range from 1.20mm to 3.70 mm (Perfil’ev 1966) (Table 1). The Cretaceous fossils are also smaller than the New World Dominican amber phlebotomine fossils Trichopygomyia killickorum Filho et al. (2004) (adult length = 1.58 mm) and Pintoyia falcaorum Brazil and Filho (2002) (adult length = 1.46 mm). As- suming that the size of the adults is correlated with those of the larvae, then the lengths of the larval stages of the fossil species likely would be smaller than the corresponding sizes of extant larvae. A comparison of the lengths of the fossil larvae with those of extant larvae of Phlebotomus (Phlebotomus) —papatasi Scopoh, P. (Adlerius) arabicus Theodor, Sergentomyia (Sergentomyia) bedfordi Newstead, S. (Sergentomyia) hamoni Abonnenc, S. (Rondaniomyia) ingrami Newstead, and S. (Rondaniomyia) dur- eni Parrot (Table 1) shows that they are definitely post-first instars and possibly even fourth instars since their lengths overlap with those of extant fourth stage larvae. Extant first stage sand fly larvae have rounded heads with an almost spherical third antennal segment (Perfi- Vev 1966). The fossil larvae have elon- gated heads and the third antennal segment is oblong-oval (Fig. 5), which is typical of 24 to 4th larval instars. Both the body and head of the fossil larvae are light colored, which is similar to the yellow color of some fourth stage Lutzo- myia Franga larvae (Ward 1976). The third instars of Lutzomyia (Psychodopy- gus) panamensis (Shannon) also have light-colored head capsules similar to those of the fossil larvae (Lawyer and Perkins 2000). Voiding the gut contents just before pupation also gives the body a whitish appearance (Lawyer and Per- kins 2000). An interesting feature of the fossil larvae is the presence of only one pair of caudal setae on the terminal abdominal segment. The only known extant sand fly 790 Table 1. sand flies. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Lengths (mm) of larval instars and adults of some extant Phlebotomus and Sergentomyia Species Ist Stage 2nd Stage 3rd Stage 4th Stage Adult Reference P. arabicus 0.6—1.2 1.42.5 2.3-3.4 3.3-4.6 2.5—3.0 Volf1.2 P. papatasi 0.5—1.0 1.3-1.4 - 2.5—3.2 Perfil’ev, 1966 P. papatasi 0.4—0.8 2.3-2.8 3.2-3.5 3.64.2 2.4-3.3 R.L. Jacobson2 S. bedfordi 0.3-0.7 0.8—1.0 — 2.1—2.8 1.82.4 Trouillet, 1977! S. ingrami 0.9-1.0 — -= 2.2-2.6 1.6—2.4 Trouillet, 1979! S. dureni 0.6—1.0 == — 2.2-2.6 1.6-2.4 Trouillet, 1979! S. hamoni 0.5—0.6 — _- 2.32.6 2.32.6 Trouillet, 1979! 1 Adult measurements taken from Abonnenc (1972). 2 Unpublished observations. larvae which possess a single pair of caudal setae in the post-first instars is Phlebotomus tobbi from the Tonian Greek islands and Syria (Killick-Ken- drick et al. 1989, 1992). However, P. tobbi has four downwardly directed spinulose setae on the ventral surface of the ninth segment, which differs from the single pair of subdorsal spinulose setae 1n the fossil larvae. Mature larvae of New World Brumptomyia Franga and Parrot species have two caudal setae (Theodor 1965, Young and Duncan 1994) while Lutzomyia larvae have two pairs of caudal setae on post-first instars (Ward 1976, Quate and Vockeroth 1981, Law- yer and Perkins 2000). Some mormiuine moth fly larvae (Psychodinae) possess a pair of caudal filaments but they are shorter and differ from sand fly setae (Jezek 1989). Reduction is generally the rule in the evolutionary process. Therefore, Early Cretaceous Old World post-first instar sand fly larvae would be expected to possess at least two, if not more pairs, of caudal setae. The fossils could represent an ancient Old World lineage continued today by P. tobii (Killick-Kendrick et al. 1989, 1992), and though this species is absent from Myanmar today, the sub- genus, Larroussius occurs in the region (Lewis 1978). However, the fossil could represent a completely separate extinct lineage. Now The fruiting bodies of the coral fungi associated with the fossil larvae have been described as Palaeoclavaria burmi- tis (Hymenomycetes: Palaeoclavariaceae) (Poinar and Brown 2003). A perforation in one of the fruiting bodies adjacent to some extruded fungal debris and the close association of larva No. 2 to a second fruiting body suggests an intimate re- lationship between the larvae and the fungus. It is possible that the sand fly eggs were deposited on or close to the fruiting body, which then served as both a source of humidity and food for the eclosed larvae. This fungus probably developed around wounds or in rotting portions of the trunk or branches of the resin-pro- ducing tree, which is one of the de- velopmental habitats of extant sand flies (Hanson 1961, Rutledge and Mosses 1972). It is possible that these, as well as the closed sporocarps of other fungi, were a food source for Early Cretaceous sand flies. Today some fungi, especially those with closed fruiting bodies, are important developmental habitats for primitive flies such as fungus gnats (Mycetophilidae and Sciaridae), gall midges (Cecidomyliidae), crane flies (Tipulidae), scavenger flies (Scatopsidae), and wood gnats (Anisopo- didae) (Bruns 1984). Fungi as food for extant sand flies is mentioned by Perfil’ev (1966) and Wermelinger and Zanuncio (2001). In a study of the larval feeding habitats of Panamanian sand flies, the VOLUME 108, NUMBER 4 preferred sites were sheltered areas be- tween tree buttresses, which were also the resting habitat for adult sand flies (Han- son 1961). The substrate in these areas was composed of organic matter from plant and animal remains. It is possible that the sand fly larvae feed on fungi developing in and on this_ organic matter. The small size of the Early Cretaceous sand flies may indicate an early period in the establishment of adult and larval feeding niches. If fungal fruiting bodies were an important food source for Early Cretaceous sand flies, then their occur- rence would have been seasonally vari- able and unpredictable, which would favor an increased feeding rate and reduced developmental times. The cau- dal setae may have been partially ex- posed on the outside of the fruiting body, possibly serving as detectors of ap- proaching danger since recent studies suggest that they have a sensory function (Pessoa et al. 2001). Just when phlebotomine sand flies evolved is still unresolved. The oldest known fossils occur in 120 million year old Lebanese amber (Azar et al. 1999); however, it is not known whether these were biting forms since descriptions are based only on males. The oldest evidence of a haematophagous habit in sand flies is in the Burmese amber Pa/aeomyia burmitis that contains within its body not only leishmanial parasites but also infected vertebrate blood cells (Poinar and Poinar 2004a, b). ACKNOWLEDGMENTS We thank Professor Y. Schlein for confirmation of the identity of the sand fly larvae, Dr. P. Volf for providing P. arabicus larval data, J. B. Lambert and Y. Wu, Department of Chemistry, Northwestern University, for their anal- ysis of the amber, and Roberta Poinar for comments on an earlier version of this manuscript. 791 LITERATURE CITED Abonnenc, E. 1972. Les Phlébotomes de la Région Ethopienne (Diptera, Psychodidae). Cahiers O. R.S. T. O. M. Serie Entomologie Médicale et Parasitologie 55: 1-289. Andrade Filho; J. D=.A. L. Faleao, and R. P: Brazil. 2004. A new phlebotomine fossil species Trichopygomyia_ killickorum sp. n. (Diptera: Psychodidae) found in the Dominican Re- public amber. Parasite 11: 71-73. Azar, DS Da IN; Pemichot; “and “AS Nelly 20088 Psychodids from the Cretaceous Ambers of Lebanon and France, with a Discussion of Eophlebotomus Cockerell, 1920 (Diptera, Psychodidae). Annals of the Ento- mological Society of America 96: 117-126. Azar, D., A. Nel, M. Solignac, J.-C. Paicheler, and F. Bouchet. 1999. New genera and species of psychodoid flies from the Lower Cretaceous amber of Lebanon. Palaeontology 42: 1101— 1136. Brazil, R. P. and J. D. Andrade Filho. 2002. Description of Pintomyia (Pifanomyia) fal- caorum sp. n. (Diptera: Psychodidae: Phlebo- tominae), a fossil sand fly from Dominican amber. Memorias do Instituto Oswaldo Cruz, Rio de Janeiro 97: 501—503. Bruns, T. D. 1984. Insect Mycophagy in the Boletales: Fungivore diversity and the mush- room habitat, pp. 91-129. In Wheeler, Q. and M. Blackwell, eds. Fungus—insect relation- ships. Columbia University Press, New York. Cruickshank, R. D. and K. Ko. 2003. Geology of an amber locality in the Hukawng Valley, northern Myanmar. Journal of Asian Earth Sciences 21: 441-455. Duckhouse, D. A. 2000. Redescription and re- evaluation of the Burmese amber psychodid Eophlebotomus connectens Cockerell and _ its phylogenetic position (Diptera: Psychodidae). Systematic Entomology 25: 503—S09. Hanson, W. J. 1961. The breeding places of Phlebotomus in Panama (Diptera, Psychodi- dae). Annals of the Entomological Society of America 54: 317-322. Hennig, W. 1972. Insektenfossilien aus der unter Kreide. IV. Psychodidae (Phlebotominae), mit inner kritischen Ubersicht iiber das phylogen- etische System der familie und die bisher beschriebenen Fossilien (Diptera). Stuttgarter Beitrage zur Naturkunde 241: 1—69. Jezek, J. 1987. Description of new Mormiuine moth- flies (Diptera, Psychodidae) from Abkhazia. Acta Entomologica Bohemoslovaca 84: 261—272. Killick-Kendrick, R., M. Killick-Kendrick, N. Léger, B. Pesson, G. Madulo-Leblond, and A. M. Page. 1989. Absence of outer caudal connectens setae on all larval instars of Phlebotomus tobbi from the Ionian Greek islands. Medical and Veterinary Entomology 3: 131-135. Killick-Kendrick, R., M. K. Sirdar, and M. Killick- Kendrick. 1992. Lack of outer caudal setae as a constant feature of Phlebotomus tobbi larvae. Medical and Veterinary Entomology 6: 171-172. Lawyer, P. G. and P. V. Perkins. 2000. Leishmaniasis and Trypanosomiasis, pp. 231—298. Jn Eldridge, B. F. and J, D. Edman, eds. Medical Entomol- ogy. Kluwer Academic Publishers, Dordrecht. Lewis, D. J. 1978. The phlebotomine sandflies (Diptera:Psychodidae) of the Oriental Region. Bulletin of the British Museum (Natural History), Entomology series 37: 217-337. Mitchell, A. H. G. 1993. Cretaceous-Cenozoic tectonic events in the western Myanmar (Burma)-Assam region. Journal of the Geo- logical Society 150: 1089-1102. Perfil’ev, P. P. 1966. Phlebotomidae (Sandflies). Vol. 3, No. 2, Fauna of U.S.S.R. (Diptera). Academy of Sciences, Moscow. 363 pp. (Trans- lated from Russian by the Israel program for scientific translations, Jerusalem, 1968). Pessoa, F. A. C., R. G. Queiroz, and R. D. Ward. 2001. External morphology of sensory struc- tures of fourth instar larvae of Neotropical species of phlebotomine sand flies (Diptera: Psychodidae) under scanning electron micros- copy. Memorias do Instituto Oswaldo Cruz, Rio de Janeiro 96: 1103-1106. Poinar, Jr., G. O. 2004. Palaeomyia burmitis gen. n., sp. n. (Phlebotomidae: Diptera), a new genus of Cretaceous sand flies with evidence of blood sucking habits. Proceedings of the Entomolog- ical Society of Washington 106: 598-605. Poinar, Jr., G. O. and A. E. Brown. 2003. A non- gilled hymenomycete in Cretaceous amber. Mycological Research 107: 763-768. Poinar, Jr., G. O. and R. Poinar. 2004a. Paleo- leishmania proterus n. gen., n. sp., (Trypano- somatidae: Kinetoplastida) from Cretaceous Burmese amber. Protista 155: 305-310. . 2004b. Evidence of vector-borne disease of Early Cretaceous reptiles. Vector-Borne and Zoonotic Diseases 4: 281—284. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Quate, L. W. and J. R. Vockeroth. 1981. Psycho- didae, pp. 153-190. In McAlpine, J. F., ed. Manual of Nearctic Diptera, Monograph 27, Vol. 1. Agriculture Canada, Research Branch, Ottawa. Rutledge, L. C. and H. L. Mosses. 1972. Biology of immature sandflies at the bases of trees in Panama. Environmental Entomology 1: 300-309. Theodor, O. 1965. On the classification of Amer- ican Phlebotominae. Journal of Medical Ento- mology 2: 171-197. Trouillet, J. 1977. Sergentomyia (Sergentomyia) bedfordi firmatus Parrot et Malbrant, 1945 et Sergentomyia (Sergentomyia) bedfordi medius Kirk et Lewis, 1950 (Diptera, Phlebotomidae). Etude morphologique des stades pré-imagi- naux et notes bio-écologiques. Annales de Parasitologie Humaine et Comparée 52: 35—52. ——. 1979. Sergentomyia (Rondanomyia) in- grami Newstead, 1914, Sergentomyia ( Ronda- nomyia) dureni Parrot, 1934 et Sergentomyia hamoni Abonnenc, 1958 (Diptera Phlebotomi- dae). Etude morphologique des stades pré- imaginaux et notes bio-écologiques. Annales de Parasitologie Humaine et Comparée 54: 353-73. Warburg, A., K. Ostrovska, and P. G. Lawyer. 1991. Pathogens of phlebotomine sandflies: A review. Parassitologia 33: 519-526. Ward, R. D. 1976. The immature stages of some phlebotomine sandflies from Brazil (Diptera: Psychodidae). Systematic Entomology 1: 227-240. Wermelinger, E. D. and J. C. Zanuncio. 2001. Development of Lutzomyia intermedia and Lutzomyia longipalpis (Diptera: Psychodidae: Phlebotominae) larvae in different diets. Bra- ailian Journal of Biology 61: 405-408. Young, D. G. and M. A. Duncan. 1994. Guide to the Identification and Geographic Distribution of Lutzomyia Sand flies in Mexico, the West Indies, Central and South America (Diptera: Psychodidae). Memoirs of the American En- tomological Institute, Number 54, 881 pp. PROC. ENTOMOL. SOC. WASH. 108(4), 2006, pp. 793-807 REVISION OF THE NORTH AMERICAN APHID GENUS BIPERSONA HOTTES (HEMIPTERA: APHIDIDAE) GARY L. MILLER AND ANDREW S. JENSEN (GLM) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, Bldg. 005, BARC-W, Beltsville, MD 20705, U.S.A. (e-mail: gmiller@sel.barc.usda.gov); (ASJ) 9344 Road S NE, Moses Lake, WA 98837, U.S.A. and Adjunct Faculty, Department of Entomology, Washington State University, Pullman, WA 99164, U.S.A. (e-mail: ajensen@televar.com) Abstract.—Adult female apterae and alates of the genus Bipersona Hottes are described and illustrated. Bipersona torticauda (Gillette), previously a junior synonym of Bipersona ochrocentri (Cockerell), is now considered a valid species. Key Words: aphid, thistle, Bipersona torticauda, Bipersona ochrocentri The aphid genus Bipersona Hottes (1926) was proposed for a group of thistle-inhabiting aphids with unusual morphology of the cauda and anal plate. The cauda of these species is elongate, narrow, and appears twisted or con- stricted along its length, and the anal plate has a posteriorly directed projec- tion. Bipersona lives on various species of thistles in the genus Cirsium (Asteraceae) and the aphids invariably are attended by ants. The recent synonymy (Eastop and Blackman 2005) of Bipersona hottesi Knowlton and Smith (= Eomacrosiphum nigromaculosum MacDougall) resulted in a monotypic genus. Samples sent to one of us (GLM) prompted further examination of the types of the recognized and synonymized species listed under Bipersona by Re- maudiere and Remaudiere (1997) and Eastop and Blackman (2005). After that examination, it was certain that several specimens housed in the National Muse- um of Natural History Aphidoidea Collection collected from Arizona and New Mexico are not referable to Biper- sona torticauda (Gillette). A thorough study of Bipersona was therefore un- dertaken for clarification. MATERIALS AND METHODS Synoptic descriptions are taken from original descriptions and specimens bor- rowed from various museums and pri- vate collections across North America including: the Essig Museum of Ento- mology (EMEC), University of Califor- nia, Berkeley, CA; Colorado State Uni- versity Collection (CSUC), Fort Collins, CO; the Florida State Collection of Arthropods (FSCA), Gainesville, FL; the Illinois Natural History Survey (INHS), Champaign, IL; National Museum of Natural History Aphidoidea Collection (USNM), Beltsville, MD; University of Minnesota Insect Collection (UMSP), St. Paul, MN; Utah State University Ento- mological Museum (EMUS), Logan, UT; Canadian National Collection of Insects (CNCI), Ottawa, Canada; and individual collections of F. W. Quednau 794 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (FWQC), Sainte-Foy, Canada, and the collection of the second author (ASJC). Codens (except individual collections) are as in Bejsak-Collorado-Mansfield (2004). Morphological terms and struc- tures adapted from Stoetzel et al. (1999) are used in this work. Measurements are presented in millimeters as minimum and maximum ranges of representative specimens. Drawings were begun using a camera lucida, and completed either by hand or by computer-aided illustra- tion programs (Adobe Illustrator® or Adobe Photoshop®). Most specimens had been cleared and mounted in Canada balsam, while others were mounted in gum-based mountants. The information under Specimens Ex- amined is organized to conserve space. Abbreviations for apterae, alatae, ovi- parae, fundatrix, and immatures are listed ass ap. alee ‘ove. tund?, “and “mA. “re- spectively. If a collection was made at the same locality, but on a different date as a previously listed collection, duplicat- ed information is not repeated. Months are designated as roman numerals. For example, the documentation provided for a particular locality may be recorded as: NEBRASKA: War Bonnet Creek, V-26- 2003, on Cirsium nebraskense [= Cirsium cCanescens|, 1). "©; Beard coll” (al) USNM; IV-10-1988, VII-29-1989, I-21- (991) on Cirsium sp. 15 ap: on’ 15 7sk) USNM. In this hypothetical example, the second collection was also found at War Bonnet Creek, even though ““War Bonnet Creek” was not repeated. Collection data that are the same except for collection date are simply listed sequentially. In examples where microscope slides record county, they are listed first within each state then chronologically. Other collec- tion localities are listed chronologically. When specimens are mounted on a single slide (sl.), it is not written as such but is assumed. Bracketed ([ ]) text represents supplemental information by the current authors for clarification purposes. RESULTS During the time this paper was in review, Eastop and Blackman (2005) placed Bipersona hottesi Knowlton and Smith (1936) as a junior synonym of Eomacrosiphon nigromaculosom (Mac- Dougall). Their decision was based on “no significant differences between this species [E. nigromaculosom] and the de- scription of Bipersona hottesi found on wild Rosa in Utah, other than the twisted cauda of the latter’? and they believed the condition of the cauda “was an artefact of the mounting process” (Eastop and Blackman 2005). We concur with Eastop and Blackman’s (2005) synonymy. After we examined in detail the type slide of B. hottesi (USNM), another specific should be mentioned. While Knowlton and Smith’s (1936) description of B. hottesi is based on two apterous viviparous specimens on a single slide, the slide in fact contains three species as follows: three alate viviparae of Macrosiphum euphorbiae (Thomas), one apterous vivi- para of Wahlgreniella nervata (Gillette), and two apterous viviparae of E. nigro- maculosum. The latter two specimens were those described as B. hottesi. The caudas of these two specimens are badly deformed so that they appear constricted in several places, or twisted, as in Bipersona. Bipersona Hottes 1926 Type species: Aphis torticauda Gillette 1907: 389-391. By original designa- tion. Diagnosis.—Large aphids (body length (bl) 2-3+ mm), normally dark red to brownish grey in life, with nymphs pink, orange, or red. Adults sometimes with paler or orange head and thorax and darker reddish to brown behind. Antennae shorter than body, dark brown to black, with numerous sensoria on antennal segment (a.s.) II] and variable numbers on a.s. IV in apterous and alate VOLUME 108, NUMBER 4 viviparae, with a few sensoria sometimes present on a.s. V of alate viviparae. Siphunculi dark brown to black with apical portion with fine polygonal re- ticulation, the remainder roughly imbri- cate. Cauda elongate, tapering with 4-8 lateral and subapical setae, and with about 3 asymmetrical constrictions in vivipara, making it appear twisted or convoluted in dorsal aspect (Figs. IF, 3E). Cauda of ovipara and male shorter and less contorted. Anal plate with a conical posteriorly directed projection (Figs. 1C, 3D). Genital plate with nu- merous setae in addition to the normal two primary and row of posterior setae (Fig. 1D). Notes.—This genus was described by Hottes in 1926 for Aphis torticauda Gillette. Hottes only mentioned Aphis torticauda, despite the fact that other species now referred to this genus had been described: Aphis ochrocentri Cock- erell and Aphis cnici Williams. It is possible that Hottes was unaware of these other names. Hottes pointed out that the genus can be recognized easily by the peculiar structure of the cauda and anal plate, but we note that Artemi- saphis Knowlton and Roberts has a sim- ilar anal plate. Artemisaphis normally has setae on the siphunculi, another unusual character. We have noticed setae on the siphunculi of a few Bipersona specimens, suggesting another link be- tween Artemisaphis and Bipersona. Arte- misaphis has at times been placed as a subgenus of Obtusicauda Soliman, most recently restored to genus rank by Robinson and Halbert (1989) because of its unique cauda and anal plate and the fact that it is tended by ants, while species of Obtusicauda are rarely ant attended. In short, Bipersona is likely a close relative of Artemisaphis, and possibly a slightly more distant relative of Obtusicauda. With the exception of some _ host records from Hypericum (see Discus- G95 sion), Bipersona aphids feed only on various thistles and like other species of thistle-feeding aphids [e.g., Uroleucon longirostre (Gillette and Palmer) and Brachcaudus cardui (L.)|, Bipersona has a long, thin ultimate rostral segment. Bipersona is unusual among the Macro- siphini for its long cornicles and ant- tended association. Most species of aphids with ant attendance have short cornicles and caudas. Bipersona ochrocentri (Cockerell 1903) (Fig. 1) Aphis ochrocentri Cockerell 1903: 248; Bipersona ochrocentri: Palmer 1952: 247; Eastop and Hille Ris Lambers 1976: 106; Remaudiere and Remaudiere 1997: Tile Type material.—Since Cockerell’s (1903) type material is evidently lost, we have designated a neotype for clarification. From material collected approximately 25 miles from Cockerell’s (1903) type locality, we have selected an aptera (far left position) as the neotype and have placed a label on the front of the slide with a map of the position of the specimen: left label states “Bipersona torticauda Gill., on thistle, Las Vegas, No Mex, July <25;-1963, Hortons é Nielsen, 63-17506, GN-105-63”; right label states “NEOTYPE Bipersona ochrocentri (Cockerell) det. G. L. Miller & A. S. Jensen 2005” (USNM). The right label also includes a map for clarifica- tion. See also Specimens Studied. Field features.—Cockerell (1903) not- ed the apterae with “‘head, thorax and most of under parts pale pink; abdomen brownish plumbeous, delicately prui- nose, more or less shining on disc; two segments anterior to cauda pale green,” cauda black, “femora with basal half yellowish-white, apical half black; tibiae and tarsi, antennae and (slender) nectar- ies black.’”? Alatae were of similar color- ation but darker (Cockerell 1903). Ap- 796 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0.1mm Fig. 1. Bipersona ochrocentri. A, Right side, aptera dorsum of head and antennal segment I; left side, aptera venter of head and antennal segment I. B, Utimate rostral segment. C, Anal plate with posterior projection. D, Genital plate of aptera. E, Cornicle of aptera. F, Cauda of aptera. G, Right side, alata dorsum of head and antennal segment I; left side, alata venter of head and antennal segment I. H, Alata antennal segments I-VI. VOLUME 108, NUMBER 4 terous and alate specimens collected by one of us (ASJ) were dark red. Biology.—Cockerell (1903) supplied no notes on biology of this species other than its association with a host plant (Cirsium ochrocentrum A. Gray). The only other information that applies to B. ochrocentri is from our own observations in western Arizona where these aphids were collected on Cirsium sp. in a very arid habitat and were ant tended. Bipersona ochrocentri are able to move quickly (not unlike many Macrosipho- niella species) as compared with other aphids. Recognition characters.—Aptera: Body length 2.50—3.48; width through eyes, 0.61— 0.67 Head (Fig. 1A) sclerotized, smooth dorsally and with some spicules ventrally; dorsal head setae stout, not much longer than diameter of their corresponding basal sockets, tips blunt. Antenna shorter than body, setae less than half width of segment; segment HII 0.49-0.67 long with 9-30 secondary sensoria; IV 0.30-0.43 long without sensoria; V 0.25—0.38 without secondary sensoria; base of VI 0.11—0.15 long; terminal process, 0.46—0.55 long. Rostrum extending past metacoxae; ulti- mate segment (Fig. 1B) 0.30—0.41 long, nearly 4.5 times as long as wide at base, with 7-13 accessory setae. Pronotum and occasionally mesonotum with marginal tubercles. Hind tibia 1.26—1.74 long; hind tarsus IT 0.13—0.19 long. Abdomen dorsally with faint fine reticulation and small in- tersegmental sclerites that appear as clusters of cells, dorsal setae stout, without basal sclerite, segment VIII with dorsal tergite that extends from lateral margins with 4-6 setae; ventrally spiculose; anal plate (Fig. 1C) with posterior projection and numerous setae, genital plate (Fig. 1D) with numerous setae; abdominal marginal tubercles usually present on segment II-V. Cornicle (Fig. 1E) 0.51—0.68 long, sca- brous, tapering slightly with 10-12 rows of reticulations and small flange apically. Cauda (Fig. 1F) 0.14—0.20 long, elongate, TH contorted with 2-3 pairs of lateral setae and a subapical seta. Alata (Fig. 1G): Similar to aptera except: body length 2.27—3.25; width through eyes 0.56—0.61. Antennal (Fig. 1H) segment ITI 0.50—0.61 long, with 26— 41 secondary sensoria; IV 0.30—0.40 long, with I-10 secondary sensoria; V 0.24-0.35 long, without secondary sensoria; base of VI 0.10—0.14 long; terminal process 0.48—0.65 long. Ulti- mate rostral segment 0.33—0.40 long, with 7—10 accessory setae. Hind tibia 1.30—1.51 long; hind tarsus II 0.15—0.18 long. Cornicle 0.35—0.56 long. Cauda 0.10—0.18 long. Notes.—This species was first de- scribed by Cockerell in 1903, apparently based on few specimens. His descriptions lacked almost all details considered necessary today. Extensive search for the types of this species was unsuccessful, forcing us to designate a neotype. We were faced with two distinctive species and had to use Cockerell’s vague de- scription to decide which of the two species was in fact B. ochrocentri. The one decisive character provided by Cock- erell was the numbers and shape of the sensoria on the antennae of the alate vivipara of B. ochrocentri. Our specimens from New Mexico and Arizona closely matched this part of the description, while the specimens of B. torticauda have differently shaped sensoria and more of them. In addition, Cockerell described B. ochrocentri from the same region of New Mexico from which some of our specimens were collected. One possibility was that both Cockerell and Gillette had described the same species. This seemed unlikely based on the antennal characters just mentioned, but also because of the following statement made by) Palmer s(@1952)ein reference? tovithe distinction between B. ochrocentri and B. torticauda: ““This species seems very close to Aphis ochrocentri, judging from the original description, but Gillette after 798 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON examining type specimens considered the species as distinct but recorded no de- tailed comparison.” Palmer’s statement about Gillette’s examination of types implies that Gillette was able to see types of B. ochrocentri. Since Palmer (1952) noted that the types of B. ochrocentri were “unavailable” and we were not able to locate them either, Gillette’s opinion is the only other evidence we have that the two descriptions refer to different species. Palmer also refers to a note in Cock- erell (1893), which she listed under collections of B. ochrocentri. This collec- tion was listed by Cockerell (1893) as an unidentified Aphis species from Colora- do. It is likely that this collection was in fact B. torticauda, considering it was collected in Colorado, not New Mexico or Arizona. Specimens examined.—UNITED STATES OF AMERICA: ARIZONA, Yavapai Co., I-40, exit 109, VII-7-2004, on’ Cirsium spt, A. Jensen’ coll. G*ap:;'3 ale) “Primm: Sen's SIyOUSNMe NEW MEXICO, 1.5 mi. N. Elida, Presidio, VIII-15-1952, on Canada thistle [Cirsium arvense]|, J. H. Russell coll. (6 ap., 2 al., on 3 sl.) USNM; Las Vegas, VII-25- 1963, on thistle, Horton & Nielsen coll. (2 ap., | imm., separate label on right side with “NEOTYPE” and map) USNM;; Roswell, VII-17-[no year given], on thistle, [no collector given] (2 al., 1 imm.{[one specimen on _ slide is not conspecific] ) USNM. Bipersona torticauda (Gillette, 1907) (Figs. 2-3) Aphis torticauda Gillette 1907: 389. Aphis cnici Williams 1911: 40 (preoccu- pied); Davis 1911: 12. Aphis williamsi Soliman placement name). Bipersona torticauda: Hottes 1926: 115; Gillette and Palmer 1934: 143; Palmer 1952:99247; -Eastop) sand Mille; aRas 9272497" (sez Lambers 1976: 106; Remaudiére and Remaudieéere 1997: 77. Systematic notes.—Williams (1911) described Aphis cnici from Cnicus sp. in Nebraska. Soliman (1927) noted that the name was preoccupied by Aphis cnici Schrank (1801), and established the name Aphis williamsi for the aphid described by Williams. We have exam- ined the types of Aphis torticauda and Aphis cnici Williams (= Aphis williamsi Soliman), and concluded that they are conspecific in agreement with Hottes (1930). Bipersona torticauda, as defined here, is a variable species living in a wide range of habitats, on different host plants, and exhibiting color variation. Gillette (1907) also noted differences in color but believed it was only a variation. We considered the possibility that more than one species may be involved, but lacking useful characters for morpholog- ical separation, we decided to retain all B. torticauda-like specimens under one name. In addition, with our work, B. torticauda 1s restored as the type species versus that of B. ochrocentri (1.e., Eastop and Hille Ris Lambers 1976, Remaudiére and Remaudiere 1997). Type material.—Gillette (1907) did not designate types within his treatment of Aphis torticauda (= B. torticauda). However, within his species description, he did record collection dates and localities for the various life cycle stages on which he based his description (June 14, July 22, October 26, and October 27, 1907). From the original collection series we were able identify specimens on four microscope slides that probably repre- sent the syntypes on which he based his descriptions. However, several inconsis- tencies exist with collection dates listed on these slide labels versus those pub- lished by Gillette in 1907. They are as follows: (1) three of the slides are labeled as July 23, 1907 and not July 22, 1907, and (2) one slide is labeled as September VOLUME 108, NUMBER 4 27, 1907 and not October 27, 1907 as published in Gillette (1907). These dis- parities may represent a printer’s typo- graphic or transcription error. This theory may be borne out in the corre- spondence between Gillette and Per- gande dated November 29, 1907 (Na- tional Aphid Collection card file record 124013) where Pergande recorded receipt of a single slide from Gillette that contained specimens he collected on July 23 and October 26, 1907. Since the Gillette (1907) description of the apter- ous male is from October 26, we believe this slide represents this material. Ex- plaining the discrepancy of the slide datedi= September 127; 01907. is™ more hypothetical. Gillette’s description of apterous oviparae from October 27, 1907 stated that “‘several specimens were taken in copula.” Examination of the September 27, 1907 specimens revealed both apterous oviparae and apterous males. Again, these disparities may represent a _ printer's typographic or transcription error. From the above material and for nomenclatural stability, we designate a lectotype specimen. Ex- cept for one specimen which is not congeneric (and noted of the slide label), all other specimens are paralectoypes. From the 18 syntypes we have selected an aptera (at the 9 o’clock position) as the lectotype and have placed a label on the back of the slide with a map of the position of the specimen. Information for the three microscope slides that contain the lectotype and paralectotypes are as follows: (slide 1.) front left label (with map and arrows) states “‘(Aphis) Bipersona torticauda, n. sp. Type, Apt. Vives paratype, (?);-C. P» Gillette’; top label “U.S. Nat. Mus. Cat No 42817”; right label “on Carduus 7-23-07 Ft. @ollins,;Colo:, Coll. C. P: Gillette Colo. Agr. Exp. Sta. Ac. 7-23-07”; back left label ‘‘Bipersona_ torticauda (Gillette) conf. det. G. L. Miller & A. S. Jensen 2005”; back right label “LECTOTYPE 799 & PARALECTOTYPES. (with map and arrows) det. G. L. Miller & A. S. Jensen 2005” (USNM); (slide 2.) front left label “124014 PARALECTOTYPES det. G. L. Miller & A. S. Jensen 2005; front right label “Aphis 6 3 22 torticauda n.sp. Ft. C. Colo., from Carduus sp., Collected by C.P.G., Co-Types’’; back left label ‘‘Bi- persona torticauda (Gillette) conf. det. G. L. Miller & A. S. Jensen 2005, CO: Fort Collins, on Carduus sp., 23 July & 26 Oct. 1907, C. P. Gillette coll; back night label “Collection information from aphid card file. Evidently, Gillette made the slide from multiple collections. GLM 24 May 2005” (USNM, as per National Aphid Collection card file record 124013 dated 7 October 1907): and (slide 3): front left label ““B. torticauda (Gill.), Fort Collins, 9-27-07, Colo., ¢ 2Type, CP:G."; bottom center label) “Meta- type’; front right label ““PARALECTO- TYPES (with map and arrows) det. G. L. Miller & A. S. Jensen 2005; back right label “‘Bipersona_ torticauda_ (Gillette) conf. det. G. L. Miller & A. S. Jensen 2005(CSUC): Three slides containing 18 specimens of B. torticauda were sent to the USNM in 1930 (USNM Cat. No. 42817) that were variously labeled with type, para- type, metatype, and neotype. While we consider two of the slides as part of the original type series as detailed above, the third slide from Poudre Canon, Colora- do on V-30-1929 should not be consid- ered part of the type series. We have also studied 18 syntypes on three slides labeled “COTYPE” of Aphis cnici Wil- liams (USNM) from NEBRASKA, War Bonnet Cr., VI-23-1890, on Cnicus sp., T. A. Williams coll., (8 al., 8 imm. on 3 sl.) USNM. Field features (Fig. 2).—Aptera: color variable with ‘“‘entire dorsum, including head, unicolorous-red; eyes, distal ends of femora, tibiae, and all of tarsi, cornicles and antennae black or black- ish” (Gillette 1907) or “‘brick-red to red- 800 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2: tending ants on Cirsium flodmanii. Photograph courtesy of T. Rand. Colony of Bipersona torticauda and ochre” with appendages and cornicles black and anal plate and cauda “‘reddish to dusky distally” (Palmer 1952). How- ever, one of us (ASJ) collected specimens which ranged from black with orange thorax to dark red with an orange head. Alata: coloration similar to apterae (Gillette 1907) or brick red to ochre red with brownish thoracic lobes and dusky to black appendages (Palmer 1952). Ovipara: similar to apterae except anal plates as conspicuously black (Gillette 1907). Apterous male: “‘sordid yellowish- brown or greenish-rufous with pro- nounced tinge of rufous upon head and thorax” and “antennae, eyes, distal halves of femora and tibiae, tarsi, corni- cles, beak and anal plates black or blackish” (Gillette 1907). Biology.—Gillette (1907) provided good notes on the biology of this species: “Nol eggs swere- "seen (until aticr the appearance of the wingless fall brood of males during the latter part of September and October. This louse is always well attended by ants, and I have been unable to find eggs upon the food-plants in the field, but when the lice are brought into the laboratory and kept for a few days upon thistle, eggs are laid in considerable numbers. They are light green in colour at first, but become polished black in a few days. I cannot help wondering if the ants carry away the eggs.” The second author has collected this species several times, and found it tended by ants in every case (Fig. 2). Similarly to B. ochrocentri, these aphids are fast-mov- ing, tending to withdraw their stylets and run down the plant stem at the slightest disturbance. In spring, the alate viviparae that have settled on bolting thistle plants become hugely gravid with nymphs, explaining Williams’ (1911) statement that, ‘““This is the largest aphis that I have ever seen.” Gillette’s (1907) remarks about alate males in July were in error since Palmer (1952) noted the specimens were actually shriveled viviparae. We have confirmed her findings. Recognition characters.—Aptera: Body length 2.45—3.73; width through eyes, 0.56— 0.72. Head (Fig. 3A) sclerotized, smooth; dorsal head setae long, approximately 5—7 times longer than diameter of their corre- sponding basal sockets; tips of dorsal head setae pointed. Antenna shorter than body, setae more than half width of segment; segment III 0.53—-0.90 long with 21-42 secondary sensoria; IV 0.34—-0.70 long with 0-9 secondary sensoria; V 0.32—0.45 long with 0-2 (usually absent) sensoria; base of VI 0.11—-0.17 long; terminal process, 0.44- 0.65 long. Rostrum extending past meta- coxae; ultimate segment (Fig. 3B) 0.26— 0.38 long, nearly 4 times as long as wide at base, with 11—19 accessory setae. Pronotum and occasionally mesonotum and metano- tum with marginal tubercles. Hind tibia 1.40-2.20 long; hind tarsus II 0.16—0.20 VOLUME 108, NUMBER 4 long. Abdomen dorsally with faint fine reticulation and small intersegmental scler- ites that appear as clusters of cells, dorsal setae often with faint associated basal sclerite, segment VIII with dorsal tergite that extends from lateral margins with 6-8 setae; ventrally spiculose; anal plate (Fig. 3D) with posterior projection and numerous setae; genital plate with numer- ous setae; abdominal marginal tubercles usually present on segment I-V. Cornicle (Fig. 3C) 0.45—0.84 long, scabrous, taper- ing slightly with 10-12 rows of reticulations and small flange apically. Cauda (Fig. 3E) 0.11—0.23 long, elongate, contorted with 3— 4 pairs of lateral setae and a subapical seta. Alata (Fig. 3F): Similar to apterous female except: body length 2.13—3.46; width through eyes 0.52—0.78. Antennal (Fig. 3G) segment III 0.52-0.92 long, with 36—63 secondary sensoria; IV 0.35— 0.75 long, with 10-18 secondary senso- ria; V 0.33—0.54 long, with 0—S secondary sensoria; base of VI 0.12—0.19 long; terminal process 0.50—0.79 long. Ulti- mate rostral segment 0.26—0.40 long, with 11—17 accessory setae. Hind tibia 1.27—2.32 long; hind tarsus II 0.15—0.22 long. Cornicle 0.40—0.74 long. Cauda 0.09—0.20 long. Notes.—Bipersona torticauda can be distinguished from B. ochrocentri pri- marily by its longer setae on the head, antennae and elsewhere, as depicted in Figs. 3A, F, and G. The presence of lateral tubercles on abdominal tergite I in B. torticauda also may be a diagnostic character. As noted in the descriptions, B. torticauda tends to have more sensoria on antennal segments III and IV, but there is substantial overlap in the ranges. There is also a subtle difference in shape of the secondary sensoria between the species, with the sensoria of B. torticauda being less protuberant than in B. ochro- centri. Specimens examined.—CANADA: BRI- TISH COLUMBIA, Westbank, on Cir- sium sp., VI-9-54 [1954], G. L. Ayre coll., 801 (1 al.) CNCI; on Hypericum sp., VII-9- 54[1954], Ayre-Wilson coll., (1 ap.) CNCI; Canal Flats, on thistle, VII-12- 1955, [no collector given], (1 ap.,1 al. on 2 sl.) CNCI; Oliver, Meyers Flats, on thistle, VI-29-1959, L.A.K. [Leonard Kelton] coll., (1 ap.) CNCI; Keremeos, on Cirsium sp., VII-17-1965, W. R. Richards: coll 4G, ap e2eal ones sis) CNCTI; 2 mi. S. Spencer Bridge, [no host given], VII-4-1966, F. Andrews coll., (1 ap.) EMEC; Grand Forks, [X-05-1984, on Hypericum perforatum, [no collector given), (2) apyvon 2 Esl) "-ECNGEZSAS= KATCHEWAN, 5 mi. W. of Furness, VIII-6-40[1940], on scotch thistle [prob- ably Cirsium sp., E. Maw, personal communication], A. R. Brooks coll., (1 ap ul al on 2 si) (ENGL TUNEED STATES OF AMERICA: COLORA- DO, Lake Co., Hwy 24 N. of Buena Vista, VII-4-2004, on Cirsium undulatum, ALS Jensenicollz'@ ap alle inam:on 3 sl.) ASJC; Weld Co., Central Plains Exp. Range, 8 km N. Nann, VIII-17, 25- 1976, on Cirsium megacephalum [Cirsium undulatum), Je “A Scott tcolla 1 ap») USNM;; Ft. Collins, VI-30-98 [1898], on Cnieusisp.,.C. PiGalG RP sGillettelicolle (14 al. on 2 sl., slides have ““Metatype”’ labels) CSUC; X-3-06 [1906], no host listed pEs@ Bales Cy Brage| coll (oval ap. ¢, 2 indeterminable) CSUC; VII-23- 07 [1907], on Carduus sp., C. P. Gillette coll. (Qap-, Ssal-slide labelihas 5 types and “paratype” written, separate label with “U.S. Nat. Mus. Cat. No. 42817, separate label on back with ““LECTO- TYPE and? ~PARALE@TOTYPES7 and map) USNM; on Carduus sp., C. P. Gillette:colls (Ci oy,, 1 ap. 2 ale ap: 3, slide has ““Types” written on label, separate label on front ““PARALECTO- TYPES”) USNM; VII-29 & X-3-1907, on Carduus spa. ©. P > Gillette coll 7Gs ov., 2 ap., 2 al., 3 imm., 1 indeterminable on 2 sl., both slides have ““Types”’ labeled on them) USNM; IX-27-07 [1907], no host data, "Co PG, Cy ee Gillette) colle PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 108, NUMBER 4 (3 ov., 4 &, slide has ‘‘Types” written on label and separate “‘Metatype”’ label, one imm. specimen on slide is not conspecif- ic, separate label on front with ““PARA- LECTOTYPES” and map) CSUC; X-3- 07 (1907)7 on: Carduus sp.,'C.P. G.[C.P. Gillette] coll., (5 ov., 1 ap. 4, slide has “Types” and “Paratype” labels) CSUC; XOLS-O7 1907S ons Carduusssp:;*C. (P: Gillette-coll; (3"ov.,-2 ap. front slide label has “Types” and “paratypes” written, separate label with “U.S. Nat. Mus. Cat. No. 42817”) USNM; VI-8- 1908, on “‘Cardus’’[Carduus sp.?] L.C.B. [L. C. Bragg] coll., (1 ap., 13 al. on 4 sl., one slide from ESUC labeled ‘“‘Meta- type’) CSUC, ESUC, USNM;; VI-12-08 (OOS, noehest dlisted, ol: CBs [ec Brase}-and ©.) Pi Gy [C.P) Gillette] colls., (4 al., slide has “‘Metatype”’ label) CSUC; VII-16-08 [1908], on Carduus sp., no collector listed, (1 ap., 3 al., slide has ““Metatype” label) UMSP; V-13-09 [1909], on thistle, L.C.B. [L. C. Bragg] coll., (6 fund., 2 imm. on 2 sl., slides have “Types” and “Metatype”’ label) CSUC; WiE12-097 [1909], on thistle. C: PL G. [C. P. Gillette] coll., (4 al., 2 ap., slide has “Metatype” label) CSUC; V-22-09 [1909] on thistle. L.@.B:. [LY C. Bragg] coll., (5 fund. 2 imm.on 3 sl., one slide has “‘Metatype” label) CSUC; V-20-10 MOrO} on thistle, WC: Br (LC: Brage] coll., (1 al., 5 ap., slide has ““Metatype”’ label) CSUC; V-26-11, on thistle, L.C.B. [L. C. Bragg] coll., (3 ap., 2 al., 1 imm.on LOMSsl:) (WSNM: = VIFFI-11 [1911]; on tistics h2C@B.. (i. Cr Brage] coll. (4 ap., 20 al.) CNCI, UMSP; VIII-10-1912, on Carduus sp., L. C. Bragg coll., (4 ap.) EMEC; VIII-22-12 [1912], on Cnicus sp., 803 L.C.B. [L. C. Bragg] coll., (14 ap., 1 al., 4 imm. on 2 sl., one slide has ““Metatype”’ label) CSUC; [X-22-15 [1915], on Cnicus SpsuBragescoll® (Gkapival mo0rov7 imm. on 2 sl., CSUC slide has “Meta- type’ label) CSUC, USNM; VIII-27- 20(1920). Mon “thistle, \CeuPs 3Ge [Coy P! Gillette] *collhuG ap:, <1) imm:) ,e@NCI VATIENG=21) a[t920|) nostothers hostiror collector data, (10 ap., 2 imm., slide has ““Metatype” label) CSUC; VIII-1-22 (1922). on ‘Carduustsp:) ©; P. (G., (CanP: Gillette] coll., (7 ap., slide has “‘Meta- type”’ label) CSUC; VII-29-25 [1925], on Cirsium filipendulum [Cirsium engelman- nii|